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Cursorial

Cursorial refers to the suite of anatomical, physiological, and biomechanical adaptations in animals that facilitate rapid and efficient running, often over long distances or at high speeds, distinguishing cursorial organisms from those specialized for other forms of locomotion such as climbing or swimming.^[1] These adaptations primarily involve modifications to the skeletal structure, musculature, and posture of the limbs, enabling cursorial animals—such as cheetahs, horses, and ostriches—to achieve superior stride lengths and rates while minimizing energy expenditure.^[2] The term originates from the Latin cursor, meaning "runner," and is most commonly applied in zoology to describe terrestrial vertebrates and invertebrates evolved for cursorial locomotion in open habitats like plains and grasslands.^[3] Key anatomical features of cursorial animals include a digitigrade or unguligrade posture, where the animal runs on its toes rather than the full foot, which elevates the body and lengthens stride by reducing contact time with the ground.^[1] Limb elongation is prominent, particularly in the lower segments: the metapodials (foot bones) are often extended and fused into structures like the cannon bone in equids and bovids, while the lower leg bones (tibia, fibula, radius, ulna) are slender and lightweight to decrease rotational inertia and enhance speed.^[2] Hindlimbs typically bear more propulsive force, with powerful extensors for thrust, whereas forelimbs are adapted for shock absorption and steering; in highly cursorial quadrupeds, all four limbs contribute to locomotion but with specialized roles.^[4] Muscle attachments near joints further boost stride rate by allowing rapid contraction, and reductions in digit number—often to two or one (hoofed ungulates)—streamline the feet for stability on hard substrates.^[1] Examples of cursorial animals span mammals, birds, and even some insects and dinosaurs. Among mammals, felids like the cheetah exemplify extreme cursoriality with speeds up to 100 km/h, achieved through flexible spines and elongated limbs, though sustained running is limited by muscle fatigue rather than overheating.^[5] Equids such as horses and artiodactyls like deer and antelopes represent endurance runners, with specialized tarsal bones like the astragalus and calcaneum articulating to provide pivot points for efficient galloping.^[2] In birds, ratites like ostriches possess cursorial legs with pneumatic bones for lightness and powerful digital flexors. Cursorial traits have evolved convergently across taxa, often in response to predation pressures or foraging needs in open environments, but they compromise other abilities like maneuverability in dense terrain.^[6] Biomechanically, cursorial locomotion optimizes speed through the formula speed = stride length × stride frequency, where adaptations maximize unsupported flight phases in gaits like bounding or galloping.^[1] Trabecular bone density in limbs increases to withstand high-impact stresses, balancing lightness with structural integrity, as seen in comparative studies of mammalian skeletons.^[7] In dinosaurs, theoretical models highlight similar correlates like elongated metatarsals in theropods for predatory pursuits.^[8] Overall, cursoriality represents an evolutionary trade-off favoring terrestrial speed at the expense of versatility, influencing ecological roles from pursuit predators to migratory herbivores.^[9]

Definition and Overview

Definition

Cursorial locomotion refers to a form of terrestrial movement in which organisms are specially adapted for efficient running, emphasizing either high speed, endurance, or a combination of both across land surfaces.^[1] This adaptation enables sustained horizontal progression over open terrain, often characterized by a flowing gait that optimizes stride length and frequency for minimal energy expenditure during prolonged activity.^[10] Cursorial organisms typically exhibit a streamlined body plan with limbs proportioned to reduce mass relative to length, facilitating rapid acceleration and deceleration without compromising stability.^[11] Unlike saltatorial locomotion, which prioritizes jumping and leaping for propulsion over obstacles or short bursts, cursorial movement avoids vertical emphasis and instead supports continuous, ground-level travel. Similarly, it differs from scansorial adaptations geared toward climbing vertical or inclined surfaces, focusing solely on planar efficiency rather than gripping or ascending capabilities.^[1] These distinctions highlight cursoriality's specialization for open habitats where evasion of predators or pursuit of prey demands consistent velocity over distance.^[12] Representative examples illustrate this spectrum: the cheetah (Acinonyx jubatus) embodies speed-oriented cursoriality, achieving bursts exceeding 100 km/h through explosive sprints, while the pronghorn antelope (Antilocapra americana) exemplifies endurance, maintaining speeds of 55-65 km/h over several kilometers.^[1] In contrast, non-cursorial species like the African elephant (Loxodonta africana), with its graviportal structure optimized for weight-bearing rather than agility, exhibit lumbering gaits ill-suited for rapid terrestrial pursuit due to their bulk.^[13]

Etymology and Historical Usage

The term "cursorial" originates from the Latin cursor, meaning "runner" or "messenger," and entered the English scientific lexicon in the early to mid-19th century as an adjective denoting adaptations suited to running.^[14]^[15] This etymological root reflects its application to organisms or anatomical features optimized for terrestrial locomotion, drawing from Late Latin cursōrius ("of running").^[14] Historically, the term gained prominence in zoological descriptions during the 1830s and 1840s, particularly in studies of mammalian anatomy and locomotion. It was employed qualitatively to describe animals exhibiting morphological and behavioral traits for fast or sustained running, such as elongated limbs and reduced body mass relative to graviportal (weight-supporting) forms, with early focus on large terrestrial mammals like equids and bovids.^[11] These usages appeared in anatomical treatises emphasizing functional adaptations for speed over endurance or weight-bearing, marking a shift toward classifying locomotor strategies in comparative biology.^[11] By the 20th century, the concept of cursoriality evolved from these qualitative assessments to incorporate quantitative biomechanical analyses, including limb bone ratios, stride efficiency, and inferred velocities, to differentiate it more precisely from other locomotor modes like graviportal.^[11] Seminal works by anatomists such as R. McN. Alexander in the 1970s and 1980s formalized this distinction, using physical models to evaluate cursorial traits across vertebrates and highlighting a continuum rather than binary categories.^[8] This refinement provided a foundation for modern definitions in biology, bridging historical nomenclature with empirical metrics of performance.^[11]

Cursorial Adaptations

General Morphological Adaptations

Cursorial locomotion relies on a suite of morphological adaptations that prioritize speed, endurance, and efficiency in terrestrial movement across various animal groups. Central to these are elongated limbs, which extend stride length and facilitate higher velocities by increasing the distance covered per step. Distal limb segments, such as the tibia and metatarsals, often show disproportionate elongation relative to proximal elements like the femur, allowing for greater leverage and reduced contact time with the ground.^[16] Additionally, lightweight skeletons minimize inertial costs during acceleration and sustained running; in avian cursorials, pneumatized bones filled with air sacs reduce overall skeletal mass while maintaining structural integrity.^[17] Powerful hindlimb musculature, including enlarged extensors like the gastrocnemius and plantaris, generates the propulsive force needed for rapid starts and maintenance of high speeds, with muscle attachments optimized for powerful knee and ankle extension.^[18] Physiological traits complement these structural features to support prolonged activity. Efficient oxygen delivery to working muscles enables aerobic metabolism during extended runs. Biomechanical principles underlying cursorial form emphasize streamlined motion. The parasagittal limb posture aligns limbs beneath the body's midline, directing force vectorially forward and minimizing energy-wasting lateral excursions.^[19] This configuration reduces rotational inertia around the body's long axis and enhances stability at speed, with reduced lateral movement contributing to overall locomotor economy.^[20] Quantitative assessments of cursoriality often involve limb proportion indices relative to body mass. For instance, hindlimb length scales positively allometrically with body mass (exponent ≈0.37-0.40), but highly cursorial forms exhibit elevated ratios, such as metatarsal-to-femur lengths exceeding 0.8, indicating specialized elongation for speed.^[21] The cursorial limb proportion score, derived from deviations in lower-leg length beyond allometric predictions, further quantifies this, with positive values (>0) denoting enhanced cursorial potential in diverse taxa.^[22]

Adaptations in Vertebrates

In vertebrates adapted for cursorial locomotion, skeletal modifications prioritize stability, weight reduction, and efficient force transmission during high-speed running. The vertebral column often exhibits increased rigidity through reinforced zygapophyseal joints and reduced intervertebral flexibility, particularly in the lumbar region, enabling a more horizontal posture that maintains dorso-ventral stability while minimizing lateral sway at speed.^[23] In ungulates, the astragalus bone features a characteristic double-pulley morphology at its trochlear surface, which locks the ankle joint for straight-line propulsion and enhances shock absorption during impacts, while the calcaneus provides a robust lever for tendon attachment.^[24] Fused or reduced tarsal elements, such as the coalescence of the navicular and cuboid bones in advanced perissodactyls, further lighten the structure and streamline hindlimb motion.^[25] Muscular adaptations in cursorial vertebrates emphasize powerful propulsion and energy efficiency. Hindlimb muscles, including enlarged gluteals and hamstrings, generate rapid acceleration by extending the hip and knee joints forcefully; for instance, in lagomorphs and equids, these muscles constitute a larger proportion of body mass compared to non-cursorial relatives, supporting sustained trotting or galloping.^[26] Elastic tendons, notably the Achilles tendon in mammals, act as biological springs by storing strain energy during stance phase and releasing it in the swing phase, reducing the energetic cost of locomotion, with studies showing up to 35% savings in mechanical energy in some mammals.^[27] This tendon lengthening scales with body size across mammals, allowing larger cursorials to recover more elastic energy per stride despite higher absolute loads.^[28] Sensory and neural enhancements ensure precise control and environmental awareness during rapid movement. Cursorial vertebrates often possess refined vestibular systems, with enlarged semicircular canals in the inner ear providing heightened sensitivity to angular accelerations, as seen in cheetahs where canal radius correlates with peak sprint velocities exceeding 100 km/h.^[29] Visual adaptations include panoramic fields of view and elevated acuity in open-habitat species; for example, equine prey animals like horses achieve nearly 360-degree field of view through laterally placed eyes, with binocular overlap of approximately 65 degrees, facilitating early predator detection at distances over 100 meters.^[30] A key vertebrate-specific metric of cursoriality is forelimb-hindlimb asymmetry, where hindlimbs are typically longer than forelimbs to optimize thrust generation while forelimbs focus on stability and shock absorption. This disparity is evident in taxa like cervids and canids, where elongated hindlimb segments (e.g., femur and tibia) increase stride length without compromising maneuverability.^[31]

Adaptations in Arthropods

Arthropods exhibit cursorial adaptations centered on their chitinous exoskeleton, which provides structural support while minimizing mass to facilitate rapid locomotion. In cursorial spiders such as wolf spiders (family Lycosidae), the exoskeleton features lightweight reinforcement in the legs through layered chitin cuticles that balance rigidity and flexibility, allowing for high-speed pursuits without excessive weight. This contrasts with more heavily armored non-cursorial arthropods, where reduced sclerotization in the prosoma and opisthosoma minimizes drag and enhances agility during ground hunting.^[32] Leg morphology in cursorial arthropods is optimized for stride length and stability. Insects like tiger beetles (Cicindelidae) possess elongated tarsi and metatarsi, forming long, thin segments that enable rapid, alternating strides for burst speeds essential in predatory chases. In spiders, the eight-legged configuration supports an alternating tetrapod gait, where legs move in anti-phase sets to maintain dynamic stability on uneven terrain, preventing tipping during acceleration. This multi-legged alternation provides greater base support than the bipedal or quadrupedal patterns seen in vertebrate cursorials, adapting to the arthropod's sprawled posture.^[33]^[34] Muscular and hydraulic systems enable powerful leg movements in the absence of traditional extensors. Spiders rely on hemolymph pressure generated in the prosoma to hydraulically extend legs, with contractions of dorsoventral muscles driving fluid into joint lacunae for propulsion during escape or hunting bursts. Complementing this, fast-twitch flexor muscles in proximal joints produce rapid contractions, achieving speeds up to about 1 m/s in species like huntsman spiders (Sparassidae), sufficient for capturing evasive prey.^[32]^[35] In insects, similar hydraulic assistance via hemocoel pressure aids leg extension alongside direct muscle action in elongated limbs. Sensory adaptations enhance navigation and prey detection during high-speed runs. Tactile hairs (setae) on spider tarsi and metatarsi detect micro-obstacles and terrain irregularities by deflecting at low forces (in the μN range), triggering leg adjustments to avoid collisions in low-light conditions. Compound eyes in cursorial insects like tiger beetles provide wide-field motion detection through ommatidial arrays, enabling quick responses to moving targets while maintaining forward momentum. These integumentary sensors integrate with hydraulic feedback for precise control over rough substrates.^[36]^[37]

Cursorial Taxa

Mammals

Mammalian cursorials represent a diverse array of species adapted for rapid terrestrial locomotion. These animals have evolved specialized traits enabling high-speed running across open habitats, contributing significantly to ecosystem dynamics through predation, herbivory, and migration. Cursoriality in mammals is particularly prominent in orders such as Perissodactyla, Artiodactyla, and Carnivora, where species exploit varied strategies for survival in expansive environments like grasslands and savannas.^[38]^[39] In Perissodactyla, such as horses (Equidae), cursorial adaptations include single-toed hooves that distribute weight evenly on firm, flat terrains, facilitating sustained speeds over long distances. Artiodactyls, including antelopes (Bovidae), feature cloven hooves that enhance agility and traction on uneven or soft ground, allowing quick maneuvers during pursuits or evasions. Within Carnivora, predators like cheetahs (Felidae) possess semi-retractable claws that provide grip during acceleration, enabling explosive sprints essential for hunting. These family-specific traits underscore the breadth of cursorial diversity among mammals.^[1]^[40]^[41] Cursorial mammals occupy distinct ecological niches, with predatory forms relying on burst speed to capture prey and herbivorous prey species emphasizing endurance to outlast pursuers. In savanna ecosystems, for instance, cursorial predators like cheetahs use their velocity to close distances on fleet-footed artiodactyls, maintaining population balances through selective hunting. Conversely, prey cursorials, such as various antelopes, leverage stamina to evade threats over extended chases, influencing vegetation patterns via grazing migrations. This predator-prey dichotomy highlights the role of cursoriality in shaping community structures.^[12]^[42] Representative examples illustrate these adaptations vividly: the pronghorn antelope (Antilocapra americana), an artiodactyl, holds the record for mammalian endurance running, sustaining an average speed of 65 km/h over 11 kilometers to escape predators.^[43] Domesticated greyhound dogs (Canis lupus familiaris), selectively bred within Carnivora for racing, exemplify sprint-oriented cursoriality, achieving bursts exceeding 70 km/h through streamlined builds optimized for short-distance pursuits.^[44] For comparative context, mammalian cursorials like pronghorns parallel avian speedsters such as ostriches in endurance capabilities across open plains.

Birds

Cursorial birds are primarily flightless or ground-dwelling species that have evolved bipedal locomotion for efficient running across open terrains, relying on speed and endurance rather than flight for survival.^[45] These adaptations are most prominent in the ratite order (Palaeognathae), which includes large, flightless birds like ostriches, emus, and rheas, as well as some ground-running species in other orders such as the Cuculiformes.^[46] Among ratites, the ostrich (Struthio camelus) exemplifies extreme cursorial specialization, with its two-toed feet reducing weight and enhancing stride efficiency, allowing sustained speeds of up to 50 km/h and bursts reaching 70 km/h.^[47] The greater rhea (Rhea americana), a South American plains specialist, achieves similar velocities of up to 60 km/h using three-toed feet for better traction on grassy terrains.^[48] In Australia, the emu (Dromaius novaehollandiae) serves as an endurance runner, capable of maintaining 48 km/h over long distances to cover vast arid landscapes.^[49] These birds feature reduced, vestigial wings that provide balance during high-speed maneuvers rather than lift, paired with powerfully muscled legs that constitute a significant portion of their body mass.^[50] Ground-running birds outside ratites, such as the greater roadrunner (Geococcyx californianus) in the Cuculiformes order, demonstrate cursorial traits for short bursts, reaching 32 km/h with zygodactyl feet that offer superior grip on desert soils for chasing prey and evading predators.^[51] These adaptations prioritize terrestrial agility, with elongated hindlimbs and specialized tendons that store elastic energy for explosive acceleration.^[52] Ecologically, cursorial birds like ratites forage in open plains and grasslands, using their speed to access scattered vegetation and insects while avoiding aerial predation through ground-based evasion.^[53] Ostriches and rheas, for instance, form loose groups in these habitats to enhance vigilance, relying on rapid flightless sprints—often zigzagging for stability—to outpace terrestrial threats.^[46] This bipedal strategy contrasts with mammalian endurance runners by emphasizing stride length over quadrupedal stability, enabling efficient energy use in expansive, predator-rich environments.^[45]

Other Groups

Among reptiles, cursorial adaptations are prominent in certain lizards, such as fringe-toed lizards (Uma spp.), which employ a sprawling, spider-like running gait enhanced by toe fringes that increase surface area for propulsion on sandy substrates.^[54] Some snakes achieve high terrestrial speeds through lateral undulation, a mode of locomotion where body waves push against environmental points of contact to generate forward thrust, enabling rapid escapes.^[55] In insects, ground beetles (Carabidae) exemplify cursorial forms with elongated legs that facilitate swift running for predation and evasion, often at nocturnal speeds exceeding 1 m/s.^[56] Cockroaches (Periplaneta americana) demonstrate remarkable burst speeds of up to 1.5 m/s during quadrupedal or bipedal locomotion, transitioning gaits at high velocities to maximize escape performance.^[57] In extinct groups, many theropod dinosaurs displayed cursorial adaptations, such as elongated metatarsals, facilitating high-speed predatory pursuits.^[8] Cursorial traits appear in minor groups like crustaceans, where land crabs (e.g., Gecarcoidea spp.) possess modified pereopods with strengthened joints and reduced webbing for stable, rapid terrestrial ambulation.^[58] Rare amphibians, such as the natterjack toad (Epidalea calamita), exhibit cursorial behavior through intermittent running rather than hopping, supported by a lateral-sequence gait for efficient ground coverage.^[59] Overall, such adaptations are less prevalent in reptiles, arthropods, and other minor taxa compared to mammals and birds, primarily due to scaling constraints where smaller body sizes limit stride length and energetic efficiency in sustained running.^[60]

Evolutionary Aspects

Origins and Development

Cursorial traits, characterized by adaptations for efficient terrestrial running, first emerged in the Permian period among therapsids, the ancestral group to mammals, where elongated limbs and shifts toward more erect postures began to facilitate faster locomotion compared to earlier sprawling synapsids.^[61] These early developments are evident in fossil therapsids like dicynodonts, which displayed preliminary changes in limb orientation that presaged cursorial capabilities, though full parasagittal posture evolved later.^[62] In parallel, Triassic archosaurs, precursors to birds and crocodilians, exhibited nascent cursorial features; for instance, the stem-archosaur Euparkeria capensis from approximately 250 million years ago possessed intermediate limb morphologies with elongated hindlimbs suited for rapid quadrupedal terrestrial movement, marking a key step in archosaur locomotor evolution.^[63] The developmental trajectory of cursorial locomotion involved a major transition from sprawling to parasagittal limb postures during the Mesozoic era, occurring independently in synapsid and archosauromorph lineages as terrestrial environments diversified.^[61] This shift, which reduced lateral limb excursion and improved stride efficiency, is documented in therapsid fossils showing progressive forelimb reconfiguration.^[64] Following the Cretaceous-Paleogene extinction event around 66 million years ago, cursorial traits co-evolved with the expansion of open habitats, as surviving mammals radiated into grasslands and underwent adaptive changes in tarsal and limb morphology to enhance speed and endurance in less cluttered environments.^[65] Fossil evidence highlights these origins and stages, with Euparkeria skeletons revealing elongated hindlimbs and ankle structures suited for rapid terrestrial movement, bridging early diapsid reptiles to more derived cursorial forms.^[63] In the mammalian lineage, the horse family (Equidae) provides a well-preserved record of progressive cursorial refinement from the Eocene epoch onward; early equids like Hyracotherium (around 55 million years ago) had four functional digits for versatile forest locomotion, but by the Miocene (approximately 20 million years ago), taxa such as Merychippus showed digit reduction to three, culminating in the monodactyl Equus by the Pleistocene, where a single robust toe minimized rotational inertia and supported high-speed running on open plains.^[66] Gaps in the fossil record persist, particularly for mid-Mesozoic transitions, due to limited preservation of soft tissues and intermediate forms, complicating precise timelines for posture shifts.^[67] Phylogenetically, cursorial traits exhibit convergent evolution across multiple vertebrate lineages rather than forming a monophyletic group, arising independently in therapsids/mammals and archosaurs/birds in response to similar selective pressures for speed in expanding terrestrial niches.^[61] This distributed pattern underscores the non-homologous nature of cursorial adaptations, with modern examples like ungulate mammals and ground birds representing diverse outcomes of these ancient evolutionary trajectories.^[68]

Implications in Evolutionary Theory

The expansion of open ecosystems, such as savannas during the Miocene, played a pivotal role in driving speciation among cursorial taxa by favoring locomotor specializations that enabled exploitation of vast, predator-rich landscapes. In placental mammals, the mid-Eocene onset of grasslands prompted a shift toward unguligrade cursoriality in herbivores, allowing larger body sizes and efficient grazing, while digitigrade adaptations in carnivores supported predation on fleet-footed prey, leading to concatenated distributions of body sizes and locomotor modes that promoted diversification. This biotic interplay, exemplified by an evolutionary "arms race" between long-limbed ungulates and pursuing carnivores, resulted in peak morphological disparity around 16–12 million years ago, underscoring cursoriality's contribution to lineage splitting in expansive habitats.^[69] Selective pressures from predation and seasonal migration strongly favored cursorial traits, enhancing survival through sustained speed and endurance, though these came at the cost of versatility in other locomotor modes. In open environments, the need to outrun predators like felids drove elongation of distal limb segments in species such as hares, prioritizing velocity over maneuverability in dense cover.^[70] Similarly, long-distance migrations imposed energetic demands that selected for efficient running gaits in ungulates, but this specialization often reduced climbing proficiency by promoting hinge-like joints and reduced proximal limb robustness, limiting access to arboreal refuges. Across diverse taxa including mammals and birds, these pressures illustrate how cursoriality facilitated niche partitioning while enforcing functional trade-offs. In contemporary contexts, cursorial adaptations pose conservation challenges amid habitat fragmentation, which disrupts migration routes for endurance specialists like pronghorn antelope, potentially elevating extinction risks by confining populations to suboptimal patches.^[71] Biomechanical modeling further illuminates these dynamics, simulating how cursorial limb proportions influence evolutionary trajectories in simulations of theropod and mammalian locomotion, revealing performance optima under varying environmental selective regimes. Cursorial traits often functioned as exaptations, repurposed from ancestral roles; for instance, the bipedal hindlimb structure inherited by birds from cursorial theropod dinosaurs initially supported terrestrial sprinting before facilitating flight and perching. The "cursorial hypothesis" for avian flight origins, positing that proto-wings evolved from running leaps for balance and propulsion, has faced critique for inadequately accounting for aerodynamic inefficiencies in ground-based ascent and the gliding-suited feathers of early avians like Archaeopteryx.

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Functional anatomy and muscle moment arms of the pelvic limb of ...
Sep 24, 2008 · Greyhounds are elite sprinting athletes, having undergone intense artificial selection for high-speed running and aerobic stamina. They reach ...
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Bipedal locomotion in ratites (Paleognatiform): Examples of cursorial ...
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The world's fastest animals | Live Science
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12 Fastest-Running Birds in the World by Land Speed - Avibirds.com
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Greater Roadrunner Overview, All About Birds, Cornell Lab of ...
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Greater Roadrunner - American Bird Conservancy
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Mechanics of a rapid running insect: two-, four- and six-legged ...
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From sprawling to parasagittal locomotion in Therapsida
Oct 5, 2022 · Most recent mammals walk with both limb pairs moved in a parasagittal plane. The swinging scapula provides an additional leg segment that adds ...
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Quantitative biomechanical assessment of locomotor capabilities of ...
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https://pmc.ncbi.nlm.nih.gov/articles/PMC5577487/
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Quantitative assessment of tarsal morphology illuminates locomotor ...
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Mechanics of evolutionary digit reduction in fossil horses (Equidae)
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Adaptive landscapes unveil the complex evolutionary path from ...
The 'sprawling-parasagittal' postural transition is a key part of mammalian evolution, associated with sweeping reorganization of the postcranial skeleton ...
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Cursorial ecomorphology and temporal patterns in theropod ...
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