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An unusual pattern among the scaling laws in nature is that the fastest animals are neither the largest, nor the smallest, but rather intermediately sized. Because of the enormous diversity in animal shape, the mechanisms underlying this have long been difficult to determine. To address this, we challenge predictive human musculoskeletal simulations, scaled in mass from the size of a mouse (0.1 kg) to the size of an elephant (2000 kg), to move as fast as possible. Our models replicate patterns observed across extant animals including: (i) an intermediate optimal body mass for speed; (ii) a reduction in the cost of transport with increasing size; and (iii) crouched postures at smaller body masses and upright postures at larger body masses. Finally, we use our models to determine the mechanical limitations of speed with size, showing larger animals may be limited by their ability to produce muscular force while smaller animals are likely limited by their ability to produce larger ground reaction forces. Despite their bipedal gait, our models replicate patterns observed across quadrupedal animals, suggesting these biological phenomena likely represent general rules and are not the result of phylogenetic or other ecological factors that typically hinder comparative studies., (© 2024. The Author(s).)

The shape of skeletal muscle varies remarkably-with important implications for locomotor performance. In many muscles, the fibres are arranged at an angle relative to the tendons' line of action, termed the pennation angle. These pennate muscles allow more sarcomeres to be packed side by side, enabling the muscle to generate higher maximum forces for a given muscle size. Historically, the physiological cross-sectional area (PCSA) has been used to capture both the size and arrangement of muscle fibres, and is one of the best predictors of a muscles capacity to produce force. However, the anatomical and mechanical implications of PCSA remain ambiguous as misinterpretations have limited our ability to understand the mechanical advantage of pennate muscle designs. We developed geometric models to resolve the mechanistic and functional impacts of pennation angle across a range of muscle shapes and sizes. Comparisons among model predictions and empirical data on human lower limb muscles demonstrated how a pennate arrangement of fibres allows muscles to produce up to six times more isometric force when compared with non-pennate muscles of the same volume. We show that in muscles much longer than thick, an optimal pennation angle exists at which isometric force is maximized. Using empirically informed geometric models we demonstrate the functional significance of a pennate muscle design and provide a new parameter, pennation mechanical advantage, which quantifies this performance improvement., Competing Interests: We declare we have no competing interests., (© 2024 The Authors.)

Animal performance fundamentally influences behaviour, ecology, and evolution. It typically varies monotonously with size. A notable exception is maximum running speed; the fastest animals are of intermediate size. Here we show that this peculiar allometry results from the competition between two musculoskeletal constraints: the kinetic energy capacity, which dominates in small animals, and the work capacity, which reigns supreme in large animals. The ratio of both capacities defines the physiological similarity index Γ, a dimensionless number akin to the Reynolds number in fluid mechanics. The scaling of Γ indicates a transition from a dominance of muscle forces to a dominance of inertial forces as animals grow in size; its magnitude defines conditions of "dynamic similarity" that enable comparison and estimates of locomotor performance across extant and extinct animals; and the physical parameters that define it highlight opportunities for adaptations in musculoskeletal "design" that depart from the eternal null hypothesis of geometric similarity. The physiological similarity index challenges the Froude number as prevailing dynamic similarity condition, reveals that the differential growth of muscle and weight forces central to classic scaling theory is of secondary importance for the majority of terrestrial animals, and suggests avenues for comparative analyses of locomotor systems., (© 2024. The Author(s).)

Skeletal muscles bulge when they contract. These three-dimensional shape changes, coupled with fiber rotation, influence a muscle's mechanical performance by uncoupling fiber velocity from muscle belly velocity (i.e., gearing). Muscle shape change and gearing are likely mediated by the interaction between internal muscle properties and contractile forces. Muscles with greater stiffness and intermuscular fat, due to aging or disuse, may limit a muscle's ability to bulge in width, subsequently causing higher gearing. The aim of this study was to determine the influence of internal muscle properties on shape change, fiber rotation, and gearing in the medial (MG) and lateral gastrocnemii (LG) during isometric plantar flexion contractions. Multimodal imaging techniques were used to measure muscle shear modulus, intramuscular fat, and fat-corrected physiological cross-sectional area (PCSA) at rest, as well as synchronous muscle architecture changes during submaximal and maximal contractions in the MG and LG of 20 young (24 ± 3 yr) and 13 older (70 ± 4 yr) participants. Fat-corrected PCSA was positively associated with fiber rotation, gearing, and changes in thickness during submaximal contractions, but it was negatively associated with changes in thickness at maximal contractions. Muscle stiffness and intramuscular fat were related to muscle bulging and reduced fiber rotation, respectively, but only at high forces. Furthermore, the MG and LG had varied internal muscle properties, which may relate to the differing shape changes, fiber rotations, and gearing behaviors observed at each contraction level. These results indicate that internal muscle properties may play an important role in mediating in vivo muscle shape change and gearing, especially during high-force contractions. NEW & NOTEWORTHY Here, we measured internal muscle properties in vivo to determine their influence on the varying shape change and gearing behaviors in the synergistic gastrocnemii muscles. These relationships have previously only been hypothesized or examined within isolated muscles during supramaximal contractions. Our results contribute to a more comprehensive understanding of the factors that influence a muscle's mechanical response to force with implications for preventing or treating muscle deficits associated with aging, disease, and disuse.

This paper considers the effects of claw morphology on the gripping efficiency of arboreal (Varanus varius) and burrowing (Varanus gouldii and Varanus panoptes) lizards. To ensure a purely morphological comparison between the lizards, we circumvent the material effects of claws from different species, by modelling and testing claw replicates of the same material properties. We correlate climbing efficiency to critical morphological features including; claw height (hc), width (wc), length (lc), curvature () and tip angle (γ), which are expressed as ratios to normalise mechanically beneficial claw structures. We find that there is strong correlation between the static grip force Fsg and the claw aspect and the cross-sectional rigidity ratio , and milder correlation (i.e. higher scatter) with the profile rigidity ratio . These correlations are also true for the interlocking grip force Fint over different shaped and sized protuberances, though we note that certain protuberance size-shape couplings are of detriment to the repeatability of Fint. Of the three lizard species, the claws of the arboreal (V. varius) are found to be superior to those of the burrower lizards (V. gouldii and V. panoptes) as a result of the V. varius claws having a smaller aspect, a higher cross-sectional rigidity ratio and a small profile rigidity ratio, which are deemed noteworthy morphological parameters that influence a claw's ability to grip effectively., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2023. Published by The Company of Biologists Ltd.)

Among terrestrial mammals, the largest, the 3 tonne African elephant, is one-million times heavier than the smallest, the 3 g pygmy shrew. Body mass is the most obvious and arguably the most fundamental characteristic of an animal, impacting many important attributes of its life history and biology. Although evolution may guide animals to different sizes, shapes, energetic profiles or ecological niches, it is the laws of physics that limit biological processes and, in turn, affect how animals interact with their environment. Consideration of scaling helps us to understand why elephants are not merely scaled-up shrews, but rather have modified body proportions, posture and locomotor style to mitigate the consequences of their large size. Scaling offers a quantitative lens into how biological features vary compared with predictions based on physical laws. In this Review, we provide an introduction to scaling and its historical context, focusing on two fields that are strongly represented in experimental biology: physiology and biomechanics. We show how scaling has been used to explore metabolic energy use with changes in body size. We discuss the musculoskeletal and biomechanical adaptations that animals use to mitigate the consequences of size, and provide insights into the scaling of mechanical and energetic demands of animal locomotion. For each field, we discuss empirical measurements, fundamental scaling theories and the importance of considering phylogenetic relationships when performing scaling analyses. Finally, we provide forward-looking perspectives focused on improving our understanding of the diversity of form and function in relation to size., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2023. Published by The Company of Biologists Ltd.)

Many climbing animals use direction-dependent adhesives to attach to vertical or inclined surfaces. These structures adhere when activated via a pull but detach when pushed. Therefore, a challenge arises when a change in climbing direction causes external forces such as gravity to change its acting orientation upon the lizard. To investigate how specialized climbers adjust, we studied kinematics and dynamics of six Hemidactylus frenatus geckos climbing head-up and head-down a vertical racetrack. We found that limbs functionally swap their adhesive role: feet above the centre of mass (COM) generated adhesive forces, feet below the COM compressive forces, both equal in magnitude across directions. To investigate how lizards perform this swap, despite the constraint of their direction-dependent adhesives, we analysed kinematic adjustments across multiple smaller levels of hierarchy: limbs, feet and toes. All levels contributed: the hindfoot angle was reoriented realigning the adhesive structure, the hindlimb centre range of motion was further protracted and the hindfoot toe spreading was reduced. Notably, all three variables were adjustments of hindlimbs, suggesting that they make a more flexible contribution in upward versus downward climbing, while forelimbs may be anatomically or functionally constrained. The relevance of multilevel dynamic adjustments might inform the development of performant gaits for legged climbing robots.

Semelparity is a breeding strategy whereby an individual invests large amounts of resources into a single breeding season, leading to the death of the individual. Male northern quolls ( Dasyurus hallucatus ) are the largest known mammal to experience a post-breeding die-off; however, the cause of their death is unknown, dissimilar from causes in other semelparous dasyurids. To identify potential differences between male northern quolls that breed once, and females that can breed for up to four seasons, the behaviours, activity budgets, speeds and distances travelled were examined. Northern quolls were captured on Groote Eylandt off the coast of the Northern Territory, Australia, and were fitted with accelerometers. A machine learning algorithm (Self-organizing Map) was trained on more than 76 h of recorded footage of quoll behaviours and used to predict behaviours in 42 days of data from wild roaming quolls (7M : 6F). Male northern quolls were more active (male 1.27 g , s.d. = 0.41; female 1.18 g, s.d. = 0.36), spent more time walking (13.09% male: 8.93% female) and engaged in less lying/resting behaviour than female northern quolls (7.67% male: 23.65% female). Reduced resting behaviour among males could explain the post-breeding death as the deterioration in appearance reflects that reported for sleep-deprived rodents., (© 2023 The Authors.)

Manoeuvrability, the ability to make rapid changes in direction, is central to animal locomotion. Turning performance may depend on the ability to successfully complete key challenges including: withstanding additional lateral forces, maintaining sufficient friction, lateral leaning during a turn and rotating the body to align with the new heading. We filmed high-speed turning in domestic dogs (Canis lupus familiaris) to quantify turning performance and explore how performance varies with body size and shape. Maximal speed decreased with higher angular velocity, greater centripetal acceleration and smaller turning radii, supporting a force limit for wider turns and a friction limit for sharp turns. Variation in turning ability with size was complex: medium sized dogs produced greater centripetal forces, had relatively higher friction coefficients, and generally aligned the body better with the heading compared with smaller and larger bodied dogs. Body shape also had a complex pattern, with longer forelimbs but shorter hindlimbs being associated with better turning ability. Further, although more crouched forelimbs were associated with an increased ability to realign the body in the direction of movement, more upright hindlimbs were related to greater centripetal and tangential accelerations. Thus, we demonstrate that these biomechanical challenges to turning can vary not only with changes in speed or turning radius, but also with changes in morphology. These results will have significant implications for understanding the link between form and function in locomotory studies, but also in predicting the outcome of predator-prey encounters., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2022. Published by The Company of Biologists Ltd.)

A considerable biomechanical challenge faces larger terrestrial animals as the demands of body support scale with body mass (Mb), while muscle force capacity is proportional to muscle cross-sectional area, which scales with Mb2/3. How muscles adjust to this challenge might be best understood by examining varanids, which vary by five orders of magnitude in size without substantial changes in posture or body proportions. Muscle mass, fascicle length and physiological cross-sectional area all scale with positive allometry, but it remains unclear, however, how muscles become larger in this clade. Do larger varanids have more muscle fibres, or does individual fibre cross-sectional area (fCSA) increase? It is also unknown if larger animals compensate by increasing the proportion of fast-twitch (higher glycogen concentration) fibres, which can produce higher force per unit area than slow-twitch fibres. We investigated muscle fibre area and glycogen concentration in hindlimb muscles from varanids ranging from 105 g to 40,000 g. We found that fCSA increased with modest positive scaling against body mass (Mb0.197) among all our samples, and ∝Mb0.278 among a subset of our data consisting of never-frozen samples only. The proportion of low-glycogen fibres decreased significantly in some muscles but not others. We compared our results with the scaling of fCSA in different groups. Considering species means, fCSA scaled more steeply in invertebrates (∝Mb0.575), fish (∝Mb0.347) and other reptiles (∝Mb0.308) compared with varanids (∝Mb0.267), which had a slightly higher scaling exponent than birds (∝Mb0.134) and mammals (∝Mb0.122). This suggests that, while fCSA generally increases with body size, the extent of this scaling is taxon specific, and may relate to broad differences in locomotor function, metabolism and habitat between different clades., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2022. Published by The Company of Biologists Ltd.)

Robotic systems for complex tasks, such as search and rescue or exploration, are limited for wheeled designs, thus the study of legged locomotion for robotic applications has become increasingly important. To successfully navigate in regions with rough terrain, a robot must not only be able to negotiate obstacles, but also climb steep inclines. Following the principles of biomimetics, we developed a modular bio-inspired climbing robot, named X4, which mimics the lizard's bauplan including an actuated spine, shoulders, and feet which interlock with the surface via claws. We included the ability to modify gait and hardware parameters and simultaneously collect data with the robot's sensors on climbed distance, slip occurrence and efficiency. We first explored the speed-stability trade-off and its interaction with limb swing phase dynamics, finding a sigmoidal pattern of limb movement resulted in the greatest distance travelled. By modifying foot orientation, we found two optima for both speed and stability, suggesting multiple stable configurations. We varied spine and limb range of motion, again showing two possible optimum configurations, and finally varied the centre of pro- and retraction on climbing performance, showing an advantage for protracted limbs during the stride. We then stacked optimal regions of performance and show that combining optimal dynamic patterns with either foot angles or ROM configurations have the greatest performance, but further optima stacking resulted in a decrease in performance, suggesting complex interactions between kinematic parameters. The search of optimal parameter configurations might not only be beneficial to improve robotic in-field operations but may also further the study of the locomotive evolution of climbing of animals, like lizards or insects., (© 2021 IOP Publishing Ltd.)

When muscles contract and change length, they also bulge in thickness and/or width. These shape changes extend the functional range of skeletal muscle by allowing individual muscle fibres to shorten at different velocities than the whole muscle. Age-related differences in muscle architecture and tissue properties influence how older muscles change shape and architecture during contractions, yet this remains unexplored in active older adults. The aim of this study was to quantify and compare in vivo muscle architecture and shape changes in the medial (MG) and lateral (LG) gastrocnemii of active younger and older adults during isometric plantarflexion contractions. Fifteen younger (21 ± 2y) and 15 older (70 ± 3y) participants performed contractions at 20%, 40%, 60%, 80%, and 100% of maximum voluntary contraction (MVC). B-mode ultrasound was used to measure fascicle length, pennation angle and muscle thickness in MG and LG. We found no influence of age on changes in normalized fascicle length and thickness, or absolute change in pennation angle during contractions. With increasing contraction level, MG and LG fascicle shortening (P < 0.001) and rotation (P < 0.001) increased. However, the change in muscle thickness increased at higher contraction levels in LG, and not MG. Similarly, increased changes in pennation angle were associated with increased muscle thickness in LG, but not MG at 80% and 100% MVC. These results suggest that (1) gastrocnemii shape changes are similar in active older and younger adults at matched levels of effort, and (2) the relationship between pennation angle and muscle thickness can differ between synergistics (LG and MG) and across contraction levels., (Copyright © 2021. Published by Elsevier Ltd.)

Tail movement is an important component of vertebrate locomotion and likely contributes to dynamic stability during steady-state locomotion. Previous results suggest that the tail plays a significant role in lizard locomotion, but little data are available on tail motion during locomotion and how it differs with morphological, ecological, and phylogenetic parameters. We collected high-speed vertical climbing and horizontal locomotion video data from 43 lizard species from four taxonomic groups (Agamidae, Gekkota, Scincidae, and Varanidae) across four habitats. We introduce a new semi-automated and generalizable analysis pipeline for tail and spine motion analysis including markerless pose-estimation, semi-automated kinematic recognition, and muti-species data analysis. We found that step length relative to snout-vent length (SVL) increased with tail length relative to SVL. Examining spine cycles agnostic to limb stride phase, we found that ranges of inter-tail bending compared with inter-spine bending increased with relative tail length, while ranges of tail deflection relative to spine deflection increased with relative speed. Considering stepwise strides, we found the angular velocity and acceleration of the tail center of mass increased with relative speed. These results will provide general insights into the biomechanics of tails in sprawling locomotion enabling biomimetic applications in robotics, and a better understanding of vertebrate form and function. We look forward to adding more species, behaviors, and locomotor speeds to our analysis pipeline through collaboration with other research groups., (© The Author(s) 2021. 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.)

Collecting quantitative information on animal behaviours is difficult, especially from cryptic species or species that alter natural behaviours under observation. Using harness-mounted tri-axial accelerometers free-roaming domestic cats (Felis Catus) we developed a methodology that can precisely classify finer-scale behaviours. We further tested the effect of a prey-protector device designed to reduce prey capture. We aligned accelerometer traces collected at 50 Hz with video files (60 fps) and labelled 12 individual behaviours, then trained a supervised machine-learning algorithm using Kohonen super self-organising maps (SOM). The SOM was able to predict individual behaviours with a ~ 99.6% overall accuracy, which was slightly better than for random forest estimates using the same dataset (98.9%). There was a significant effect of sample size, with precision and sensitivity decreasing rapidly below 2000 1-s observations. We were also able to detect a behaviour specific reduction in the predictability when cats were fitted with the prey-protector device indicating it altered biomechanical gait. Our results can be applied in movement ecology, zoology and conservation, where habitat specific movement performance between predators or prey may be critical to managing species of conservation significance, or in veterinary and agricultural fields, where early detection of movement pathologies can improve animal welfare.

Locomotion is a key aspect associated with ecologically relevant tasks for many organisms, therefore, survival often depends on their ability to perform well at these tasks. Despite this significance, we have little idea how different performance tasks are weighted when increased performance in one task comes at the cost of decreased performance in another. Additionally, the ability for natural systems to become optimized to perform a specific task can be limited by structural, historic or functional constraints. Climbing lizards provide a good example of these constraints as climbing ability likely requires the optimization of tasks which may conflict with one another such as increasing speed, avoiding falls and reducing the cost of transport (COT). Understanding how modifications to the lizard bauplan can influence these tasks may allow us to understand the relative weighting of different performance objectives among species. Here, we reconstruct multiple performance landscapes of climbing locomotion using a 10 d.f. robot based upon the lizard bauplan, including an actuated spine, shoulders and feet, the latter which interlock with the surface via claws. This design allows us to independently vary speed, foot angles and range of motion (ROM), while simultaneously collecting data on climbed distance, stability and efficiency. We first demonstrate a trade-off between speed and stability, with high speeds resulting in decreased stability and low speeds an increased COT. By varying foot orientation of fore- and hindfeet independently, we found geckos converge on a narrow optimum of foot angles (fore 20°, hind 100°) for both speed and stability, but avoid a secondary wider optimum (fore -20°, hind -50°) highlighting a possible constraint. Modifying the spine and limb ROM revealed a gradient in performance. Evolutionary modifications in movement among extant species over time appear to follow this gradient towards areas which promote speed and efficiency.

In our everyday lives, we negotiate complex and unpredictable environments. Yet, much of our knowledge regarding locomotion has come from studies conducted under steady-state conditions. We have previously shown that humans rely on the ankle joint to absorb energy and recover from perturbations; however, the muscle-tendon unit (MTU) behaviour and motor control strategies that accompany these joint-level responses are not yet understood. In this study, we determined how neuromuscular control and plantar flexor MTU dynamics are modulated to maintain stability during unexpected vertical perturbations. Participants performed steady-state hopping and, at an unknown time, we elicited an unexpected perturbation via rapid removal of a platform. In addition to kinematics and kinetics, we measured gastrocnemius and soleus muscle activations using electromyography and in vivo fascicle dynamics using B-mode ultrasound. Here, we show that an unexpected drop in ground height introduces an automatic phase shift in the timing of plantar flexor muscle activity relative to MTU length changes. This altered timing initiates a cascade of responses including increased MTU and fascicle length changes and increased muscle forces which, when taken together, enables the plantar flexors to effectively dissipate energy. Our results also show another mechanism, whereby increased co-activation of the plantar- and dorsiflexors enables shortening of the plantar flexor fascicles prior to ground contact. This co-activation improves the capacity of the plantar flexors to rapidly absorb energy upon ground contact, and may also aid in the avoidance of potentially damaging muscle strains. Our study provides novel insight into how humans alter their neural control to modulate in vivo muscle-tendon interaction dynamics in response to unexpected perturbations. These data provide essential insight to help guide design of lower-limb assistive devices that can perform within varied and unpredictable environments.

Geometric scaling predicts a major challenge for legged, terrestrial locomotion. Locomotor support requirements scale identically with body mass ( α M 1 ), while force-generation capacity should scale α M 2/3 as it depends on muscle cross-sectional area. Mammals compensate with more upright limb postures at larger sizes, but it remains unknown how sprawling tetrapods deal with this challenge. Varanid lizards are an ideal group to address this question because they cover an enormous body size range while maintaining a similar bent-limb posture and body proportions. This study reports the scaling of ground reaction forces and duty factor for varanid lizards ranging from 7 g to 37 kg. Impulses (force×time) ( α M 0.99-1.34 ) and peak forces ( α M 0.73-1.00 ) scaled higher than expected. Duty factor scaled α M 0.04 and was higher for the hindlimb than the forelimb. The proportion of vertical impulse to total impulse increased with body size, and impulses decreased while peak forces increased with speed.

There is a functional trade-off in the design of skeletal muscle. Muscle strength depends on the number of muscle fibers in parallel, while shortening velocity and operational distance depend on fascicle length, leading to a trade-off between the maximum force a muscle can produce and its ability to change length and contract rapidly. This trade-off becomes even more pronounced as animals increase in size because muscle strength scales with area (length 2 ) while body mass scales with volume (length 3 ). In order to understand this muscle trade-off and how animals deal with the biomechanical consequences of size, we investigated muscle properties in the pectoral girdle of varanid lizards. Varanids are an ideal group to study the scaling of muscle properties because they retain similar body proportions and posture across five orders of magnitude in body mass and are highly active, terrestrially adapted reptiles. We measured muscle mass, physiological cross-sectional area, fascicle length, proximal and distal tendon lengths, and proximal and distal moment arms for 27 pectoral girdle muscles in 13 individuals across 8 species ranging from 64 g to 40 kg. Standard and phylogenetically informed reduced major axis regression was used to investigate how muscle architecture properties scale with body size. Allometric growth was widespread for muscle mass (scaling exponent >1), physiological cross-sectional area (scaling exponent >0.66), but not tendon length (scaling exponent >0.33). Positive allometry for muscle mass was universal among muscles responsible for translating the trunk forward and flexing the elbow, and nearly universal among humeral protractors and wrist flexors. Positive allometry for PCSA was also common among trunk translators and humeral protractors, though less so than muscle mass. Positive scaling for fascicle length was not widespread, but common among humeral protractors. A higher proportion of pectoral girdle muscles scaled with positive allometry than our previous work showed for the pelvic girdle, suggesting that the center of mass may move cranially with body size in varanids, or that the pectoral girdle may assume a more dominant role in locomotion in larger species. Scaling exponents for physiological cross-sectional area among muscles primarily associated with propulsion or with a dual role were generally higher than those associated primarily with support against gravity, suggesting that locomotor demands have at least an equal influence on muscle architecture as body support. Overall, these results suggest that larger varanids compensate for the increased biomechanical demands of locomotion and body support at higher body sizes by developing larger pectoral muscles with higher physiological cross-sectional areas. The isometric scaling rates for fascicle length among locomotion-oriented pectoral girdle muscles suggest that larger varanids may be forced to use shorter stride lengths, but this problem may be circumvented by increases in limb excursion afforded by the sliding coracosternal joint., (© 2020 Anatomical Society.)

Urbanization significantly impacts the health and viability of wildlife populations yet it is not well understood how urban landscapes differ from non-urban landscapes with regard to their effects on wildlife. This study investigated the physiological response of eastern grey kangaroos (Macropus giganteus) to land use at a landscape scale. Using fecal glucocorticoid metabolites (FGM) we compared stress levels of kangaroo populations in urban and non-urban environments. We modeled FGM concentrations from 24 kangaroo populations against land use (urban or non-urban) and other anthropogenic and environmental factors, using a linear modeling approach. We found that land use was a significant predictor of FGM concentrations in eastern grey kangaroos with significant differences in concentrations between urban and non-urban populations. However, the direction of the relationship differed between northern and southern regions of Australia. In the northern study sites, kangaroos in urban areas had significantly higher FGM levels than their non-urban counterparts. In contrast, in southern sites, where kangaroos occur in high densities in many urban areas, urban kangaroos had lower FGM concentrations than non-urban kangaroos. Rainfall and temperature were also significant predictors of FGM and the direction of the relationship was consistent across both regions. These results are consistent with the contrasting abundance and persistence of kangaroo populations within the urban matrix between the two study regions. In the northern region many populations have declined over the last two decades and are fragmented, also occurring at lower densities than in southern sites. Our study indicates that it is the characteristics of urban environments, rather than the urban environment per se, which determines the extent of impacts of urbanization on kangaroos. This research provides insights into how the design of urban landscapes can influence large mammal populations., (© 2019 by the Ecological Society of America.)

From the camel's toes to the horse's hooves, the diversity in foot morphology among mammals is striking. One distinguishing feature is the presence of fat pads, which may play a role in reducing foot pressures, or may be related to habitat specialization. The camelid family provides a useful paradigm to explore this as within this phylogenetically constrained group we see prominent (camels) and greatly reduced (alpacas) fat pads. We found similar scaling of vertical ground reaction force with body mass, but camels had larger foot contact areas, which increased with velocity, unlike alpacas, meaning camels had relatively lower foot pressures. Further, variation between specific regions under the foot was greater in alpacas than camels. Together, these results provide strong evidence for the role of fat pads in reducing relative peak locomotor foot pressures, suggesting that the fat pad role in habitat specialization remains difficult to disentangle.

The morphology and locomotor performance of a species can determine their inherent fitness within a habitat type. Koalas have an unusual morphology for marsupials, with several key adaptations suggested to increase stability in arboreal environments. We quantified the kinematics of their movement over ground and along narrow arboreal trackways to determine the extent to which their locomotion resembled that of primates, occupying similar niches, or basal marsupials from which they evolved. On the ground, the locomotion of koalas resembled a combination of marsupial behaviours and primate-like mechanics. For example, their fastest strides were bounding type gaits with a top speed of 2.78 m s -1 (mean 1.20 m s -1 ), resembling marsupials, while the relatively longer stride length was reflective of primate locomotion. Speed was increased using equal modification of stride length and frequency. On narrow substrates, koalas took longer but slower strides (mean 0.42 m s -1 ), adopting diagonally coupled gaits including both lateral and diagonal sequence gaits, the latter being a strategy distinctive among arboreal primates. The use of diagonally coupled gaits in the arboreal environment is likely only possible because of the unique gripping hand morphology of both the fore and hind feet of koalas. These results suggest that during ground locomotion, they use marsupial-like strategies but alternate to primate-like strategies when moving amongst branches, maximising stability in these environments. The locomotion strategies of koalas provide key insights into an independent evolutionary branch for an arboreal specialist, highlighting how locomotor strategies can convergently evolve between distant lineages., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

Characterisation of an organism's performance in different habitats provides insight into the conditions that allow it to survive and reproduce. In recent years, the northern quoll ( Dasyurus hallucatus ) - a medium-sized semi-arboreal marsupial native to northern Australia - has undergone significant population declines within open forest, woodland and riparian habitats, but less so in rocky areas. To help understand this decline, we quantified the biomechanical performance of wild northern quolls as they ran up inclined narrow (13 mm pole) and inclined wide (90 mm platform) substrates. We predicted that quolls may possess biomechanical adaptations to increase stability on narrow surfaces, which are more common in rocky habitats. Our results showed that quolls have some biomechanical characteristics consistent with a stability advantage on narrow surfaces. This includes the coupled use of limb pairs, as indicated via a decrease in footfall time, and an ability to produce corrective torques to counteract the toppling moments commonly encountered during gait on narrow surfaces. However, speed was constrained on narrow surfaces, and quolls did not adopt diagonal sequence gaits, unlike true arboreal specialists such as primates. In comparison with key predators, such as cats and dogs, northern quolls appear inferior in terrestrial environments but have a stability advantage at higher speeds on narrow supports. This may partially explain the heterogeneous declines in northern quoll populations among various habitats on mainland Australia., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

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.

Neuromotor deficits are an important sign of manganese (Mn) toxicity in humans and laboratory animals. However, the impacts of Mn exposure on the motor function of wild animals remains largely unknown. Here, we assessed the impact of chronic exposure to Mn from active mining operations on Groote Eylandt, Australia on the motor function of the semi-arboreal northern quoll (Dasyurus hallucatus), an endangered species. The three motor tests conducted-maximum sprint speed on a straight run, manoeuvrability around a corner, and motor control on a balance beam-showed that elevated Mn body burden did not diminish performance of these traits. However, quolls with higher Mn body burden approached a corner at a significantly narrower range of speeds, due to a significantly lower maximum approach speed. Slower speeds approaching a turn may reduce success at catching prey and avoiding predators. Given that maximum sprint speed on a straight run was not affected by Mn body burden, but maximum speed entering a corner was, slower speeds approaching a turn may reflect compensation for otherwise impaired performance in the turn., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Certain lizards are known to run bipedally. Modelling studies suggest bipedalism in lizards may be a consequence of a caudal shift in the body centre of mass, combined with quick bursts of acceleration, causing a torque moment at the hip lifting the front of the body. However, some lizards appear to run bipedally sooner and for longer than expected from these models, suggesting positive selection for bipedal locomotion. While differences in morphology may contribute to bipedal locomotion, changes in kinematic variables may also contribute to extended bipedal sequences, such as changes to the body orientation, tail lifting and changes to the ground reaction force profile. We examined these mechanisms among eight Australian agamid lizards. Our analysis revealed that angular acceleration of the trunk about the hip, and of the tail about the hip were both important predictors of extended bipedal running, along with increased temporal asymmetry of the ground reaction force profile. These results highlight important dynamic movements during locomotion, which may not only stabilize bipedal strides, but also to de-stabilize quadrupedal strides in agamid lizards, in order to temporarily switch to, and extend a bipedal sequence., (© 2018 The Author(s).)

Movement speed can underpin an animal's probability of success in ecological tasks. Prey often use agility to outmanoeuvre predators; however, faster speeds increase inertia and reduce agility. Agility is also constrained by grip, as the foot must have sufficient friction with the ground to apply the forces required for turning. Consequently, ground surface should affect optimum turning speed. We tested the speed-agility trade-off in buff-footed antechinus ( Antechinus mysticus ) on two different surfaces. Antechinus used slower turning speeds over smaller turning radii on both surfaces, as predicted by the speed-agility trade-off. Slipping was 64% more likely on the low-friction surface, and had a higher probability of occurring the faster the antechinus were running before the turn. However, antechinus compensated for differences in surface friction by using slower pre-turn speeds as their amount of experience on the low-friction surface increased, which consequently reduced their probability of slipping. Conversely, on the high-friction surface, antechinus used faster pre-turn speeds in later trials, which had no effect on their probability of slipping. Overall, antechinus used larger turning radii (0.733±0.062 versus 0.576±0.051 m) and slower pre-turn (1.595±0.058 versus 2.174±0.050 m s -1 ) and turning speeds (1.649±0.061 versus 2.01±0.054 m s -1 ) on the low-friction surface. Our results demonstrate the interactive effect of surface friction and the speed-agility trade-off on speed choice. To predict wild animals' movement speeds, future studies should examine the interactions between biomechanical trade-offs and terrain, and quantify the costs of motor mistakes in different ecological activities., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists 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.

While snake venoms have been the subject of intense study, comparatively little work has been done on lizard venoms. In this study, we have examined the structural and functional diversification of anguimorph lizard venoms and associated toxins, and related these results to dentition and predatory ecology. Venom composition was shown to be highly variable across the 20 species of Heloderma , Lanthanotus , and Varanus included in our study. While kallikrein enzymes were ubiquitous, they were also a particularly multifunctional toxin type, with differential activities on enzyme substrates and also ability to degrade alpha or beta chains of fibrinogen that reflects structural variability. Examination of other toxin types also revealed similar variability in their presence and activity levels. The high level of venom chemistry variation in varanid lizards compared to that of helodermatid lizards suggests that venom may be subject to different selection pressures in these two families. These results not only contribute to our understanding of venom evolution but also reveal anguimorph lizard venoms to be rich sources of novel bioactive molecules with potential as drug design and development lead compounds., Competing Interests: The authors declare no conflict of interest.

How extinct, non-avian theropod dinosaurs locomoted is a subject of considerable interest, as is the manner in which it evolved on the line leading to birds. Fossil footprints provide the most direct evidence for answering these questions. In this study, step width-the mediolateral (transverse) distance between successive footfalls-was investigated with respect to speed (stride length) in non-avian theropod trackways of Late Triassic age. Comparable kinematic data were also collected for humans and 11 species of ground-dwelling birds. Permutation tests of the slope on a plot of step width against stride length showed that step width decreased continuously with increasing speed in the extinct theropods ( p < 0.001), as well as the five tallest bird species studied ( p < 0.01). Humans, by contrast, showed an abrupt decrease in step width at the walk-run transition. In the modern bipeds, these patterns reflect the use of either a discontinuous locomotor repertoire, characterized by distinct gaits (humans), or a continuous locomotor repertoire, where walking smoothly transitions into running (birds). The non-avian theropods are consequently inferred to have had a continuous locomotor repertoire, possibly including grounded running. Thus, features that characterize avian terrestrial locomotion had begun to evolve early in theropod history., (© 2017 The Author(s).)

Many hemipteran bugs can jump explosively from plant substrates, which can be very smooth. We therefore analysed the jumping performance of froghoppers ( Philaenus spumarius, Aphrophoridae) and leafhoppers ( Aphrodes bicinctus/makarovi, Cicadellidae) taking off from smooth (glass) and rough (sandpaper, 30 µm asperity size) surfaces. On glass, the propulsive hind legs of Philaenus froghoppers slipped, resulting in uncontrolled jumps with a fast forward spin, a steeper angle and only a quarter of the velocity compared with jumps from rough surfaces. By contrast, Aphrodes leafhoppers took off without their propulsive hind legs slipping, and reached low take-off angles and high velocities on both substrates. This difference in jumping ability from smooth surfaces can be explained not only by the lower acceleration of the long-legged leafhoppers, but also by the presence of 2-9 soft pad-like structures (platellae) on their hind tarsi, which are absent in froghoppers. High-speed videos of jumping showed that platellae contact the surface briefly (approx. 3 ms) during the acceleration phase. Friction force measurements on individual hind tarsi on glass revealed that at low sliding speeds, both pushing and pulling forces were small, and insufficient to explain the recorded jumps. Only when the tarsi were pushed with higher velocities did the contact area of the platellae increase markedly, and high friction forces were produced, consistent with the observed jumps. Our findings show that leafhoppers have special adhesive footpads for jumping from smooth surfaces, which achieve firm grip and rapid control of attachment/detachment by combining anisotropic friction with velocity dependence., (© 2017 The Authors.)

Cancellous bone is very sensitive to its prevailing mechanical environment, and study of its architecture has previously aided interpretations of locomotor biomechanics in extinct animals or archaeological populations. However, quantification of architectural features may be compromised by poor preservation in fossil and archaeological specimens, such as post mortem cracking or fracturing. In this study, the effects of post mortem cracks on the quantification of cancellous bone fabric were investigated through the simulation of cracks in otherwise undamaged modern bone samples. The effect on both scalar (degree of fabric anisotropy, fabric elongation index) and vector (principal fabric directions) variables was assessed through comparing the results of architectural analyses of cracked vs. non-cracked samples. Error was found to decrease as the relative size of the crack decreased, and as the orientation of the crack approached the orientation of the primary fabric direction. However, even in the best-case scenario simulated, error remained substantial, with at least 18% of simulations showing a > 10% error when scalar variables were considered, and at least 6.7% of simulations showing a > 10° error when vector variables were considered. As a 10% (scalar) or 10° (vector) difference is probably too large for reliable interpretation of a fossil or archaeological specimen, these results suggest that cracks should be avoided if possible when analysing cancellous bone architecture in such specimens., (© 2016 Anatomical Society.)

The survival of both the hunter and the hunted often comes down to speed. Yet how fast an animal can run is intricately linked to its size, such that the fastest animals are not the biggest nor the smallest. The ability to maintain high speeds is dependent on the body's capacity to withstand the high stresses involved with locomotion. Yet even when standing still, scaling principles would suggest that the mechanical stress an animal feels will increase in greater demand than its body can support. So if big animals want to be fast, they must find solutions to overcome these high stresses. This article explores the ways in which extant animals mitigate size-related increases in musculoskeletal stress in an effort to help understand where all the giants have gone., Competing Interests: The authors have declared that no competing interests exist.

The short-beaked echidna (Tachyglossus aculeatus) is a monotreme and therefore provides a unique combination of phylogenetic history, morphological differentiation and ecological specialisation for a mammal. The echidna has a unique appendicular skeleton, a highly specialised myrmecophagous lifestyle and a mode of locomotion that is neither typically mammalian nor reptilian, but has aspects of both lineages. We therefore were interested in the interactions of locomotor biomechanics, ecology and movements for wild, free-living short-beaked echidnas. To assess locomotion in its complex natural environment, we attached both GPS and accelerometer loggers to the back of echidnas in both spring and summer. We found that the locomotor biomechanics of echidnas is unique, with lower stride length and stride frequency than reported for similar-sized mammals. Speed modulation is primarily accomplished through changes in stride frequency, with a mean of 1.39 Hz and a maximum of 2.31 Hz. Daily activity period was linked to ambient air temperature, which restricted daytime activity during the hotter summer months. Echidnas had longer activity periods and longer digging bouts in spring compared with summer. In summer, echidnas had higher walking speeds than in spring, perhaps because of the shorter time suitable for activity. Echidnas spent, on average, 12% of their time digging, which indicates their potential to excavate up to 204 m 3 of soil a year. This information highlights the important contribution towards ecosystem health, via bioturbation, of this widespread Australian monotreme., (© 2016. Published by The Company of Biologists Ltd.)

Elephants, the largest living land mammals, have evolved a specialized foot morphology to help reduce locomotor pressures while supporting their large body mass. Peak pressures that could cause tissue damage are mitigated passively by the anatomy of elephants' feet, yet this mechanism does not seem to work well for some captive animals. This study tests how foot pressures vary among African and Asian elephants from habitats where natural substrates predominate but where foot care protocols differ. Variations in pressure patterns might be related to differences in husbandry, including but not limited to trimming and the substrates that elephants typically stand and move on. Both species' samples exhibited the highest concentration of peak pressures on the lateral digits of their feet (which tend to develop more disease in elephants) and lower pressures around the heel. The trajectories of the foot's centre of pressure were also similar, confirming that when walking at similar speeds, both species load their feet laterally at impact and then shift their weight medially throughout the step until toe-off. Overall, we found evidence of variations in foot pressure patterns that might be attributable to husbandry and other causes, deserving further examination using broader, more comparable samples.

Background: The functional design of skeletal muscles is shaped by conflicting selective pressures between support and propulsion, which becomes even more important as animals get larger. If larger animals were geometrically scaled up versions of smaller animals, increases in body size would cause an increase in musculoskeletal stress, a result of the greater scaling of mass in comparison to area. In large animals these stresses would come dangerously close to points of failure. By examining the architecture of 22 hindlimb muscles in 27 individuals from 9 species of varanid lizards ranging from the tiny 7.6 g Varanus brevicauda to the giant 40 kg Varanus komodoensis, we present a comprehensive dataset on the scaling of musculoskeletal architecture in monitor lizards (varanids), providing information about the phylogenetic constraints and adaptations of locomotor muscles in sprawling tetrapods., Results: Scaling results for muscle mass, pennation and physiological cross-sectional area (PCSA), all suggest that larger varanids increase the relative force-generating capacity of femur adductors, knee flexors and ankle plantarflexors, with scaling exponents greater than geometric similarity predicts. Thus varanids mitigate the size-related increases in stress by increasing muscle mass and PCSA rather than adopting a more upright posture with size as is shown in other animals. As well as the scaling effects of muscle properties with body mass, the variation in muscle architecture with changes in hindlimb posture were also prominent. Within varanids, posture varies with habitat preference. Climbing lizards display a sprawling posture while terrestrial lizards display a more upright posture. Sprawling species required larger PCSAs and muscle masses in femur retractors, knee flexors, and ankle plantarflexors in order to support the body., Conclusions: Both size and posture-related muscle changes all suggest an increased role in support over propulsion, leading to a decrease in locomotor performance which has previously been shown with increases in size. These estimates suggest the giant Pleistocene varanid lizard (Varanus megalania priscus) would likely not have been able to outrun early humans with which it co-habitated the Australian landmass with.

Organismal functions are size-dependent whenever body surfaces supply body volumes. Larger organisms can develop strongly folded internal surfaces for enhanced diffusion, but in many cases areas cannot be folded so that their enlargement is constrained by anatomy, presenting a problem for larger animals. Here, we study the allometry of adhesive pad area in 225 climbing animal species, covering more than seven orders of magnitude in weight. Across all taxa, adhesive pad area showed extreme positive allometry and scaled with weight, implying a 200-fold increase of relative pad area from mites to geckos. However, allometric scaling coefficients for pad area systematically decreased with taxonomic level and were close to isometry when evolutionary history was accounted for, indicating that the substantial anatomical changes required to achieve this increase in relative pad area are limited by phylogenetic constraints. Using a comparative phylogenetic approach, we found that the departure from isometry is almost exclusively caused by large differences in size-corrected pad area between arthropods and vertebrates. To mitigate the expected decrease of weight-specific adhesion within closely related taxa where pad area scaled close to isometry, data for several taxa suggest that the pads' adhesive strength increased for larger animals. The combination of adjustments in relative pad area for distantly related taxa and changes in adhesive strength for closely related groups helps explain how climbing with adhesive pads has evolved in animals varying over seven orders of magnitude in body weight. Our results illustrate the size limits of adhesion-based climbing, with profound implications for large-scale bio-inspired adhesives.

The ability for prey to escape a pursuing predator is dependent both on the prey's speed away from the threat and on their ability to rapidly change directions, or maneuverability. Given that the biomechanical trade-off between speed and maneuverability limits the simultaneous maximization of both performance traits, animals should not select their fastest possible speeds when running away from a pursuing predator but rather a speed that maximizes the probability of successful escape. We explored how variation in the relationship between speed and maneuverability-or the shape of the trade-off-affects the optimal choice of speed for escaping predators. We used tablet-based games that simulated interactions between predators and prey (human subjects acting as predators attempting to capture "prey" moving across a screen). By defining a specific relationship between speed and maneuverability, we could test the survival of each of the possible behavioral choices available to this phenotype, i.e., the best combination of speed and maneuverability for prey fitness, based on their ability to escape. We found that the shape of the trade-off function affected the prey's optimal speed for success in escaping, the prey's maximum performance in escaping, and the breadth of speeds over which the prey's performance was high. The optimal speed for escape varied only when the trade-off between speed and maneuverability was non-linear. Phenotypes possessing trade-off functions for which maneuverability was only compromised at high speeds exhibited lower optimal speeds. Phenotypes that exhibited greater increases in maneuverability for any decrease in speed were more likely to have broader ranges of performance, meaning that individuals could attain their maximum performance across a broader range of speeds. We also found that there was a differential response of the subject's learning to these different components of locomotion. With increased experience through repeated trials, subjects were able to successfully catch faster and faster dots. However, no improvement was observed in the subject's ability to capture more maneuverable prey. Our work highlights the costs of high-speed movement on other traits, including maneuverability, which make the use of an animal's fastest speeds unlikely, even when attempting to escape predators. By investigating the shape of the trade-off functions between speed and maneuverability and the way the environment and morphology mediates this trade-off, we can begin to understand why animals choose to move at the speeds they do when they are running away from predators or attempting to capture prey., (© The Author 2015. 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.)

An animal's movement speed affects all behaviors and underlies the intensity of an activity, the time it takes to complete it, and the probability of successfully completing it, but which factors determine how fast or slow an animal chooses to move? Despite the critical importance of an animal's choice of speed (hereafter designated as "speed-choice"), we still lack a framework for understanding and predicting how fast animals should move in nature. In this article, we develop a framework for predicting speed that is applicable to any animal-including humans-performing any behavior where choice of speed occurs. To inspire new research in this area, we (1) detail the main factors likely to affect speed-choice, including organismal constraints (i.e., energetic, physiological, and biomechanical) and environmental constraints (i.e., predation intensity and abiotic factors); (2) discuss the value of optimal foraging theory in developing models of speed-choice; and (3) describe how optimality models might be integrated with the range of potential organismal and environmental constraints to predict speed. We show that by utilizing optimality theory it is possible to provide quantitative predictions of optimal speeds across different ecological contexts. However, the usefulness of any predictive models is still entirely dependent on being able to provide relevant mathematical functions to insert into such models. We still lack basic knowledge about how an animal's speed affects its motor control, maneuverability, observational skills, and vulnerability to predators. Studies exploring these gaps in knowledge will help facilitate the field of optimal performance and allow us to adequately parameterize models predicting the speed-choice of animals, which represents one of the most basic of all behavioral decisions., (© The Author 2015. 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.)

Burrowing is an important form of locomotion in reptiles, but no study has examined the energetic cost of burrowing for reptiles. This is significant because burrowing is the most energetically expensive mode of locomotion undertaken by animals and many burrowing species therefore show specialisations for their subterranean lifestyle. We examined the effect of temperature and substrate characteristics (coarse sand or fine sand) on the net energetic cost of burrowing (NCOB) and burrowing rate in two species of the Egernia group of skinks (Liopholis striata and Liopholis inornata) compared with other burrowing animals. We further tested for morphological specialisations among burrowing species by comparing the relationship between body shape and retreat preference in Egernia group skinks. For L. striata and L. inornata, NCOB is 350 times more expensive than the predicted cost of pedestrian terrestrial locomotion. Temperature had a positive effect on burrowing rate for both species, and a negative effect on NCOB for L. striata but not L. inornata. Both NCOB and burrowing rate were independent of substrate type. Burrows constructed by skinks had a smaller cross-sectional area than those constructed by mammals of comparable mass, and NCOB of skinks was lower than that of mammals of similar mass. After accounting for body size, retreat preference was significantly correlated with body shape in Egernia group skinks. Species of Egernia group skinks that use burrows for retreats have narrower bodies and shorter front limbs than other species. We conclude that the morphological specialisations of burrowing skinks allow them to construct relatively narrow burrows, thereby reducing NCOB and the total cost of constructing their burrow retreats., (© 2015. Published by The Company of Biologists Ltd.)

The origin of bipedal locomotion in lizards is unclear. Modeling studies have suggested that bipedalism may be an exaptation, a byproduct of features originally designed to increase maneuverability, which were only later exploited. Measurement of the body center of mass (BCOM) in 124 species of lizards confirms a significant rearward shift among bipedal lineages. Further racetrack trials showed a significant acceleration threshold between bipedal and quadrupedal runs. These suggest good general support for a passive bipedal model, in which the combination of these features lead to passive lifting of the front of the body. However, variation in morphology could only account for 56% of the variation in acceleration thresholds, suggesting that dynamics have a significant influence on bipedalism. Deviation from the passive bipedal model was compared with node age, supporting an increase in the influence of dynamics over time. Together, these results show that bipedalism may have first arisen as a consequence of acceleration and a rearward shift in the BCOM, but subsequent linages have exploited this consequence to become bipedal more often, suggesting that bipedalism in lizards may convey some advantage. Exploitation of bipedalism was also associated with increased rates of phenotypic diversity, suggesting exploiting bipedalism may promote adaptive radiation., (© 2014 The Author(s). Evolution © 2014 The Society for the Study of Evolution.)

Adaptations promoting greater performance in one habitat are thought to reduce performance in others. However, there are many examples of animals in which, despite habitat differences, such predicted differences in performance do not occur. One such example is the relationship between locomotory performance to habitat for varanid lizards. To explain the lack of difference in locomotor performance we examined detailed observations of the kinematics of each lizard's stride. Differences in kinematics were greatest between climbing and non-climbing species. For terrestrial lizards, the kinematics indicated that increased femur adduction, femur rotation and ankle angle all contributed positively to changes in stride length, but they were constrained for climbing species, probably because of biomechanical restrictions on the centre of mass height (to increase stability on vertical surfaces). Despite climbing species having restricted stride length, no differences have been previously reported in sprint speed between climbing and non-climbing varanids. This is best explained by climbing varanids using an alternative speed modulation strategy of varying stride frequency to avoid the potential trade-off of speed versus stability on vertical surfaces. Thus, by measuring the relevant biomechanics for lizard strides, we have shown how kinematic differences among species can mask performance differences typically associated with habitat variation.

Rowing is demanding, in part, because drag on the oars increases as the square of their speed. Hence, as muscles shorten faster, their force capacity falls, whereas drag rises. How do frogs resolve this dilemma to swim rapidly? We predicted that shortening velocity cannot exceed a terminal velocity where muscle and fluid torques balance. This terminal velocity, which is below Vmax, depends on gear ratio (GR = outlever/inlever) and webbed foot area. Perhaps such properties of swimmers are 'tuned', enabling shortening speeds of approximately 0.3Vmax for maximal power. Predictions were tested using a 'musculo-robotic' Xenopus laevis foot driven either by a living in vitro or computational in silico plantaris longus muscle. Experiments verified predictions. Our principle finding is that GR ranges from 11.5 to 20 near the predicted optimum for rowing (GR ≈ 11). However, gearing influences muscle power more strongly than foot area. No single morphology is optimal for producing muscle power. Rather, the 'optimal' GR decreases with foot size, implying that rowing ability need not compromise jumping (and vice versa). Thus, despite our neglect of additional forces (e.g. added mass), our model predicts pairings of physiological and morphological properties to confer effective rowing. Beyond frogs, the model may apply across a range of size and complexity from aquatic insects to human-powered rowing.

Studies of the muscle force-velocity relationship and its derived n-shaped power-velocity curve offer important insights into muscular limits of performance. Given the power is maximal at 1/3 V(max), geometric scaling of muscle force coupled with fluid drag force implies that this optimal muscle-shortening velocity for power cannot be maintained across the natural body-size range. Instead, muscle velocity may decrease with increasing body size, conferring a similar n-shaped power curve with body size. Here we examine swimming speed and muscle function in the aquatic frog Xenopus laevis. Swimming speed shows an n-shaped scaling relationship, peaking at 47.35 g. Further, in vitro muscle function of the ankle extensor plantaris longus also shows an optimal body mass for muscle power output (47.27 g), reflecting that of swimming speed. These findings suggest that in drag-based aquatic systems, muscle-environment interactions vary with body size, limiting both the muscle's potential to produce power and the swimming speed.