Dinosaur locomotion: beyond the bones

The skeletons of dinosaurs, particularly large-bodied, enigmatic forms like Tyrannosaurus and Triceratops, are awe inspiring and have been brought ‘back-to-life’ in numerous animated and motion picture productions. But how did these animals really stand and move in life and how rigorously can scientists reconstruct their locomotion? In recent years, traditional approaches, such as morphological comparisons and limb bone scaling, have been combined with new computational modelling methods to provide rich insight into dinosaur locomotion and how it may have changed during the group's long evolutionary history. However, many significant challenges remain and currently it is still difficult to constrain how any one dinosaur moved. Unknown factors, such as body mass and muscle anatomy, make it difficult to reliably reconstruct the motions and capabilities (e.g. running speeds) of dinosaurs based on data available from fossils. Thus, future work must continue to integrate information on dinosaur anatomy with principles of locomotion established in living animals, as well as embracing a range of methodological approaches.

Paleobiology

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 challenges. Building on decades of prior research, we outline and critically discuss a complete workflow for biomechanical analysis of extinct species, using locomotor biomechanics in the Triassic theropod dinosaur Coelophysis as a case study. We progress from the digital capture of fossil bone morphology to creating rigged skeletal models, to reconstructing musculature and soft tissue volumes, to the development of computational musculoskeletal models, and finally to the execution of biomechanical simulations. Using a three-dimensional musculoskeletal model comprising 33 muscles, a static inverse simulation of the mid-stance of running shows that ...

Anatomical record (Hoboken, N.J. : 2007), 2011

Fossil Record, 1999

Speeds of walking dinosaurs that left fossil trackways have been estimated using the stride length times natural pendulum frequency of the limbs. In a detailed analysis of limb movements in walking Asian elephants and giraffes, however, distinct differences between actual limb movements and the predicted limb movements using only gravity as driving force were observed. Additionally, stride frequency was highly variable. Swing time was fairly constant, but especially at high walking speeds, much shorter than half the natural pendulum period. An analysis of hip and shoulder movements during walking showed that limb swinging was influenced by accelerations of hip and shoulder joints especially at high walking speeds. These results suggest an economical fast walking mechanism that could have been utilised by large dinosaurs to increase maximum speeds of locomotion. These findings throw new light on the dynamics of large vertebrates and can be used to improve speed estimates in large dinosaurs .

Ichnos, 1994

Page 1. Ichnos, v. 3, pp. 193-202, 1994 An International Journal for Plant and Animal Traces Limping Dinosaurs? Trackway evidence for abnormal gaits Martin G. Lockley,1 Adrian P. Hunt,1 Joaquin Moratalla,2 and Masaki Matsukawa3 ...

Paleobiology, 2005

Muscle moment arms are important determinants of muscle function; however, it is challenging to determine moment arms by inspecting bone specimens alone, as muscles have curvilinear paths that change as joints rotate. The goals of this study were to (1) develop a three-dimensional graphics-based model of the musculoskeletal system of the Cretaceous theropod dinosaur Tyrannosaurus rex that predicts muscle-tendon unit paths, lengths, and moment arms for a range of limb positions; (2) use the model to determine how the T. rex hindlimb muscle moment arms varied between crouched and upright poses; (3) compare the predicted moment arms with previous assessments of muscle function in dinosaurs; (4) evaluate how the magnitudes of these moment arms compare with those in other animals; and (5) integrate these findings with previous biomechanical studies to produce a revised appraisal of stance, gait, and speed in T. rex. The musculoskeletal model includes ten degrees of joint freedom (flexion/extension, ab/adduction, or medial/ lateral rotation) and 33 main muscle groups crossing the hip, knee, ankle, and toe joints of each hindlimb. The model was developed by acquiring and processing bone geometric data, defining joint rotation axes, justifying muscle attachment sites, and specifying muscle-tendon geometry and paths. Flexor and extensor muscle moment arms about all of the main limb joints were estimated, and limb orientation was statically varied to characterize how the muscle moment arms changed. We used sensitivity analysis of uncertain parameters, such as muscle origin and insertion centroids, to deterimine how much our conclusions depend on the muscle reconstruction we adopted. This shows that a specific amount of error in the reconstruction (e.g., position of muscle origins) can have a greater, lesser, similar, or no effect on the moment arms, depending on complex interactions between components of the musculoskeletal geometry. We found that more upright poses would have improved mechanical advantage of the muscles considerably. Our analysis shows that previously assumed moment arm values were generally conservatively high. Our results for muscle moment arms are generally lower than the values predicted by scaling data from extant taxa, suggesting that T. rex did not have the allometrically large muscle moment arms that might be expected in a proficient runner. The information provided by the model is important for determining how T. rex stood and walked, and how the muscles of a 4000-7000 kg biped might have worked in comparison with extant bipeds such as birds and humans. Our model thus strengthens the conclusion that T. rex was not an exceptionally fast runner, and supports the inference that more upright (although not completely columnar) poses are more plausible for T. rex. These results confirm general principles about the relationship between size, limb orientation, and locomotor mechanics: exceptionally big animals have a more limited range of locomotor abilities and tend to adopt more upright poses that improve extensor muscle effective mechanical advantage. This model builds on previous phylogenetically based muscle reconstructions and so moves closer to a fully dynamic, three-dimensional model of stance, gait, and speed in T. rex.

Journal of Vertebrate …, 2009

Ornithischian dinosaurs were primitively bipedal with forelimbs modified for grasping, but quadrupedalism evolved in the clade on at least three occasions independently. Outside of Ornithischia, quadrupedality from bipedal ancestors has only evolved on two other occasions, making this one of the rarest locomotory transitions in tetrapod evolutionary history. The osteological and myological changes associated with these transitions have only recently been documented, and the biomechanical consequences of these changes remain to be examined. Here, we review previous approaches to understanding locomotion in extinct animals, which can be broadly split into form-function approaches using analogy based on extant animals, limb-bone scaling, and computational approaches. We then carry out the first systematic attempt to quantify changes in locomotor muscle function in bipedal and quadrupedal ornithischian dinosaurs. Using three-dimensional computational modelling of the major pelvic locomotor muscle moment arms, we examine similarities and differences among individual taxa, between quadrupedal and bipedal taxa, and among taxa representing the three major ornithischian lineages (Thyreophora, Ornithopoda, Marginocephalia). Our results suggest that the ceratopsid Chasmosaurus and the ornithopod Hypsilophodon have relatively low moment arms for most muscles and most functions, perhaps suggesting poor locomotor performance in these taxa. Quadrupeds have higher abductor moment arms than bipeds, which we suggest is due to the overall wider bodies of the quadrupeds modelled. A peak in extensor moment arms at more extended hip angles and lower medial rotator moment arms in quadrupeds than in bipeds may be due to a more columnar hindlimb and loss of medial rotation as a form of lateral limb support in quadrupeds. We are not able to identify trends in moment arm evolution across Ornithischia as a whole, suggesting that the bipedal ancestry of ornithischians did not constrain the development of quadrupedal locomotion via a limited number of functional pathways. Functional anatomy appears to have had a greater effect on moment arms than phylogeny, and the differences identified between individual taxa and individual clades may relate to differences in locomotor performance required for living in different environments or for clade-specific behaviours.

The known relationships of speed, gait and body size (derived mainly from mammals) are used to determine the gaits and theoretical maximum speeds of dinosaurs. Speed estimates are made for 62 dinosaurs (representing 51 genera) and are supplemented with information from comparative anatomy and from dinosaur trackways. It is concluded that smaller bipedal dinosaurs were capable of running at speeds up to 35 or 40 km/h. The so~called "ostrich dinosaurs" are credited with maximum speeds less than 60 kin/h, and possibly as low as 35 or 40 km/h. Larger bipedal dinosaurs were probably restricted to walking or slow trotting gaits, with maximum speeds in the range 15--20 km/h. Most quadrupedal dinosaurs seem to have been restricted to a walking gait. Stegosaurs and ankylosaurs may have had maximum speeds as low as 6--8 km/h. Sauropods may have attained 12--17 kin/h, and some ceratopsians may have been capable of trotting at speeds up to 25 km/h. On a weight-for-weight basis the speeds of dinosaurs are generally lower than those of mammals.