The biomechanical importance of the scaphoid-centrale fusion during simulated knuckle-walking and its implications for human locomotor evolution

© 2020, The Author(s). Inferring the locomotor behaviour of the last common ancestor (LCA) of humans and African apes is still a divisive issue. An African great-ape-like ancestor using knuckle-walking is still the most parsimonious hypothesis for the LCA, despite diverse conflicting lines of evidence. Crucial to this hypothesis is the role of the centrale in the hominoid wrist, since the fusion of this bone with the scaphoid is among the clearest morphological synapomorphies of African apes and hominins. However, the exact functional significance of this fusion remains unclear. We address this question by carrying out finite element simulations of the hominoid wrist during knuckle-walking by virtually generating fused and unfused morphologies in a sample of hominoids. Finite element analysis was applied to test the hypothesis that a fused scaphoid-centrale better withstands the loads derived from knuckle-walking. The results show that fused morphologies display lower stress values, hence supporting a biomechanical explanation for the fusion as a functional adaptation for knuckle-walking. This functional interpretation for the fusion contrasts with the current inferred positional behaviour of the earliest hominins, thus suggesting that this morphology was probably retained from an LCA that exhibited knuckle-walking as part of its locomotor repertoire and that was probably later exapted for other functions

Knee function through finite element analysis and the role of Miocene hominoids in our understanding of the origin of antipronograde behaviours: the Pierolapithecus catalaunicus patella as a case study

Although extensive research has been carried out in recent years on the origin and evolution of human bipedalism, a full understanding of this question is far from settled. Miocene hominoids are key to a better understanding of the locomotor types observed in living apes and humans. Pierolapithecus catalaunicus, an extinct stem great ape from the middle Miocene (c. 12.0 Ma) of the Vallès-Penedès Basin (north-eastern Iberian Peninsula), is the first undoubted hominoid with an orthograde (erect) body plan. Its locomotor repertoire included above-branch quadrupedalism and other antipronograde behaviours. Elucidating the adaptive features present in the Pierolapithecus skeleton and its associated biomechanics helps us to better understand the origin of hominoid orthogrady. This work represents a new biomechanical perspective on Pierolapithecus locomotion, by studying its patella and comparing it with those drawn from a large sample of extant anthropoids. This is the first time that the biomechanical patellar performance in living non-human anthropoids and a stem hominid has been studied using finite element analysis (FEA). Differences in stress distribution are found depending on body plan and the presence/absence of a distal apex, probably due to dissimilar biomechanical performances. Pierolapithecus’ biomechanical response mainly resembles that of great apes, suggesting a similar knee joint use in mechanical terms. These results underpin previous studies on Pierolapithecus, favouring the idea that a relevant degree of some antipronograde behaviour may have made up part of its locomotor repertoire. Moreover, our results corroborate the presence of modern great ape-like knee biomechanical performances back in the Miocene

Multibody dynamics analysis (MDA) as a numerical modelling tool to reconstruct the function and palaeobiology of extinct organisms

Recent advances in computer technology have substantially changed the field of palaeontology in the last two decades. Palaeontologists now have a whole new arsenal of powerful digital techniques available to study fossil organisms in unprecedented detail and to test hypotheses regarding function and behaviour. Multibody dynamics analysis (MDA) is one of these techniques and although it originated as a tool used in the engineering and automotive industry, it holds great potential to address palaeontological questions as well. MDA allows the simulation of dynamic movements in complex objects consisting of multiple linked components. As such, this technique is ideally suited to model biological structures and to obtain quantifiable results that can be used to test the function of musculoskeletal systems rigorously. However, despite these advantages, MDA has seen a slow uptake by the palaeontological community. The most likely reason for this lies in the steep learning curve and complexity of the method. This paper provides an overview of the underlying principles of MDA and outlines the main steps involved in conducting analyses. A number of recent studies using MDA to reconstruct the palaeobiology of fossil organisms are presented and the potential for future studies is discussed. Similar to other computational techniques, including finite element analysis and computational fluid dynamics, the non‐invasive and exploratory power of MDA makes it ideally suited to study the form and function in vertebrates for which no modern analogues exist

Evolution of axial regionalization in Aves during the Mesozoic and its impact on the survival of modern lineages to K/Pgmass extinction

Archivo que contiene el resumen y la presentación del estudio presentado en este congreso.The axial column of Neornithes (modern birds) is characterized by regional fusions in caudal vertebrae (pygostyle), lumbosacrals (synsacrum), and thoracics (notarium in several taxa) that provide a rigid and stable axis during flight. Such a configuration integrates into a body plan highly suited for wing-assisted locomotion (with feathered forelimbs, modified girdles, and crouched limbs) that evolved from running dinosaurs and stem birds over the last ~150 million years. Shifts in count numbers and fusion of vertebrae have had paramount implications on the avian diversification and flight refinement. However, how the organization of precaudal vertebrae evolved across the dinosaur–bird lineage, and how and when the highly tuned axial column of neornithines was acquired are unexplored. Here, we quantify vertebral numbers in pennaraptoran dinosaurs –including Aves—, and show how the axial configuration of birds was driven from different shifts between two primary developmental mechanisms of body-axis organization: segmentation and homeotic regionalization. We demonstrate that the configuration highly tuned for flight of modern birds was not fully acquired until the appearance of Neornithes. The acquisition of a trunk-sacrum configuration more efficient to deal with stresses derived from the flapping flight could be a key factor in the survivorship of neornithines and the extinction of non-neornithine birds during the end-Cretaceous mass extinction event.Ministerio de Ciencia, Innovación y Universidades (proyectos CGL2015-68300-P y PID2019-111185GB-I00) Junta de Andalucía (proyectos P18-FR3193 y PAIDI-DOC-00095) Natural History Museum of Los ángeles County (project ‘Aerodynamics of early birds’) Universidad de Málaga. Campus de Excelencia Internacional Andalucía Tech

Comparative 3D analyses and palaeoecology of giant early amphibians (Temnospondyli: Stereospondyli)

Macroevolutionary, palaeoecological and biomechanical analyses in deep time offer the possibility to decipher the structural constraints, ecomorphological patterns and evolutionary history of extinct groups. Here, 3D comparative biomechanical analyses of the extinct giant early amphibian group of stereospondyls together with living lissamphibians and crocodiles, shows that: i) stereospondyls had peculiar palaeoecological niches with proper bites and stress patterns very different than those of giant salamanders and crocodiles; ii) their extinction may be correlated with the appearance of neosuchians, which display morphofunctional innovations. Stereospondyls weathered the end-Permian mass extinction, re-radiated, acquired gigantic sizes and dominated (semi) aquatic ecosystems during the Triassic. Because these ecosystems are today occupied by crocodilians, and stereospondyls are extinct amphibians, their palaeobiology is a matter of an intensive debate: stereospondyls were a priori compared with putative living analogous such as giant salamanders and/or crocodilians and our new results try to close this debate.Peer ReviewedPostprint (published version

Cranial biomechanics in basal urodeles: the Siberian salamander (Salamandrella keyserlingii) and its evolutionary and developmental implications

Developmental changes in salamander skulls, before and after metamorphosis, afect the feeding capabilities of these animals. How changes in cranial morphology and tissue properties afect the function of the skull are key to decipher the early evolutionary history of the crown-group of salamanders. Here, 3D cranial biomechanics of the adult Salamandrella keyserlingii were analyzed under diferent tissue properties and ossifcation sequences of the cranial skeleton. This helped unravel that: (a) Mechanical properties of tissues (as bone, cartilage or connective tissue) imply a consensus between the stifness required to perform a function versus the fxation (and displacement) required with the surrounding skeletal elements. (b) Changes on the ossifcation pattern, producing fontanelles as a result of bone loss or failure to ossify, represent a trend toward simplifcation potentially helping to distribute stress through the skull, but may also imply a major destabilization of the skull. (c) Bone loss may be originated due to biomechanical optimization and potential reduction of developmental costs. (d) Hynobiids are excellent models for biomechanical reconstruction of extinct early urodeles

Biomechanics and hydrodynamics of prey capture in the Chinese giant salamander reveal a high-performance jaw-powered suction feeding mechanism

During the evolutionary transition from fish to tetrapods, a shift from uni- to bidirectional suction feeding systems followed a reduction in the gill apparatus. Such a shift can still be observed during metamorphosis of salamanders, although many adult salamanders retain their aquatic lifestyle and feed by high-performance suction.Unfortunately, little is known about the interplay between jaws and hyobranchial motions to generate bidirectional suction flows. Here,we study the cranial morphology, aswell as kinematic and hydrodynamic aspects related to prey capture in the Chinese giant salamander (Andrias davidianus). Compared with fish and previously studied amphibians, A. davidianus uses an alternative suction mechanismthat mainly relies on accelerating water by separating the ‘plates’ formed by the long and broad upper and lower jaw surfaces. Computational fluid dynamics simulations, based on three-dimensional morphology and kinematical data from high-speed videos, indicate that the viscerocranial elements mainly serve to accommodate the water that was given a sufficient anterior-to-posterior impulse beforehand by powerful jawseparation.We hypothesize that this modifiedway of generating suction is primitive for salamanders, and that this behaviour could have played an important role in the evolution of terrestrial life in vertebrates by releasing mechanical constraints on the hyobranchial system

3D bite modeling and feeding mechanics of the largest living amphibian, the Chinese Giant Salamander Andrias davidianus (Amphibia:Urodela)

Biting is an integral feature of the feeding mechanism for aquatic and terrestrial salamanders to capture, fix or immobilize elusive or struggling prey. However, little information is available on how it works and the functional implications of this biting system in amphibians although such approaches might be essential to understand feeding systems performed by early tetrapods. Herein, the skull biomechanics of the Chinese giant salamander, Andrias davidianus is investigated using 3D finite element analysis. The results reveal that the prey contact position is crucial for the structural performance of the skull, which is probably related to the lack of a bony bridge between the posterior end of the maxilla and the anterior quadrato-squamosal region. Giant salamanders perform asymmetrical strikes. These strikes are unusual and specialized behavior but might indeed be beneficial in such sit-and-wait or ambush-predators to capture laterally approaching prey. However, once captured by an asymmetrical strike, large, elusive and struggling prey have to be brought to the anterior jaw region to be subdued by a strong bite. Given their basal position within extant salamanders and theirPeer ReviewedPostprint (published version

Biomechanical performance of an immature maxillary central incisor after revitalization: a finite element analysis

Aim To investigate the stress distribution in an immature maxillary incisor and the same tooth after simulated revitalization with deposition of tubular dentine or cementum by finite element analysis (FEA). Methodology A finite element model of a maxillary central incisor was developed on the basis of a mu CT scan. The tooth was segmented in two parts: a part that represented a tooth in an immature state and an apical part that represented the tissue formed after revitalization. The apical part was given the mechanical properties of dentine or cementum. The immature tooth and the same tooth reinforced by either dentine or cementum underwent simulation of biting, trauma and orthodontic movement. Von Mises stress values were compared between the scenarios and tooth segments. Results Maximum stress in the immature incisor developed apically; however, dentine- and cementum-reinforced teeth revealed the greatest stress in the external portion of the root decreasing towards the apex. Greatest mechanical stress was caused by dental trauma perpendicular to the long axis of the root followed by biting and orthodontic movement. Stress peaks were lower in the dentine-reinforced tooth compared with the cementum-reinforced tooth in all scenarios; however, median stress in the immature part was reduced irrespective of dentine or cementum deposition. Dentine reinforcement caused greater stress values in the apical segment due to absorbance of the applied force, whereas stress was not transferred towards deposited cementum. Conclusions Apposition of simulated hard tissue in a maxillary central incisor after revitalization reduced mechanical stress in the immature tooth. Formation of dentine was advantageous because, unlike cementum, it facilitated an even stress distribution throughout the root resulting in lower stress values

A biomechanical approach to understand the ecomorphological relationship between primate mandibles and diet

The relationship between primate mandibular form and diet has been previously analysed by applying a wide array of techniques and approaches. Nonetheless, most of these studies compared few species and/or infrequently aimed to elucidate function based on an explicit biomechanical framework. In this study, we generated and analysed 31 Finite Element planar models of different primate jaws under different loading scenarios (incisive, canine, premolar and molar bites) to test the hypothesis that there are significant differences in mandibular biomechanical performance due to food categories and/or food hardness. The obtained stress values show that in primates, hard food eaters have stiffer mandibles when compared to those that rely on softer diets. In addition, we find that folivores species have the weakest jaws, whilst omnivores have the strongest mandibles within the order Primates. These results are highly relevant because they show that there is a strong association between mandibular biomechanical performance, mandibular form, food hardness and diet categories and that these associations can be studied using biomechanical techniques rather than focusing solely on morphology