Maximum running speed limitations on terrestrial mammals: a theoretical approach

Here we study maximum running speed (MRS) limitations on a previously proposed model of energetic and muscle-tendon unit functions on running mammals. In the present work the MRS and some anatomical or physiological limitations are estimated for mammals with body mass between 1.5 and 300 kg. The MRS variations with body mass are discussed and compared with results of previous experimental and observational studies. The tendon strength seems to be the most relevant limitation, but leg extensor muscle mass and metabolic costs could be relevant also. The physiological maximum muscle speed seems to be less important in the body mass range studied here.     

Elastic energy savings and active energy cost in a simple model of running

The energetic economy of running benefits from tendon and other tissues that store and return elastic energy, thus saving muscles from costly mechanical work. The classic “Spring-mass” computational model successfully explains the forces, displacements and mechanical power of running, as the outcome of dynamical interactions between the body center of mass and a purely elastic spring for the leg. However, the Spring-mass model does not include active muscles and cannot explain the metabolic energy cost of running, whether on level ground or on a slope. Here we add explicit actuation and dissipation to the Spring-mass model, and show how they explain substantial active (and thus costly) work during human running, and much of the associated energetic cost. Dissipation is modeled as modest energy losses (5% of total mechanical energy for running at 3 m s^-1) from hysteresis and foot-ground collisions, that must be restored by active work each step. Even with substantial elastic energy return (59% of positive work, comparable to empirical observations), the active work could account for most of the metabolic cost of human running (about 68%, assuming human-like muscle efficiency). We also introduce a previously unappreciated energetic cost for rapid production of force, that helps explain the relatively smooth ground reaction forces of running, and why muscles might also actively perform negative work. With both work and rapid force costs, the model reproduces the energetics of human running at a range of speeds on level ground and on slopes. Although elastic return is key to energy savings, there are still losses that require restorative muscle work, which can cost substantial energy during running.     

Metabolism during flight in two species of bats, Phyllostomus hastatus and Pteropus gouldii.

The energetic cost of flight in a wind-tunnel was measured at various combinations of speed and flight angle from two species of bats whose body masses differ by almost an order of magnitude. The highest mean metabolic rate per unit body mass measured from P. hastatus (mean body mass, 0.093 kg) was 130.4 Wkg-1, and that for P. gouldii (mean body mass, 0.78 kg) was 69.6 Wkg-1. These highest metabolic rates, recorded from flying bats, are essentially the same as those predicted for flying birds of the same body masses, but are from 2.5 to 3.0 times greater than the highest metabolic rates of which similar-size exercising terrestrial mammals appear capable. The lowest mean rate of energy utilization per unit body mass P. hastatus required to sustain level flight was 94.2 Wkg-1 and that for P. gouldii was 53.4 Wkg-1. These data from flying bats together with comparable data for flying birds all fall along a straight line when plotted on double logarithmic coordinates as a function of body mass. Such data show that even the lowest metabolic requirements of bats and birds during level flight are about twice the highest metabolic capabilities of similar-size terrestrial mammals. Flying bats share with flying birds the ability to move substantially greater distance per unit energy consumed than walking or running mammals. Calculations show that P. hastatus requires only one-sixth the energy to cover a given distance as does the same-size terrestrial mammal, while P. gouldii requires one-fourth the energy of the same-size terrestrial mammal. An empirically derived equation is presented which enables one to make estimates of the metabolic rates of bats and birds during level flight in nature from body mass data alone. Metabolic data obtained in this study are compared with predictions calculated from an avian flight theory.     

Running energetics of the North American river otter: do short legs necessarily reduce efficiency on land?

Semi-aquatic mammals move between two very different media (air and water), and are subject to a greater range of physical forces (gravity, buoyancy, drag) than obligate swimmers or runners. This versatility is associated with morphological compromises that often lead to elevated locomotor energetic costs when compared to fully aquatic or terrestrial species. To understand the basis of these differences in energy expenditure, this study examined the interrelationships between limb morphology, cost of transport and biomechanics of running in a semi-aquatic mammal, the North American river otter. Oxygen consumption, preferred locomotor speeds, and stride characteristics were measured for river otters (body mass=11.1 kg, appendicular/axial length=29%) trained to run on a treadmill. To assess the effects of limb length on performance parameters, kinematic measurements were also made for a terrestrial specialist of comparable stature, the Welsh corgi dog (body mass=12.0 kg, appendicular/axial length=37%). The results were compared to predicted values for long legged terrestrial specialists. As found for other semi-aquatic mammals, the net cost of transport of running river otters (6.63 J kg(-1)min(-1) at 1.43 ms(-1)) was greater than predicted for primarily terrestrial mammals. The otters also showed a marked reduction in gait transition speed and in the range of preferred running speeds in comparison to short dogs and semi-aquatic mammals. As evident from the corgi dogs, short legs did not necessarily compromise running performance. Rather, the ability to incorporate a period of suspension during high speed running was an important compensatory mechanism for short limbs in the dogs. Such an aerial period was not observed in river otters with the result that energetic costs during running were higher and gait transition speeds slower for this versatile mammal compared to locomotor specialists.     

The evolution of body armor in mammals: plantigrade constraints of large body size

Predictions associated with opposing selection generating minimum variance in basal metabolic rate (BMR) in mammals at a constrained body mass (CBM; 358 g) were tested. The CBM is presumed to be associated with energetic constraints linked to predation and variable resources at intermediate sizes on a logarithmic mass scale. Opposing selection is thought to occur in response to energetic constraints associated with predation and unpredictable resources. As body size approaches and exceeds the CBM, mammals face increasing risks of predation and daily energy requirements. Fast running speeds may require high BMRs, but unpredictable and low resources may select for low BMRs, which also reduce foraging time and distances and thus predation risks. If these two selection forces oppose each other persistently, minimum BMR variance may result. However, extreme BMR outliers at and close to the CBM should be indicative of unbalanced selection and predator avoidance alternatives (escapers vs. defenders), and may therefore provide indirect support for opposing selection. It was confirmed that body armor in defenders evolves at and above the CBM, and armored mammals had significantly lower BMRs than their nonarmored counterparts. However, analyses comparing the BMR of escapers--the fastest nonarmored runners (Lagomorpha)--with similar-sized counterparts were inconclusive and were confounded by limb morphology associated with speed optimization. These analyses suggest that the risks and costs of predation and the speed limitations of the plantigrade foot may constrain the evolution of large body sizes in plantigrade mammals.     

A biomechanical constraint on body mass in terrestrial mammalian predators

Observations on extant mammals suggest that large body mass is selectively advantageous for a terrestrial predator on large herbivores. Yet, throughout the Cenozoic, some lineages of terrestrial mammalian predators attained greater maximal body masses than others. In order to explain this evolutionary pattern, the following biomechanical constraint on body mass is hypothesized. The stress, set up in the humerus by the bending moment of the peak ground reaction force at maximal running speed, increased with increasing body mass within a given lineage of terrestrial mammalian predators, resulting in a decreasing safety factor for the bone, until a predator could no longer attain the maximal running speed of its smaller relatives. The selective disadvantage of reduced maximal running speed prevented further increase of body mass within the lineage. This hypothesis is tested by examining the scaling of humeral dimensions and estimating maximal body masses in several lineages of terrestrial mammalian predators. Among lineages with otherwise similar postcranial skeletons, those with the more robust humeri at a given body mass attained the greater maximal body masses. Lineages with the longer deltoid ridges/deltopectoral crests of the humeri and/or the more distally located deltoid scars (suggesting the more distal insertions of the humeral flexors) at a given body mass also attained the greater maximal body masses. These results support the existence of the proposed biomechanical constraint, although paleoecological data suggest that some lineages of terrestrial mammalian predators failed to reach the limits, imposed by this constraint, because of the small size of available prey.     

Human leg model predicts ankle muscle-tendon morphology, state, roles and energetics in walking

A common feature in biological neuromuscular systems is the redundancy in joint actuation. Understanding how these redundancies are resolved in typical joint movements has been a long-standing problem in biomechanics, neuroscience and prosthetics. Many empirical studies have uncovered neural, mechanical and energetic aspects of how humans resolve these degrees of freedom to actuate leg joints for common tasks like walking. However, a unifying theoretical framework that explains the many independent empirical observations and predicts individual muscle and tendon contributions to joint actuation is yet to be established. Here we develop a computational framework to address how the ankle joint actuation problem is resolved by the neuromuscular system in walking. Our framework is founded upon the proposal that a consideration of both neural control and leg muscle-tendon morphology is critical to obtain predictive, mechanistic insight into individual muscle and tendon contributions to joint actuation. We examine kinetic, kinematic and electromyographic data from healthy walking subjects to find that human leg muscle-tendon morphology and neural activations enable a metabolically optimal realization of biological ankle mechanics in walking. This optimal realization (a) corresponds to independent empirical observations of operation and performance of the soleus and gastrocnemius muscles, (b) gives rise to an efficient load-sharing amongst ankle muscle-tendon units and (c) causes soleus and gastrocnemius muscle fibers to take on distinct mechanical roles of force generation and power production at the end of stance phase in walking. The framework outlined here suggests that the dynamical interplay between leg structure and neural control may be key to the high walking economy of humans, and has implications as a means to obtain insight into empirically inaccessible features of individual muscle and tendons in biomechanical tasks.     

Muscle mechanical advantage of human walking and running: implications for energy cost.

Muscular forces generated during locomotion depend on an animal's speed, gait, and size and underlie the energy demand to power locomotion. Changes in limb posture affect muscle forces by altering the mechanical advantage of the ground reaction force (R) and therefore the effective mechanical advantage (EMA = r/R, where r is the muscle mechanical advantage) for muscle force production. We used inverse dynamics based on force plate and kinematic recordings of humans as they walked and ran at steady speeds to examine how changes in muscle EMA affect muscle force-generating requirements at these gaits. We found a 68% decrease in knee extensor EMA when humans changed gait from a walk to a run compared with an 18% increase in hip extensor EMA and a 23% increase in ankle extensor EMA. Whereas the knee joint was extended (154-176 degrees) during much of the support phase of walking, its flexed position (134-164 degrees) during running resulted in a 5.2-fold increase in quadriceps impulse (time-integrated force during stance) needed to support body weight on the ground. This increase was associated with a 4.9-fold increase in the ground reaction force moment about the knee. In contrast, extensor impulse decreased 37% (P < 0.05) at the hip and did not change at the ankle when subjects switched from a walk to a run. We conclude that the decrease in limb mechanical advantage (mean limb extensor EMA) and increase in knee extensor impulse during running likely contribute to the higher metabolic cost of transport in running than in walking. The low mechanical advantage in running humans may also explain previous observations of a greater metabolic cost of transport for running humans compared with trotting and galloping quadrupeds of similar size.     

Radiant heat affects thermoregulation and energy expenditure during rewarming from torpor

The high expenditure of energy required for endogenous rewarming is one of the widely perceived disadvantages of torpor. However, recent evidence demonstrates that passive rewarming either by the increase of ambient temperature or by basking in the sun appears to be common in heterothermic birds and mammals. As it is presently unknown how radiant heat affects energy expenditure during rewarming from torpor and little is known about how it affects normothermic thermoregulation, we quantified the effects of radiant heat on body temperature and metabolic rate of the small (body mass 25 g) marsupial Sminthopsis macroura in the laboratory. Normothermic resting individuals exposed to radiant heat were able to maintain metabolic rates near basal levels (at 0.91 ml O(2) g(-1) h(-1)) and a constant body temperature down to an ambient temperature of 12 degrees C. In contrast, metabolic rates of individuals without access to radiant heat were 4.5-times higher at an ambient temperature of 12 degrees C and body temperature fell with ambient temperature. During radiant heat-assisted passive rewarming from torpor, animals did not employ shivering but appeared to maximise uptake of radiant heat. Their metabolic rate increased only 3.2-times with a 15- degrees C rise of body temperature (Q(10)=2.2), as predicted by Q(10) effects. In contrast, during active rewarming shivering was intensive and metabolic rates showed an 11.6-times increase. Although body temperature showed a similar absolute change between the beginning and the end of the rewarming process, the overall energetic cost during active rewarming was 6.3-times greater than that during passive, radiant heat-assisted rewarming. Our study demonstrates that energetic models assuming active rewarming from torpor at low ambient temperatures can substantially over-estimate energetic costs. The low energy expenditure during passive arousal provides an alternative explanation as to why daily torpor is common in sunny regions and suggests that the prevalence of torpor in low latitudes may have been under-estimated in the past.     

Mechanical output of the cat soleus during treadmill locomotion: in vivo vs in situ characteristics

To study the mechanical output of skeletal muscle, four adult cats were trained to run on a treadmill and then implanted under sterile conditions and anesthesia with a force transducer on the soleus tendon and EMG electrodes in the muscle belly. After a two-week recovery period, five consecutive step cycles were filmed at treadmill speeds of 0.8, 1.3 and 2.2 m s-1. Locomotion data in vivo included individual muscle force, length and velocity changes and EMG during each step cycle. Data for an average step cycle at each speed were compared to the force-velocity properties obtained on the same muscle under maximal nerve stimulation and isotonic loading conditions in situ. Results indicate that the force and power generated at a given velocity of shortening during late stance in vivo were greater at the higher speeds of locomotion than the force and power generated at the same shortening velocity in situ. Strain energy stored in the muscle-tendon unit during the yield phase in early stance is felt to be a major contributor to the muscle's enhanced mechanical output during muscle shortening in late stance.     

Running on a slope: A collision-based analysis to assess the optimal slope

When running, energy is lost during stance to redirect the center of mass of the body (COM) from downwards to upwards. The present study uses a collision-based approach to analyze how these energy losses change with slope and speed. Therefore, we evaluate separately the average collision angle, i.e. the angle of deviation from perpendicular relationship between the force and velocity vectors, during the absorptive and generative part of stance. Our results show that on the level, the collision angle of the absorptive phase is smaller than the collision angle of the generative phase, suggesting that the collision is generative to overcome energy losses by soft tissues. When running uphill, the collision becomes more and more generative as slope increases because the average upward vertical velocity of the COM becomes greater than on the level. When running downhill at a constant speed, the collision angle decreases during the generative phase and increases during the absorptive phase because the average downward vertical velocity of the COM becomes greater. As a result, the difference between the collision angles of the generative and absorptive phases observed on the level disappears on a shallow negative slope of ∼−6°, where the collision becomes 'pseudo-elastic' and collisional energy losses are minimized. At this 'optimal' slope, the metabolic energy consumption is minimal. On steeper negative slopes, the collision angle during the absorptive phase becomes greater than during the generative phase and the collision is absorptive. At all slopes, the collision becomes more generative when speed increases.     

Energetic costs of size and sexual signalling in a wolf spider

A prerequisite for honest handicaps is that there are significant condition-dependent costs in the expression of sexual traits. In the wolf spider Hygrolycosa rubrofasciata (Ohlert), sexual signalling (drumming) is costly in terms of increased mortality. Here we investigated whether this mortality may be caused by increased energy expenditure. During sexual signalling, metabolic rate was 22 times higher than at rest and four times higher than when males were actively moving. Metabolic rate per unit mass was positively related to absolute body mass during sexual signalling but not during other activities. This positive relationship is novel to any studies of metabolic rates. Indeed, it seems that the largest males can drum only 12 times per minute before reaching the maximum sustainable metabolic rate, whereas the smallest males may drum up to 39 times per minute. However, there is no relationship between body mass and drumming rate, indicating that larger males are able to compensate for the higher cost of drumming. There was a quadratic relationship between relative abdomen mass and overall body mass, which may provide a partial explanation for the increased energy expenditure of largest males while drumming. Altogether, our results indicate that sexual signalling is highly energetically demanding, which may be the main reason for the honesty of signalling in this species. In addition, the energetic costs are surprisingly strongly size dependent, which may compensate any disadvantage of small male size.     

Effects of velocity and weight support on ground reaction forces and metabolic power during running

The biomechanical and metabolic demands of human running are distinctly affected by velocity and body weight. As runners increase velocity, ground reaction forces (GRF) increase, which may increase the risk of an overuse injury, and more metabolic power is required to produce greater rates of muscular force generation. Running with weight support attenuates GRFs, but demands less metabolic power than normal weight running. We used a recently developed device (G-trainer) that uses positive air pressure around the lower body to support body weight during treadmill running. Our scientific goal was to quantify the separate and combined effects of running velocity and weight support on GRFs and metabolic power. After obtaining this basic data set, we identified velocity and weight support combinations that resulted in different peak GRFs, yet demanded the same metabolic power. Ideal combinations of velocity and weight could potentially reduce biomechanical risks by attenuating peak GRFs while maintaining aerobic and neuromuscular benefits. Indeed, we found many combinations that decreased peak vertical GRFs yet demanded the same metabolic power as running slower at normal weight. This approach of manipulating velocity and weight during running may prove effective as a training and/or rehabilitation strategy.     

Stretching Your Energetic Budget: How Tendon Compliance Affects the Metabolic Cost of Running

Muscles attach to bones via tendons that stretch and recoil, affecting muscle force generation and metabolic energy consumption. In this study, we investigated the effect of tendon compliance on the metabolic cost of running using a full-body musculoskeletal model with a detailed model of muscle energetics. We performed muscle-driven simulations of running at 2–5 m/s with tendon force–strain curves that produced between 1 and 10% strain when the muscles were developing maximum isometric force. We computed the average metabolic power consumed by each muscle when running at each speed and with each tendon compliance. Average whole-body metabolic power consumption increased as running speed increased, regardless of tendon compliance, and was lowest at each speed when tendon strain reached 2–3% as muscles were developing maximum isometric force. When running at 2 m/s, the soleus muscle consumed less metabolic power at high tendon compliance because the strain of the tendon allowed the muscle fibers to operate nearly isometrically during stance. In contrast, the medial and lateral gastrocnemii consumed less metabolic power at low tendon compliance because less compliant tendons allowed the muscle fibers to operate closer to their optimal lengths during stance. The software and simulations used in this study are freely available at simtk.org and enable examination of muscle energetics with unprecedented detail.     

Contributions of muscles to mediolateral ground reaction force over a range of walking speeds

Impaired control of mediolateral body motion during walking is an important health concern. Developing treatments to improve mediolateral control is challenging, partly because the mechanisms by which muscles modulate mediolateral ground reaction force (and thereby modulate mediolateral acceleration of the body mass center) during unimpaired walking are poorly understood. To investigate this, we examined mediolateral ground reaction forces in eight unimpaired subjects walking at four speeds and determined the contributions of muscles, gravity, and velocity-related forces to the mediolateral ground reaction force by analyzing muscle-driven simulations of these subjects. During early stance (0-6% gait cycle), peak ground reaction force on the leading foot was directed laterally and increased significantly (p<0.05) with walking speed. During early single support (14-30% gait cycle), peak ground reaction force on the stance foot was directed medially and increased significantly (p<0.01) with speed. Muscles accounted for more than 92% of the mediolateral ground reaction force over all walking speeds, whereas gravity and velocity-related forces made relatively small contributions. Muscles coordinate mediolateral acceleration via an interplay between the medial ground reaction force contributed by the abductors and the lateral ground reaction forces contributed by the knee extensors, plantarflexors, and adductors. Our findings show how muscles that contribute to forward progression and body-weight support also modulate mediolateral acceleration of the body mass center while weight is transferred from one leg to another during double support.     

Hamstring muscle kinematics and activation during overground sprinting

Hamstring muscle strain injury is one of the most commonly seen injuries in sports such as track and field, soccer, football, and rugby. The purpose of this study was to advance our understanding of the mechanisms of hamstring muscle strain injuries during over ground sprinting by investigating hamstring muscle-tendon kinematics and muscle activation. Three-dimensional videographic and electromyographic (EMG) data were collected for 20 male runners, soccer or lacrosse players performing overground sprinting at their maximum effort. Hamstring muscle-tendon lengths, elongation velocities, and linear envelop EMG data were analyzed for a running gait cycle of the dominant leg. Hamstring muscles exhibited eccentric contractions during the late stance phase as well as during the late swing phase of overground sprinting. The peak eccentric contraction speeds of the hamstring muscles were significantly greater during the late swing phase than during the late stance phase (p=0.001) while the hamstring muscle-tendon lengths at the peak eccentric contraction speeds were significantly greater during the late stance phase than during the late swing phase (p=0.001). No significant differences existed in the maximum hamstring muscle-tendon lengths between the two eccentric contractions. The potential for hamstring muscle strain injury exists during the late stance phase as well as during the late swing phases of overground sprinting.     

Task-Level Strategies for Human Sagittal-Plane Running Maneuvers Are Consistent with Robotic Control Policies

The strategies that humans use to control unsteady locomotion are not well understood. A “spring-mass” template comprised of a point mass bouncing on a sprung leg can approximate both center of mass movements and ground reaction forces during running in humans and other animals. Legged robots that operate as bouncing, “spring-mass” systems can maintain stable motion using relatively simple, distributed feedback rules. We tested whether the changes to sagittal-plane movements during five running tasks involving active changes to running height, speed, and orientation were consistent with the rules used by bouncing robots to maintain stability. Changes to running height were associated with changes to leg force but not stance duration. To change speed, humans primarily used a “pogo stick” strategy, where speed changes were associated with adjustments to fore-aft foot placement, and not a “unicycle” strategy involving systematic changes to stance leg hip moment. However, hip moments were related to changes to body orientation and angular speed. Hip moments could be described with first order proportional-derivative relationship to trunk pitch. Overall, the task-level strategies used for body control in humans were consistent with the strategies employed by bouncing robots. Identification of these behavioral strategies could lead to a better understanding of the sensorimotor mechanisms that allow for effective unsteady locomotion.     

Metabolic compensation during high energy output in fasting, lactating grey seals (Halichoerus grypus): metabolic ceilings revisited

Lactation is the most energetically expensive period for female mammals and is associated with some of the highest sustained metabolic rates (SusMR) in vertebrates (reported as total energy throughput). Females typically deal with this energy demand by increasing food intake and the structure of the alimentary tract may act as the central constraint to ceilings on SusMR at about seven times resting or standard metabolic rate (SMR). However, demands of lactation may also be met by using a form of metabolic compensation such as reducing locomotor activities or entering torpor. In some phocid seals, cetaceans and bears, females fast throughout lactation and thus cannot offset the high energetic costs of lactation through increased food intake. We demonstrate that fasting grey seal females sustain, for several weeks, one of the highest total daily energy expenditures (DEE; 7.4 x SMR) reported in mammals, while progressively reducing maintenance metabolic expenditures during lactation through means not explained by reduction in lean body mass or behavioural changes. Simultaneously, the energy-exported in milk is progressively increased, associated with increased lipoprotein lipase activity in the mammary gland, resulting in greater offspring growth. Our results suggest that females use compensatory mechanisms to help meet the extraordinary energetic costs of lactation. Additionally, although the concepts of SusMR and ceilings on total DEE may be somewhat different in fasting lactating species, our data on phocid seals demonstrate that metabolic ceilings on milk energy output, in general, are not constrained by the same kind of peripheral limitations as are other energy-consuming tissues. In phocid seals, the high ceilings on DEE during lactation, coupled with metabolic compensation, are undoubtedly important factors enabling shortened lactation.     

Consequences of forward translation of the point of force application for the mechanics of running

In running humans, the point of force application between the foot and the ground moves forwards during the stance phase. Our aim was to determine the mechanical consequences of this 'point of force translation' (POFT). We modified the planar spring-mass model of locomotion to incorporate POFT, and then compared spring-mass simulations with and without POFT. We found that, if leg stiffness is adjusted appropriately, it is possible to maintain very similar values of peak vertical ground reaction force (GRF), stance time, contact length and vertical centre of mass displacement, whether or not POFT occurs. The leg stiffness required to achieve this increased as the distance of POFT increased. Peak horizontal GRF and mechanical work per step were lower when POFT occurred. The results indicate that the lack of POFT in the traditional spring-mass model should not prevent it from providing good predictions of peak vertical GRF, stance time, contact length and vertical centre of mass displacement in running humans, if an appropriate spring stiffness is used. However, the model can be expected to overestimate peak horizontal GRF and mechanical work per step. When POFT occurs, the spring stiffness in the traditional spring-mass model is not equivalent to leg stiffness. Therefore, caution should be exercised when using spring stiffness to understand how the musculoskeletal system adapts to different running conditions. This can explain the contradictory results in the literature regarding the effect of running speed on leg stiffness.     

Optimal body size with respect to maximal speed for the yellow-spotted monitor lizard (Varanus panoptes; Varanidae)

Studies of locomotor performance often link variation in morphology with ecology. While maximum sprint speed is a commonly used performance variable, the absolute limits for this performance trait are not completely understood. Absolute maximal speed has often been shown to increase linearly with body size, but several comparative studies covering a large range of body sizes suggest that maximal speed does not increase indefinitely with body mass but rather reaches an optimum after which speed declines. Because of the comparative nature of these studies, it is difficult to determine whether this decrease is due to biomechanical constraints on maximal speed or is a consequence of phylogenetic inertia or perhaps relaxed selection for lower maximal speed at large body size. To explore this issue, we have examined intraspecific variations in morphology, maximal sprint speed, and kinematics for the yellow-spotted monitor lizard Varanus panoptes, which varied in body mass from 0.09 to 5.75 kg. We show a curvilinear relationship between body size and absolute maximal sprint speed with an optimal body mass with respect to speed of 1.245 kg. This excludes the phylogenetic inertia hypothesis, because this effect should be absent intraspecifically, while supporting the biomechanical constraints hypothesis. The relaxed selection hypothesis cannot be excluded if there is a size-based behavioral shift intraspecifically, but the biomechanical constraints hypothesis is better supported from kinematic analyses. Kinematic measurements of hind limb movement suggest that the distance moved by the body during the stance phase may limit maximum speed. This limit is thought to be imposed by a decreased ability of the bones and muscles to support body mass for larger lizards.