Finite element modelling of sound transmission in the Weberian apparatus of zebrafish (Danio rerio).

Zebrafish, an essential vertebrate model, has greatly expanded our understanding of hearing. However, one area that remains unexplored is the biomechanics of the Weberian apparatus, crucial for sound conduction and perception. Using micro-computed tomography (µCT) bioimaging, we created three-dimensional finite element models of the zebrafish Weberian ossicles. These models ranged from the exact size to scaled isometric versions with constrained geometry (1 to 10 mm in ossicular chain length). Harmonic finite element analysis of all 11 models revealed that the resonance frequency of the zebrafish's Weberian ossicular chain is approximately 900 Hz, matching their optimal hearing range. Interestingly, resonance frequency negatively correlated with size, while the ratio of peak displacement and difference of resonance frequency between tripus and scaphium remained constant. This suggests the transmission efficiency of the ossicular chain and the homogeneity of resonance frequency at both ends of the chain are not size-dependent. We conclude that the Weberian apparatus's resonance frequency can explain zebrafish's best hearing frequency, and their biomechanical characteristics are not influenced by isometric ontogeny. As the first biomechanical modelling of atympanic ear and among the few non-human ear modelling, this study provides a methodological framework for further investigations into hearing mechanisms and the hearing evolution of vertebrates.

One step further in biomechanical models in palaeontology: a nonlinear finite element analysis review.

Finite element analysis (FEA) is no longer a new technique in the fields of palaeontology, anthropology, and evolutionary biology. It is nowadays a well-established technique within the virtual functional-morphology toolkit. However, almost all the works published in these fields have only applied the most basic FEA tools i.e., linear materials in static structural problems. Linear and static approximations are commonly used because they are computationally less expensive, and the error associated with these assumptions can be accepted. Nonetheless, nonlinearities are natural to be used in biomechanical models especially when modelling soft tissues, establish contacts between separated bones or the inclusion of buckling results. The aim of this review is to, firstly, highlight the usefulness of non-linearities and secondly, showcase these FEA tool to researchers that work in functional morphology and biomechanics, as non-linearities can improve their FEA models by widening the possible applications and topics that currently are not used in palaeontology and anthropology.

Finite element analysis relating shape, material properties, and dimensions of taenioglossan radular teeth with trophic specialisations in Paludomidae (Gastropoda).

The radula, a chitinous membrane with embedded tooth rows, is the molluscan autapomorphy for feeding. The morphologies, arrangements and mechanical properties of teeth can vary between taxa, which is usually interpreted as adaptation to food. In previous studies, we proposed about trophic and other functional specialisations in taenioglossan radulae from species of African paludomid gastropods. These were based on the analysis of shape, material properties, force-resistance, and the mechanical behaviour of teeth, when interacting with an obstacle. The latter was previously simulated for one species (Spekia zonata) by the finite-element-analysis (FEA) and, for more species, observed in experiments. In the here presented work we test the previous hypotheses by applying the FEA on 3D modelled radulae, with incorporated material properties, from three additional paludomid species. These species forage either on algae attached to rocks (Lavigeria grandis), covering sand (Cleopatra johnstoni), or attached to plant surface and covering sand (Bridouxia grandidieriana). Since the analysed radulae vary greatly in their general size (e.g. width) and size of teeth between species, we additionally aimed at relating the simulated stress and strain distributions with the tooth sizes by altering the force/volume. For this purpose, we also included S. zonata again in the present study. Our FEA results show that smaller radulae are more affected by stress and strain than larger ones, when each tooth is loaded with the same force. However, the results are not fully in congruence with results from the previous breaking stress experiments, indicating that besides the parameter size, more mechanisms leading to reduced stress/strain must be present in radulae.

Finite element analysis of individual taenioglossan radular teeth (Mollusca).

Molluscs are a highly successful group of invertebrates characterised by a specialised feeding organ called the radula. The diversity of this structure is associated with distinct feeding strategies and ecological niches. However, the precise function of the radula (each tooth type and their arrangement) remains poorly understood. Here for the first time, we use a quantitative approach, Finite-Element-Analysis (FEA), to test hypotheses regarding the function of particular taenioglossan tooth types. Taenioglossan radulae are of special interest, because they are comprised of multiple teeth that are regionally distinct in their morphology. For this study we choose the freshwater gastropod species Spekia zonata, endemic to Lake Tanganyika, inhabiting and feeding on algae attached to rocks. As a member of the African paludomid species flock, the enigmatic origin and evolutionary relationships of this species has received much attention. Its chitinous radula comprises several tooth types with distinctly different shapes. We characterise the tooth's position, material properties and attachment to the radular membrane and use this data to evaluate 18 possible FEA scenarios differing in the above parameters. Our estimations of stress and strain indicate different functional loads for different teeth. We posit that the central and lateral teeth are best suitable for scratching substrate loosening ingesta, whereas the marginals are best suited for gathering food particles. Our successful approach and workflow are readily applicable to other mollusc species.

Evaluating fidelity of CT based 3D models for Zebrafish conductive hearing system.

The zebrafish Weberian apparatus is an emerging model for human conductive hearing system. Their Weberian apparatus comprises minute bones and ligamentary links, and conducts sound pressure transmission from the gas bladder to inner ear through four pairs of Weberian ossicles along the vertebral column. We herein present a methodological study using MicroCT to image the Weberian apparatus for biomechanical and morphological analysis. The aim of this work is to evaluate computational models generated from multiple MicroCT scans with different parameters, to identify the most feasible scan combination for practical (minimized scan time) yet accurate (relative to highest resolution) biomechanical simulations. We segmented and created 3D models from CT scan image stacks at 4.64 µm, 5.05 µm, 9.30 µm and 13.08 µm voxel resolutions, respectively. Then, we used geometric morphometrics analysis to quantify inter-model shape differences, as well as a series of finite element modal and harmonic analyses to simulate auditory signal vibrations. Relative to the highest resolution and most accurate model, the Model 9.30 is closest in overall geometry and biomechanical behavior of all lower resolution models. The differences in resolution and quality of the CT substantially affect the segmentation and reconstruction process of the three-dimensional model of the ossicles, and the subsequent analyses. We conclude that scan voxel resolution is a key factor influencing outcomes of biomechanical simulations of delicate and minute structures, especially when studying the harmonic response of minute ossicles connected by ligaments using finite element modeling. Furthermore, contrast variations in CT images as determined by x-ray power and scan speed, also affect fidelity in 3D models and simulation outcomes.

Broad-scale morpho-functional traits of the mandible suggest no hard food adaptation in the hominin lineage.

An on-going debate concerning the dietary adaptations of archaic hominins and early Homo has been fuelled by contradictory inferences obtained using different methodologies. This work presents an extensive comparative sample of 30 extant primate species that was assembled to perform a morpho-functional comparison of these taxa with 12 models corresponding to eight fossil hominin species. Finite Element Analysis and Geometric Morphometrics were employed to analyse chewing biomechanics and mandible morphology to, firstly, establish the variation of this clade, secondly, relate stress and shape variables, and finally, to classify fossil individuals into broad ingesta related hardness categories using a support vector machine algorithm. Our results suggest that some hominins previously assigned as hard food consumers (e.g. the members of the Paranthropus clade) in fact seem to rely more strongly on soft foods, which is consistent with most recent studies using either microwear or stable isotope analyses. By analysing morphometric and stress results in the context of the comparative framework, we conclude that in the hominin clade there were probably no hard-food specialists. Nonetheless, the biomechanical ability to comminute harder items, if required as fallback option, adds to their strategy of increased flexibility.

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

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.

Cranial suture biomechanics in Metoposaurus krasiejowensis (Temnospondyli, Stereospondyli) from the upper Triassic of Poland.

Cranial sutures connect adjacent bones of the skull and play an important role in the absorption of stresses that may occur during different activities. The Late Triassic temnospondyl amphibian Metoposaurus krasiejowensis has been extensively studied over the years in terms of skull biomechanics, but without a detailed description of the function of cranial sutures. In the present study, 34 thin sections of cranial sutures were examined in order to determine their histovariability and interpret their biomechanical role in the skull. The histological model was compared with three-dimensional-finite element analysis (FEA) simulations of the skull under bilateral and lateral biting as well as skull-raising loads for maximum and minimum principal stress. Histologically, only two sutural morphologies were recognised in the skull of Metoposaurus: interdigitated sutures (commonly associated with compressive stresses) are dominant along the entire length of the skull roof and palate; tongue-and-groove sutures (commonly associated with tensile stresses) are present across the maxilla. FEA shows a much more complex picture of stress type and distribution than predicted by sutures. Common to both methods is a predominance of compressive stresses which act on the skull during biting. The methods predict different stress regimes during biting in the posterior part of the skull: where histological analysis suggests compression, FEA predicts tension. For lateral biting and skull raising, histological and digital reconstructions show similar general patterns but with some variations.

Why ruminating ungulates chew sloppily: Biomechanics discern a phylogenetic pattern.

There is considerable debate regarding whether mandibular morphology in ungulates primarily reflects phylogenetic affinities or adaptation to specific diet. In an effort to help resolve this debate, we use three-dimensional finite element analysis (FEA) to assess the biomechanical performance of mandibles in eleven ungulate taxa with well-established but distinct dietary preferences. We found notable differences in the magnitude and the distribution of von Mises stress between Artiodactyla and Perissodactyla, with the latter displaying lower overall stress values. Additionally, within the order Artiodactyla the suborders Ruminantia and Tylopoda showed further distinctive stress patterns. Our data suggest that a strong phylogenetic signal can be detected in biomechanical performance of the ungulate mandible. In general, Perissodactyla have stiffer mandibles than Artiodactyla. This difference is more evident between Perissodactyla and ruminant species. Perissodactyla likely rely more heavily on thoroughly chewing their food upon initial ingestion, which demands higher bite forces and greater stress resistance, while ruminants shift comminution to a later state (rumination) where less mechanical effort is required by the jaw to obtain sufficient disintegration. We therefore suggest that ruminants can afford to chew sloppily regardless of ingesta, while hindgut fermenters cannot. Additionally, our data support a secondary degree of adaptation towards specific diet. We find that mandibular morphologies reflect the masticatory demands of specific ingesta within the orders Artiodactyla and Perissodactyla. Of particular note, stress patterns in the white rhinoceros (C. simum) look more like those of a general grazer than like other rhinoceros' taxa. Similarly, the camelids (Tylopoda) appear to occupy an intermediate position in the stress patterns, which reflects the more ancestral ruminating system of the Tylopoda.

Inferring locomotor behaviours in Miocene New World monkeys using finite element analysis, geometric morphometrics and machine-learning classification techniques applied to talar morphology.

The talus is one of the most commonly preserved post-cranial elements in the platyrrhine fossil record. Talar morphology can provide information about postural adaptations because it is the anatomical structure responsible for transmitting body mass forces from the leg to the foot. The aim of this study is to test whether the locomotor behaviour of fossil Miocene platyrrhines could be inferred from their talus morphology. The extant sample was classified into three different locomotor categories and then talar strength was compared using finite-element analysis. Geometric morphometrics were used to quantify talar shape and to assess its association with biomechanical strength. Finally, several machine-learning (ML) algorithms were trained using both the biomechanical and morphometric data from the extant taxa to infer the possible locomotor behaviour of the Miocene fossil sample. The obtained results show that the different locomotor categories are distinguishable using either biomechanical or morphometric data. The ML algorithms categorized most of the fossil sample as arboreal quadrupeds. This study has shown that a combined approach can contribute to the understanding of platyrrhine talar morphology and its relationship with locomotion. This approach is likely to be beneficial for determining the locomotor habits in other fossil taxa.

Analyzing the sclerocarpy adaptations of the Pitheciidae mandible.

Primates are interpreted to be ancestrally adapted to frugivory, although some modern groups show clear adaptations to other diets. Among them, pitheciids stand out for specifically predating seeds. This dietary specialization is known as sclerocarpy and refers to the extraction of seeds from surrounding hard tissues using the anterior dentition followed by the mastication of seeds by the molars. It has been proposed that Callicebus-Pithecia-Chiropotes-Cacajao represent a morphocline of increasingly specialized anatomical traits for sclerocarpic foraging. This study addresses whether there is a sclerocarpic specialization gradient in the mandibular morphology of pitheciids. Finite element analysis (FEA) was used to simulate two biting scenarios and the obtained stress values were compared between different pitheciids. Geometric morphometrics (GM) were used to display the morphological variation of this group. No support was found for the morphocline hypothesis from a biomechanical viewpoint since all pitheciins showed similar stress values and on average Chiropotes rather than Cacajao exhibited the strongest mandible. From a morphological perspective, it was found that there is indeed relative "robusticity" continuum in the pitheciid mandible for some aspects of shape as expected for the morphocline hypothesis, but this gradient could be related to other factors rather than sclerocarpic specialization. The present results are expected to contribute to a better insight regarding the ecomorphological relationship between mandibular morphology and mechanical performance among pitheciids.

Merging cranial histology and 3D-computational biomechanics: a review of the feeding ecology of a Late Triassic temnospondyl amphibian.

Finite Element Analysis (FEA) is a useful method for understanding form and function. However, modelling of fossil taxa invariably involves assumptions as a result of preservation-induced loss of information in the fossil record. To test the validity of predictions from FEA, given such assumptions, these results could be compared to independent lines of evidence for cranial mechanics. In the present study a new concept of using bone microstructure to predict stress distribution in the skull during feeding is put forward and a correlation between bone microstructure and results of computational biomechanics (FEA) is carried out. The bony framework is a product of biological optimisation; bone structure is created to meet local mechanical conditions. To test how well results from FEA correlate to cranial mechanics predicted from bone structure, the well-known temnospondyl Metoposaurus krasiejowensis was used as a model. A crucial issue to Temnospondyli is their feeding mode: did they suction feed or employ direct biting, or both? Metoposaurids have previously been characterised either as active hunters or passive bottom dwellers. In order to test the correlation between results from FEA and bone microstructure, two skulls of Metoposaurus were used, one modelled under FE analyses, while for the second one 17 dermal bone microstructure were analysed. Thus, for the first time, results predicting cranial mechanical behaviour using both methods are merged to understand the feeding strategy of Metoposaurus. Metoposaurus appears to have been an aquatic animal that exhibited a generalist feeding behaviour. This taxon may have used two foraging techniques in hunting; mainly bilateral biting and, to a lesser extent, lateral strikes. However, bone microstructure suggests that lateral biting was more frequent than suggested by Finite Element Analysis (FEA). One of the potential factors that determined its mode of life may have been water levels. During optimum water conditions, metoposaurids may have been more active ambush predators that were capable of lateral strikes of the head. The dry season required a less active mode of life when bilateral biting is particularly efficient. This, combined with their characteristically anteriorly positioned orbits, was optimal for ambush strategy. This ability to use alternative modes of food acquisition, independent of environmental conditions, might hold the key in explaining the very common occurrence of metoposaurids during the Late Triassic.

The intervals method: a new approach to analyse finite element outputs using multivariate statistics.

BACKGROUND: In this paper, we propose a new method, named the intervals' method, to analyse data from finite element models in a comparative multivariate framework. As a case study, several armadillo mandibles are analysed, showing that the proposed method is useful to distinguish and characterise biomechanical differences related to diet/ecomorphology. METHODS: The intervals' method consists of generating a set of variables, each one defined by an interval of stress values. Each variable is expressed as a percentage of the area of the mandible occupied by those stress values. Afterwards these newly generated variables can be analysed using multivariate methods. RESULTS: Applying this novel method to the biological case study of whether armadillo mandibles differ according to dietary groups, we show that the intervals' method is a powerful tool to characterize biomechanical performance and how this relates to different diets. This allows us to positively discriminate between specialist and generalist species. DISCUSSION: We show that the proposed approach is a useful methodology not affected by the characteristics of the finite element mesh. Additionally, the positive discriminating results obtained when analysing a difficult case study suggest that the proposed method could be a very useful tool for comparative studies in finite element analysis using multivariate statistical approaches.

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, affect the feeding capabilities of these animals. How changes in cranial morphology and tissue properties affect 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 different tissue properties and ossification 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 stiffness required to perform a function versus the fixation (and displacement) required with the surrounding skeletal elements. (b) Changes on the ossification pattern, producing fontanelles as a result of bone loss or failure to ossify, represent a trend toward simplification 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.

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.

Feeding biomechanics of Late Triassic metoposaurids (Amphibia: Temnospondyli): a 3D finite element analysis approach.

The Late Triassic freshwater ecosystems were occupied by different tetrapod groups including large-sized anamniotes, such as metoposaurids. Most members of this group of temnospondyls acquired gigantic sizes (up to 5 m long) with a nearly worldwide distribution. The paleoecology of metoposaurids is controversial; they have been historically considered passive, bottom-dwelling animals, waiting for prey on the bottom of rivers and lakes, or they have been suggested to be active mid-water feeders. The present study aims to expand upon the paleoecological interpretations of these animals using 3D finite element analyses (FEA). Skulls from two taxa, Metoposaurus krasiejowensis, a gigantic taxon from Europe, and Apachesaurus gregorii, a non-gigantic taxon from North America, were analyzed under different biomechanical scenarios. Both 3D models of the skulls were scaled to allow comparisons between them and reveal that the general stress distribution pattern found in both taxa is clearly similar in all scenarios. In light of our results, both previous hypotheses about the paleoecology of these animals can be partly merged: metoposaurids probably were ambush and active predators, but not the top predators of these aquatic environments. The FEA results demonstrate that they were particularly efficient at bilateral biting, and together with their characteristically anteropositioned orbits, optimal for an ambush strategy. Nonetheless, the results also show that these animals were capable of lateral strikes of the head, suggesting active hunting of prey. Regarding the important skull size differences between the taxa analyzed, our results suggest that the size reduction in the North American taxon could be related to drastic environmental changes or the increase of competitors. The size reduction might have helped them expand into new ecological niches, but they likely remained fully aquatic, as are all other metoposaurids.

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.

3D computational mechanics elucidate the evolutionary implications of orbit position and size diversity of early amphibians.

For the first time in vertebrate palaeontology, the potential of joining Finite Element Analysis (FEA) and Parametrical Analysis (PA) is used to shed new light on two different cranial parameters from the orbits to evaluate their biomechanical role and evolutionary patterns. The early tetrapod group of Stereospondyls, one of the largest groups of Temnospondyls is used as a case study because its orbits position and size vary hugely within the members of this group. An adult skull of Edingerella madagascariensis was analysed using two different cases of boundary and loading conditions in order to quantify stress and deformation response under a bilateral bite and during skull raising. Firstly, the variation of the original geometry of its orbits was introduced in the models producing new FEA results, allowing the exploration of the ecomorphology, feeding strategy and evolutionary patterns of these top predators. Secondly, the quantitative results were analysed in order to check if the orbit size and position were correlated with different stress patterns. These results revealed that in most of the cases the stress distribution is not affected by changes in the size and position of the orbit. This finding supports the high mechanical plasticity of this group during the Triassic period. The absence of mechanical constraints regarding the orbit probably promoted the ecomorphological diversity acknowledged for this group, as well as its ecological niche differentiation in the terrestrial Triassic ecosystems in clades as lydekkerinids, trematosaurs, capitosaurs or metoposaurs.

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 their "conservative" morphology, cryptobranchids may be useful models to reconstruct the feeding ecology and biomechanics of different members of early tetrapods and amphibians, with similar osteological and myological constraints.

Finite Element Analysis of the Cingulata Jaw: An Ecomorphological Approach to Armadillo's Diets.

Finite element analyses (FEA) were applied to assess the lower jaw biomechanics of cingulate xenarthrans: 14 species of armadillos as well as one Pleistocene pampathere (11 extant taxa and the extinct forms Vassallia, Eutatus and Macroeuphractus). The principal goal of this work is to comparatively assess the biomechanical capabilities of the mandible based on FEA and to relate the obtained stress patterns with diet preferences and variability, in extant and extinct species through an ecomorphology approach. The results of FEA showed that omnivorous species have stronger mandibles than insectivorous species. Moreover, this latter group of species showed high variability, including some similar biomechanical features of the insectivorous Tolypeutes matacus and Chlamyphorus truncatus to those of omnivorous species, in agreement with reported diets that include items other than insects. It remains unclear the reasons behind the stronger than expected lower jaw of Dasypus kappleri. On the other hand, the very strong mandible of the fossil taxon Vassallia maxima agrees well with the proposed herbivorous diet. Moreover, Eutatus seguini yielded a stress pattern similar to Vassalia in the posterior part of the lower jaw, but resembling that of the stoutly built Macroeuphractus outesi in the anterior part. The results highlight the need for more detailed studies on the natural history of extant armadillos. FEA proved a powerful tool for biomechanical studies in a comparative framework.