Constructing and testing hypotheses of dinosaur foot motions from fossil tracks using digitization and simulation

Whilst bones present a static view of extinct animals, fossil footprints are a direct record of the activity and motion of the track maker. Deep footprints are a particularly good record of foot motion. Such footprints rarely look like the feet that made them; the sediment being heavily disturbed by the foot motion. Because of this, such tracks are often overlooked or dismissed in preference for more foot‐like impressions. However, the deeper the foot penetrates the substrate, the more motion is captured in the sediment volume. We have used deep, penetrative, Jurassic dinosaur tracks which have been naturally split into layers, to reconstruct foot motions of animals living over 200 million years ago. We consider these reconstructions to be hypotheses of motion. To test these hypotheses, we use the Discrete Element Method, in which individual particles of substrate are simulated in response to a penetrating foot model. Simulations that produce virtual tracks morphologically similar to the fossils lend support to the motion being plausible, while simulations that result in very different final tracks serve to reject the hypothesis of motion and help generate a new hypothesis

Arched footprints preserve the motions of fossil hominin feet

The longitudinal arch of the human foot is viewed as a pivotal adaptation for bipedal walking and running. Fossil footprints from Laetoli, Tanzania, and Ileret, Kenya, are believed to provide direct evidence of longitudinally arched feet in hominins from the Pliocene and Pleistocene, respectively. We studied the dynamics of track formation using biplanar X-ray, three-dimensional animation and discrete element particle simulation. Here, we demonstrate that longitudinally arched footprints are false indicators of foot anatomy; instead they are generated through a specific pattern of foot kinematics that is characteristic of human walking. Analyses of fossil hominin tracks from Laetoli show only partial evidence of this walking style, with a similar heel strike but a different pattern of propulsion. The earliest known evidence for fully modern human-like bipedal kinematics comes from the early Pleistocene Ileret tracks, which were presumably made by members of the genus Homo. This result signals important differences in the foot kinematics recorded at Laetoli and Ileret and underscores an emerging picture of locomotor diversity within the hominin clade

Integration of biplanar X-ray, three-dimensional animation and particle simulation reveals details of human ‘track ontogeny’

The emergence of bipedalism had profound effects on human evolutionary history, but the evolution of locomotor patterns within the hominin clade remains poorly understood. Fossil tracks record in vivo behaviours of extinct hominins, and they offer great potential to reveal locomotor patterns at various times and places across the human fossil record. However, there is no consensus on how to interpret anatomical or biomechanical patterns from tracks due to limited knowledge of the complex foot–substrate interactions through which they are produced. Here, we implement engineering-based methods to understand human track formation with the ultimate goal of unlocking invaluable information on hominin locomotion from fossil tracks. We first developed biplanar X-ray and three-dimensional animation techniques that permit visualization of subsurface foot motion as tracks are produced, and that allow for direct comparisons of foot kinematics to final track morphology. We then applied the discrete element method to accurately simulate the process of human track formation, allowing for direct study of human track ontogeny. This window lets us observe how specific anatomical and/or kinematic variables shape human track morphology, and it offers a new avenue for robust hypothesis testing in order to infer patterns of foot anatomy and motion from fossil hominin tracks

It's in the loop: shared sub-surface foot kinematics in birds and other dinosaurs shed light on a new dimension of fossil track diversity

The feet of ground-dwelling birds retain many features of their dinosaurian ancestry. Experiments with living species offer insights into the complex interplay among anatomy, kinematics and substrate during the formation of Mesozoic footprints. However, a key aspect of the track-making process, sub-surface foot movement, is hindered by substrate opacity. Here, we use biplanar X-rays to image guineafowl walking through radiolucent substrates of different consistency (solid, dry granular, firm to semi-liquid muds). Despite substantial kinematic variation, the foot consistently moves in a looping pattern below ground. As the foot sinks and then withdraws, the claws of the three main toes create entry and exit paths in different locations. Sampling these paths at incremental horizons captures two-dimensional features just as fossil tracks do, allowing depth-based zones to be characterized by the presence and relative position of digit impressions. Examination of deep, penetrative tracks from the Early Jurassic confirms that bipeds had an equivalent looping response to soft substrates approximately 200 Ma. Our integration of extant and extinct evidence demonstrates the influence of substrate properties on sinking depth and sub-surface foot motion, both of which are significant sources of track variation in the fossil record of dinosaurs

Estimating Mass Properties of Dinosaurs Using Laser Imaging and 3D Computer Modelling

Body mass reconstructions of extinct vertebrates are most robust when complete to near-complete skeletons allow the reconstruction of either physical or digital models. Digital models are most efficient in terms of time and cost, and provide the facility to infinitely modify model properties non-destructively, such that sensitivity analyses can be conducted to quantify the effect of the many unknown parameters involved in reconstructions of extinct animals. In this study we use laser scanning (LiDAR) and computer modelling methods to create a range of 3D mass models of five specimens of non-avian dinosaur; two near-complete specimens of Tyrannosaurus rex, the most complete specimens of Acrocanthosaurus atokensis and Strutiomimum sedens, and a near-complete skeleton of a sub-adult Edmontosaurus annectens. LiDAR scanning allows a full mounted skeleton to be imaged resulting in a detailed 3D model in which each bone retains its spatial position and articulation. This provides a high resolution skeletal framework around which the body cavity and internal organs such as lungs and air sacs can be reconstructed. This has allowed calculation of body segment masses, centres of mass and moments or inertia for each animal. However, any soft tissue reconstruction of an extinct taxon inevitably represents a best estimate model with an unknown level of accuracy. We have therefore conducted an extensive sensitivity analysis in which the volumes of body segments and respiratory organs were varied in an attempt to constrain the likely maximum plausible range of mass parameters for each animal. Our results provide wide ranges in actual mass and inertial values, emphasizing the high level of uncertainty inevitable in such reconstructions. However, our sensitivity analysis consistently places the centre of mass well below and in front of hip joint in each animal, regardless of the chosen combination of body and respiratory structure volumes. These results emphasize that future biomechanical assessments of extinct taxa should be preceded by a detailed investigation of the plausible range of mass properties, in which sensitivity analyses are used to identify a suite of possible values to be tested as inputs in analytical models

The influence of speed and size on avian terrestrial locomotor biomechanics: predicting locomotion in extinct theropod dinosaurs

How extinct, non-avian theropod dinosaurs moved is a subject of considerable interest and controversy. A better understanding of non-avian theropod locomotion can be achieved by better understanding terrestrial locomotor biomechanics in their modern descendants, birds. Despite much research on the subject, avian terrestrial locomotion remains little explored in regards to how kinematic and kinetic factors vary together with speed and body size. Here, terrestrial locomotion was investigated in twelve species of ground-dwelling bird, spanning a 1,780-fold range in body mass, across almost their entire speed range. Particular attention was devoted to the ground reaction force (GRF), the force that the feet exert upon the ground. Comparable data for the only other extant obligate, striding biped, humans, were also collected and studied. In birds, all kinematic and kinetic parameters examined changed continuously with increasing speed, while in humans all but one of those same parameters changed abruptly at the walk-run transition. This result supports previous studies that show birds to have a highly continuous locomotor repertoire compared to humans, where discrete ‘walking’ and ‘running’ gaits are not easily distinguished based on kinematic patterns alone. The influences of speed and body size on kinematic and kinetic factors in birds are developed into a set of predictive relationships that may be applied to extinct, non-avian theropods. The resulting predictive model is able to explain 79–93% of the observed variation in kinematics and 69–83% of the observed variation in GRFs, and also performs well in extrapolation tests. However, this study also found that the location of the whole-body centre of mass may exert an important influence on the nature of the GRF, and hence some caution is warranted, in lieu of further investigation

Hybrid Model for the Analysis of Human Gait: A Non-linear Approach

In this work, a generalization of the study of the human gait was made from already existent models in the literature, like models of Keller and Kockshenev. In this hybrid model, a strategy of metabolic energy minimization is combined in a race process, with a non-linear description of the movement of the mass center’s libration, trying to reproduce the behavior of the walk-run transition. The results of the experimental data, for different speed regimes, indicate that the perimeter of the trajectory of the mass center is a relevant quantity in the quantification of this dynamic. An experimental procedure was put into practice in collaboration with the research group in Biomedical Engineering, Basic Sciences and Laboratories of the Manuela Beltrán University in Bogotá, Colombia

Divergent evolution of terrestrial locomotor abilities in extant Crocodylia

Extant Crocodylia are exceptional because they employ almost the full range of quadrupedal footfall patterns (“gaits”) used by mammals; including asymmetrical gaits such as galloping and bounding. Perhaps this capacity evolved in stem Crocodylomorpha, during the Triassic when taxa were smaller, terrestrial, and long-legged. However, confusion about which Crocodylia use asymmetrical gaits and why persists, impeding reconstructions of locomotor evolution. Our experimental gait analysis of locomotor kinematics across 42 individuals from 15 species of Crocodylia obtained 184 data points for a wide velocity range (0.15–4.35 ms−1). Our results suggest either that asymmetrical gaits are ancestral for Crocodylia and lost in the alligator lineage, or that asymmetrical gaits evolved within Crocodylia at the base of the crocodile line. Regardless, we recorded usage of asymmetrical gaits in 7 species of Crocodyloidea (crocodiles); including novel documentation of these behaviours in 5 species (3 critically endangered). Larger Crocodylia use relatively less extreme gait kinematics consistent with steeply decreasing athletic ability with size. We found differences between asymmetrical and symmetrical gaits in Crocodylia: asymmetrical gaits involved greater size-normalized stride frequencies and smaller duty factors (relative ground contact times), consistent with increased mechanical demands. Remarkably, these gaits did not differ in maximal velocities obtained: whether in Alligatoroidea or Crocodyloidea, trotting or bounding achieved similar velocities, revealing that the alligator lineage is capable of hitherto unappreciated extreme locomotor performance despite a lack of asymmetrical gait usage. Hence asymmetrical gaits have benefits other than velocity capacity that explain their prevalence in Crocodyloidea and absence in Alligatoroidea—and their broader evolution

Shake a tail feather: the evolution of the theropod tail into a stiff aerodynamic surface

Theropod dinosaurs show striking morphological and functional tail variation; e.g., a long, robust, basal theropod tail used for counterbalance, or a short, modern avian tail used as an aerodynamic surface. We used a quantitative morphological and functional analysis to reconstruct intervertebral joint stiffness in the tail along the theropod lineage to extant birds. This provides new details of the tail's morphological transformation, and for the first time quantitatively evaluates its biomechanical consequences. We observe that both dorsoventral and lateral joint stiffness decreased along the non-avian theropod lineage (between nodes Theropoda and Paraves). Our results show how the tail structure of non-avian theropods was mechanically appropriate for holding itself up against gravity and maintaining passive balance. However, as dorsoventral and lateral joint stiffness decreased, the tail may have become more effective for dynamically maintaining balance. This supports our hypothesis of a reduction of dorsoventral and lateral joint stiffness in shorter tails. Along the avian theropod lineage (Avialae to crown group birds), dorsoventral and lateral joint stiffness increased overall, which appears to contradict our null expectation. We infer that this departure in joint stiffness is specific to the tail's aerodynamic role and the functional constraints imposed by it. Increased dorsoventral and lateral joint stiffness may have facilitated a gradually improved capacity to lift, depress, and swing the tail. The associated morphological changes should have resulted in a tail capable of producing larger muscular forces to utilise larger lift forces in flight. Improved joint mobility in neornithine birds potentially permitted an increase in the range of lift force vector orientations, which might have improved flight proficiency and manoeuvrability. The tail morphology of modern birds with tail fanning capabilities originated in early ornithuromorph birds. Hence, these capabilities should have been present in the early Cretaceous, with incipient tail-fanning capacity in the earliest pygostylian birds

Dinosaur Speed Demon: The Caudal Musculature of Carnotaurus sastrei and Implications for the Evolution of South American Abelisaurids

In the South American abelisaurids Carnotaurus sastrei, Aucasaurus garridoi, and, to a lesser extent Skorpiovenator bustingorryi, the anterior caudal ribs project at a high dorsolateral inclination and have interlocking lateral tips. This unique morphology facilitated the expansion of the caudal hypaxial musculature at the expense of the epaxial musculature. Distinct ridges on the ventrolateral surfaces of the caudal ribs of Aucasaurus garridoi are interpreted as attachment scars from the intra caudofemoralis/ilio-ischiocaudalis septa, and confirm that the M. caudofemoralis of advanced South American abelisaurids originated from a portion of the caudal ribs. Digital muscle models indicate that, relative to its overall body size, Carnotaurus sastrei had a substantially larger M. caudofemoralis than any other theropod yet studied. In most non-avian theropods, as in many extant sauropsids, the M. caudofemoralis served as the primary femoral retractor muscle during the locomotive power stroke. This large investment in the M. caudofemoralis suggests that Carnotaurus sastrei had the potential for great cursorial abilities, particularly short-burst sprinting. However, the tightly interlocking morphology of the anterior caudal vertebrae implies a reduced ability to make tight turns. Examination of these vertebral traits in evolutionary context reveals a progressive sequence of increasing caudofemoral mass and tail rigidity among the Abelisauridae of South America