John R. Hutchinson

Royal Veterinary College | RVC · Department of Comparative Biomedical Sciences

The patella (kneecap) exhibits multiple evolutionary origins in birds, mammals, and lizards, and is thought to increase the mechanical advantage of the knee extensor muscles. Despite appreciable interest in the specialized anatomy and locomotion of palaeognathous birds (ratites and relatives), the structure, ontogeny and evolution of the patella in...

Crocodiles and their kin (Crocodylidae) use asymmetrical (bounding and galloping) gaits when moving rapidly. Despite being morphologically and ecologically similar, it seems alligators and their kin (Alligatoridae) do not. To investigate a possible anatomical basis for this apparent major difference in locomotor capabilities, we measured relative m...

Despite their semi-aquatic mode of life, modern crocodylians use a wide range of terrestrial locomotor behaviours, including asymmetrical gaits otherwise only found in mammals. The key to these diverse abilities may lie in the axial skeleton. Correlations between vertebral morphology and both intervertebral joint stiffness and locomotor behaviour h...

An estimated 1,170 white (Ceratotherium simum), black (Diceros bicornis), greater one-horned (Rhinoceros unicornis), and Sumatran (Dicerorhinus sumatrensis) rhinoceroses are kept in captivity worldwide, where they are a popular public attraction and serve important roles in education and conservation. Rhinoceroses in captivity are reportedly affect...

Locomotion in living birds (Neornithes) has two remarkable features: feather-assisted flight, and the use of unusually crouched hindlimbs for bipedal support and movement. When and how these defining functional traits evolved remains controversial. However, the advent of computer modelling approaches and the discoveries of exceptionally preserved k...

Lagosuchus talampayensis is a small‐bodied (~0.5 m long) Late Triassic dinosauriform archosaur from Argentina. Lagosuchus long has been a pivotal taxon for reconstructing the evolution of form and function on the dinosaur lineage. This importance is because it has a mix of ancestral archosaurian traits, such as a small pelvis with a mostly closed a...

The Kem Kem Group is a lowermost lithostratigraphic unit from the Upper Cretaceous that extends along the border between Algeria and Morocco, in the northern region of Africa. This geological unit has yielded several tetrapod fossils, including a well-represented assemblage of theropod dinosaurs, after more than eight decades of research. Here, we...

Theropods are obligate bipedal dinosaurs that appeared 230 Ma and are still extant as birds. Their history is characterized by extreme variations in body mass, with gigantism evolving convergently between many lineages. However, no quantification of hindlimb functional morphology has shown whether these body mass increases led to similar specializa...

Heavy animals incur large forces on their limb bones, due to the transmission of body weight and ground reaction forces, and the contractions of the various muscles of the limbs. This is particularly true for rhinoceroses, the heaviest extant animals capable of galloping. Several studies have examined their musculoskeletal system and the forces the...

The force a muscle generates is dependent on muscle structure, in which fibre length, pennation angle and tendon slack length all influence force production. Muscles are not preserved in the fossil record and these parameters must be estimated when constructing a musculoskeletal model. Here, we test the capability of digitally reconstructed muscles...

Piatnitzkysauridae were Jurassic theropods that represented the earliest diverging branch of Megalosauroidea, being one of the earliest lineages to have evolved moderate body size. This clade's typical body size and some unusual anatomical features raise questions about locomotor function and specializations to aid in body support; and other palaeo...

Although lameness is a common problem in elephants (Asian elephant [Elephas maximus] and African elephants Loxodonta africana and Loxodonta cyclotis) under human care, there has not been a standardized lameness assessment system to date. This study developed and evaluated a standardized system for the assessment of locomotion in elephants under hum...

Elephants are atypical of most quadrupeds in that they maintain the same lateral sequence footfall pattern across all locomotor speeds. It has been speculated that the preservation of the footfall patterns is necessary to maintain a statically stable support polygon. This should be a particularly important constraint in large, relatively slow anima...

Reptiles represent one of the most diverse groups of tetrapod vertebrates. Extant representatives of reptiles include lepidosaurs (lizards), testudines (turtles) and archosaurs (crocodiles and birds). In particular, they show an important locomotor diversity with bipedal, quadrupedal and facultatively bipedal taxa. This diversity is accompanied by...

During the Mesozoic, non-avian theropods represented one of the most successful clades globally distributed, with a wide diversity of forms. An example is the clade Megalosauroidea, which included medium- to large-bodied forms. Here, we analyse the macroevolution of the locomotor system in early Theropoda, emphasizing the Megalosauroidea. We scored...

Extant amniotes show remarkable postural diversity. Broadly speaking, limbs with erect (strongly adducted, more vertically oriented) posture are found in mammals that are particularly heavy (graviportal) or show good running skills (cursorial), while crouched (highly flexed) limbs are found in taxa with more generalized locomotion. In Reptilia, cro...

Reptiles represent one of the most diverse groups of tetrapod vertebrates. Extant representatives of reptiles include lepidosaurs (lizards), testudines (turtles) and archosaurs (crocodiles and birds). In particular, they show an important locomotor diversity with bipedal, quadrupedal and facultatively bipedal taxa. This diversity is accompanied by...

It is accepted that non-avian theropod dinosaurs, with their long muscular tails and small forelimbs, had a centre-of-mass close to the hip, while extant birds, with their reduced tails and enlarged wings have their mass centred more cranially. Transition between these states is considered crucial to two key innovations in the avian locomotor syste...

Here, we review the modern interface of three-dimensional (3D) empirical (e.g. motion capture) and theoretical (e.g. modelling and simulation) approaches to the study of terrestrial locomotion using appendages in tetrapod vertebrates. These tools span a spectrum from more empirical approaches such as XROMM, to potentially more intermediate approach...

Mammals have an evolutionary history spanning hundreds of millions of years. Today, mammals represent one of the most diverse groups of tetrapod vertebrates. In particular, they present a great postural diversity. The humerus adopts different positions: small mammals have a “crouched” posture with a quasi-horizontal humerus, while in the largest sp...

The water‐to‐land transition by the first tetrapod vertebrates represents a key stage in their evolution. Selection pressures exerted by this new environment on animals led to the emergence of new locomotor and postural strategies that favoured access to different ecological niches and contributed to their evolutionary success. Today, amniotes show...

Birds and crocodylians are the only remaining members of Archosauria (ruling reptiles) and they exhibit major differences in posture and gait, which are polar opposites in terms of locomotor strategies. Their broader lineages (Avemetatarsalia and Pseudosuchia) evolved a multitude of locomotor modes in the Triassic and Jurassic periods, including se...

Significant evolutionary shifts in locomotor behaviour often involve comparatively subtle anatomical transitions. For dinosaurian and avian evolution, medial overhang of the proximal femur has been central to discussions. However, there is an apparent conflict with regard to the evolutionary origin of the dinosaurian femoral head, with neontologica...

In vertebrates, active movement is driven by muscle forces acting on bones, either directly or through tendinous insertions. There has been much debate over how muscle size and force are reflected by the muscular attachment areas (AAs). Here we investigate the relationship between the physiological cross-sectional area (PCSA), a proxy for the force...

Joint range of motion (RoM) analyses are fundamental to our understanding of how an animal moves throughout its ecosystem. Recent technological advances allow for more detailed quantification of this RoM (e.g. including interaction of degrees of freedom) both in ex vivo joints and in vivo experiments. Both types of data have been used to draw compa...

Although shark teeth are abundant in the fossil record, their bodies are rarely preserved. Thus, our understanding of the anatomy of the extinct Otodus megalodon remains rudimentary. We used an exceptionally well-preserved fossil to create the first three-dimensional model of the body of this giant shark and used it to infer its movement and feedin...

Giraffes ( Giraffa camelopardalis ) possess specialized locomotor morphology, namely elongate and gracile distal limbs. While this contributes to their overall height and enhances feeding behavior, we propose that the combination of long limb segments and modest muscle lever arms results in low effective mechanical advantage (EMA, the ratio of in-l...

Quantifying joint range of motion (RoM), the reachable poses at a joint, has many applications in research and clinical care. Joint RoM measurements can be used to investigate the link between form and function in extant and extinct animals, to diagnose musculoskeletal disorders and injuries or monitor rehabilitation progress. However, it is diffic...

Krogh’s principle states, “For such a large number of problems there will be some animal of choice, or a few such animals, on which it can be most conveniently studied.” The downside of picking a question first and then finding an ideal organism on which to study it is that it will inevitably leave many organisms neglected. Here, we promote the inv...

Synopsis Salamanders are often used as analogs for early tetrapods in paleontological reconstructions of locomotion. However, concerns have been raised about whether this comparison is justifiable, necessitating comparisons of a broader range of early tetrapods with salamanders. Here, we test whether the osteological morphology of the hindlimb in t...

The microarchitecture of bone both results from and in turn affects the remodelling process. Bone-specific surface, for instance, is one of these important microarchitectural parameters because remodelling is also considered to be a surface-mediated phenomenon (Berli et al. 2017). An understanding of these structural parameters across the widest po...

Archosauria diversified throughout the Triassic Period before experiencing two mass extinctions near its end ∼201 Mya, leaving only the crocodile-lineage (Crocodylomorpha) and bird-lineage (Dinosauria) as survivors; along with the pterosaurian flying reptiles. About 50 years ago, the “locomotor superiority hypothesis” (LSH) proposed that dinosaurs...

Serially connected robots are promising candidates for performing tasks in confined spaces such as search and rescue in large-scale disasters. Such robots are typically limbless, and we hypothesize that the addition of limbs could improve mobility. However, a challenge in designing and controlling such devices lies in the coordination of high-dimen...

The transition from early sauropodomorphs to sauropods is of special interest given that a shift from obligatory or facultative bipedalism to an obligatory quadrupedalism is evident. In this chapter, we review and discuss the biological mechanisms underpinning such evolutionary transformations. The discovery of the South American sauropodomorph Mus...

Objectives Contrary to earlier hypotheses, a previous biomechanical analysis indicated that long-documented morphological differences between the shoulders of humans and apes do not enhance the arm-raising mechanism. Here, we investigate a different interpretation: the oblique shoulder morphology that is shared by all hominoids but humans enhances...

Biomechanical models and simulations of musculoskeletal function rely on accurate muscle parameters, such as muscle masses and lines of action, to estimate force production potential and moment arms. These parameters are often obtained through destructive techniques (i.e., dissection) in living taxa, frequently hindering the measurement of other re...

Fossil trackways provide a glimpse into the behavior of extinct animals. However, while providing information of the trackmaker size, stride, and even speed, the actual gait of the organism can be ambiguous. This is especially true of quadrupedal animals, where disparate gaits can have similar trackway patterns. Here, predictive simulation using tr...

The last common ancestor of birds and crocodylians plus all of its descendants (clade Archosauria) dominated terrestrial Mesozoic ecosystems, giving rise to disparate body plans, sizes, and modes of locomotion. As in the fields of vertebrate morphology and paleontology more generally, studies of archosaur skeletal structure have come to depend on t...

Synopsis X-ray Reconstruction of Moving Morphology (XROMM) permits researchers to see beneath the skin, usually to see musculoskeletal movements. These movements can be tracked and later used to provide information regarding the mechanics of movement. Here, we discuss “IK marker-guided rotoscoping”—a method that combines inverse kinematic solvers w...

Muscle-actuated control is a research topic of interest spanning different fields, in particular biomechanics, robotics and graphics. This type of control is particularly challenging because models are often overactuated, and dynamics are delayed and non-linear. It is however a very well tested and tuned actuation model that has undergone millions...

The evolutionary history of archosaurs and their closest relatives is characterized by a wide diversity of locomotor modes, which has even been suggested as a pivotal aspect underlying the evolutionary success of dinosaurs vs. pseudosuchians across the Triassic–Jurassic transition. This locomotor diversity (e.g., more sprawling/erect; crouched/upri...

Locomotion has influenced the ecology, evolution, and extinction of species throughout history, yet studying locomotion in the fossil record is challenging. Computational biomechanics can provide novel insight by mechanistically relating observed anatomy to whole-animal function and behavior. Here, we leverage optimal control methods to generate th...

Crocodiles are unusual reptiles which can use a broad continuum of hindlimb postures during terrestrial locomotion, ranging from a high walk to a sprawl. But how does a crocodile choose how it uses its hindlimb muscles to generate such a repertoire of movements, which in turn involves varying ranges of muscular leverage to support and move the hind...

Biomechanical musculoskeletal models rely on accurate muscle parameters, such as muscle masses and lines of action, to estimate force production and leverage of individual muscles. Unfortunately, these parameters are often obtained through destructive techniques (i.e. dissection and dice-CT scanning) in extant taxa, frequently hindering the measure...

The fourth trochanter is an attachment site for the caudofemoral musculature that is located on the posterior side of archosauriform femora. Its morphology is commonly used as a synapomorphy for various archosauriform clades such as its pendant shape in some ornithischian dinosaurs. The fourth trochanter presumably was subjected to strong mechanica...

Giant land vertebrates have evolved more than 30 times, notably in dinosaurs and mammals. The evolutionary and biomechanical perspectives considered here unify data from extant and extinct species, assessing current theory regarding how the locomotor biomechanics of giants has evolved. In terrestrial tetrapods, isometric and allometric scaling patt...

Land mammals support and move their body using their musculoskeletal system. Their musculature usually presents varying adaptations with body mass or mode of locomotion. Rhinocerotidae is an interesting clade in this regard, as they are heavy animals potentially reaching three tons but are still capable of adopting a galloping gait. However, their...

Giraffes ( Giraffa camelopardalis ) possess specialised locomotor morphology, namely elongate and gracile distal limbs. Whilst this contributes to their overall height (and enhanced feeding behaviour), we propose that the combination of long limb segments and modest muscle lever arms results in low effective mechanical advantage (EMA, the ratio of...

Jumping is a common, but demanding, behaviour that many animals employ during everyday activity. In contrast to jump-specialists such as anurans and some primates, jumping biomechanics and the factors that influence performance remains little studied for generalized species that lack marked adaptations for jumping. Computational biomechanical model...

The avian lung is highly specialized and is both functionally and morphologically distinct from that of their closest extant relatives, the crocodilians. It is highly partitioned, with a unidirectionally ventilated and immobilized gas-exchanging lung, and functionally decoupled, compliant, poorly vascularized ventilatory air-sacs. To understand the...

The arrangement and physiology of muscle fibres can strongly influence musculoskeletal function and whole-organismal performance. However, experimental investigation of muscle function during in vivo activity is typically limited to relatively few muscles in a given system. Computational models and simulations of the musculoskeletal system can part...

We developed a three‐dimensional, computational biomechanical model of a juvenile Nile crocodile (Crocodylus niloticus) pelvis and hindlimb, composed of 47 pelvic limb muscles, to investigate muscle function. We tested whether crocodiles, which are known to use a variety of limb postures during movement, use limb orientations (joint angles) that op...

Bipedal locomotion evolved along the archosaurian lineage to birds, shifting from “hip-based” to “knee-based” mechanisms. However, the roles of individual muscles in these changes and their evolutionary timings remain obscure. Using 13 three-dimensional musculoskeletal models of the hindlimbs of bird-line archosaurs, we quantify how the moment arms...

Video showing the arm abduction sequence of the gorilla musculoskeletal shoulder model.

Musculoskeletal computer models allow us to quantitatively relate morphological features to biomechanical performance. In non‐human apes, certain morphological features have long been linked to greater arm abduction potential and increased arm‐raising performance, compared to humans. Here, we present the first musculoskeletal model of a western low...

Many animals generate propulsive forces by coordinating legs, which contact and push against the surroundings, with bending of the body, which can only indirectly influence these forces. Such body–leg coordination is not commonly employed in quadrupedal robotic systems. To elucidate the role of back bending during quadrupedal locomotion, we study a...

One of the most intriguing questions in vertebrate evolution is how tetrapods gained the ability to walk on land. Although many hypotheses have been proposed, few have been rigorously tested using the fossil record. Here, we build three-dimensional musculoskeletal models of the pectoral appendage in Eusthenopteron, Acanthostega, and Pederpes and qu...

Biomechanical modelling approaches which can accurately estimate musculoskeletal functions have offered unique insights into the locomotion of extant animals, which in turn can provide the basis for simulating extinct animal movement. We tested whether Nile crocodiles, which use a variety of limb postures during movement, adopt limb orientations th...

The intersection of paleontology and biomechanics can be reciprocally illuminating, helping to improve paleobiological knowledge of extinct species and furthering our understanding of the generality of biomechanical principles derived from study of extant species. However, working with data gleaned primarily from the fossil record has its challenge...

Archosaurian reptiles (including living crocodiles and birds) had an explosive diversification of locomotor form and function since the Triassic ∼250 million years ago. Their limb muscle physiology and biomechanics are pivotal to our understanding of how their diversity and evolution relate to locomotor function. Muscle contraction velocity, force...

Throughout their 250 Myr history, archosaurian reptiles have exhibited a wide array of body sizes, shapes, and locomotor habits, especially in regard to terrestriality. These features make Archosauria a useful clade with which to study the interplay between body size, shape, and locomotor behavior, and how this interplay may have influenced locomot...

Biomechanical models can offer unique insights into animal musculoskeletal function, which can in turn provide the basis for simulating extinct animal movement. Modelling musculoskeletal function in extinct forms first ideally requires modern forms to be explored, modelled and analysed for validation purposes. For example, to model the locomotory b...

Biomechanical analyses of locomotor function in extinct taxa are challenging, as the information provided by the fossil record is often scarce and incomplete, as important soft-tissue information is seldom preserved. Individual muscles sometimes can be identified based on osteological correlates for attachment sites on the bones. Furthermore, the p...

Does the size of “muscle scars” on bones correspond to the cross-sectional areas of muscles? This question is vital for understanding morphology (e.g., musculoskeletal integration/evo-devo), biomechanics (e.g., mechanobiology) and for reconstructing musculoskeletal form and function in extinct taxa. Archosaurian reptiles famously evolved disparate...

The evolutionary history of archosaurs and their closest relatives is characterized by a wide diversity of locomotor modes. The locomotor superiority hypothesis suggests that this diversity was the main driver of early dinosaur success over pseudosuchians across the Triassic-Jurassic extinctions. The impacts of postural changes (e.g. more sprawling...

Triassic archosaurs and stem-archosaurs show a remarkable disparity in their ankle and pelvis morphologies. However, the implications of these different morphologies for specific functions are still poorly understood. Here, we present the first quantitative analysis into the locomotor abilities of a stem-archosaur applying 3D modelling techniques....