Emily Rayfield

ABSTRACT The invasion of the land by vertebrates is a key moment in the history of life and necessitated dramatic skeletal evolution. Gradual changes in lower jaw morphology across the fish–tetrapod transition were initially ascribed to a stronger lower jaw and increased bite force, shifts in feeding strategy (suction vs. biting), or different constraints imposed by contrasting environments (aquatic vs. terrestrial). However, more recent work has suggested a disjunction between morphology and function in the lower jaws of early tetrapods. This study, which incorporates early tetrapod taxa from the Late Devonian to Early Triassic, investigates the mechanical behaviour of the lower jaw under feeding loads. CT scans were segmented to remove matrix, separate individual bones, sutures, and teeth, and retrodeformed to produce 3D finite element models of the lower jaws of Eusthenopteron, Acanthostega, Greererpeton, Crassigyrinus, Megalocephalus, and Lydekkerina. Models were scaled appropriately and simplified loads and constraints applied. Stress patterns and deformation regimes were compared to finite element models of the lower jaws of extant fish, including pike (Esox) and eel (Anguilla), and to Alligator. Deformation of the lower jaw (including eversion of the working-side tooth row and dorsal bending of the balancing-side jaw ramus) and the distribution of tensile and compressive stresses were similar across most taxa. However, higher stress occurred in the balancing-side ramus of more derived tetrapods, such as Lydekkerina, and in Alligator; in contrast, the working-side ramus of fish mandibles exhibited much higher stresses than the balancing-side ramus. These differences may reflect more efficient transfer of balancing-side muscle forces across the midline due to increasingly complex symphyseal morphology in later tetrapods compared to more basal forms, such as Eusthenopteron. Alternatively, lower stresses in the working-side ramus of later tetrapods may support the suggestion that morphological changes across the fish-tetrapod transition produced a stronger lower jaw better adapted for biting.

Rodents are defined by a uniquely specialized dentition and a highly complex arrangement of jaw-closing muscles. Finite element analysis (FEA) is an ideal technique to investigate the biomechanical implications of these specializations, but it is essential to understand fully the degree of influence of the different input parameters of the FE model to have confidence in the model’s predictions. This study evaluates the sensitivity of FE models of rodent crania to elastic properties of the materials, loading direction, and the location and orientation of the models’ constraints. Three FE models were constructed of squirrel, guinea pig and rat skulls. Each was loaded to simulate biting on the incisors, and the first and the third molars, with the angle of the incisal bite varied over a range of 45 . The Young’s moduli of the bone and teeth components were varied between limits defined by findings from our own and previously published tests of material properties. Geometric morphometri...

The invasion of the land by vertebrates is a key moment in the history of life and necessitated dramatic skeletal evolution. Gradual changes in lower jaw morphology across the fish–tetrapod transition were initially ascribed to a stronger lower jaw and increased bite force, shifts in feeding strategy (suction vs. biting), or different constraints imposed by contrasting environments (aquatic vs. terrestrial). However, more recent work has suggested a disjunction between morphology and function in the lower jaws of early tetrapods. This study, which incorporates early tetrapod taxa from the Late Devonian to Early Triassic, investigates the mechanical behaviour of the lower jaw under feeding loads. CT scans were segmented to remove matrix, separate individual bones, sutures, and teeth, and retrodeformed to produce 3D finite element models of the lower jaws of Eusthenopteron, Acanthostega, Greererpeton, Crassigyrinus, Megalocephalus, and Lydekkerina. Models were scaled appropriately and...

Pliosaurs were among the largest predators in Mesozoic seas, and yet their functional anatomy and feeding biomechanics are poorly understood. A new, well-preserved pliosaur from the Kimmeridgian of Weymouth Bay (UK) revealed cranial adaptations related to feeding. Digital modelling of computed tomography scans allowed reconstruction of missing, distorted regions of the skull and of the adductor musculature, which indicated high bite forces. Size-corrected beam theory modelling showed that the snout was poorly optimised against bending and torsional stresses compared with other aquatic and terrestrial predators, suggesting that pliosaurs did not twist or shake their prey during feeding and that seizing was better performed with post-symphyseal bites. Finite element analysis identified biting-induced stress patterns in both the rostrum and lower jaws, highlighting weak areas in the rostral maxillary-premaxillary contact and the caudal mandibular symphysis. A comparatively weak skull c...

Finite element modelling is well entrenched in comparative vertebrate biomechanics as a tool to assess the mechanical design of skeletal structures and to better comprehend the complex interaction of their form-function relationships. But what makes a reliable subject-specific finite element model? To approach this question, we here present a set of convergence and sensitivity analyses and a validation study as an example, for finite element analysis (FEA) in general, of ways to ensure a reliable model. We detail how choices of element size, type and material properties in FEA influence the results of simulations. We also present an empirical model for estimating heterogeneous material properties throughout an elephant femur (but of broad applicability to FEA). We then use an ex vivo experimental validation test of a cadaveric femur to check our FEA results and find that the heterogeneous model matches the experimental results extremely well, and far better than the homogeneous mode...

Computer-aided visualization and analysis of fossils has
revolutionized the study of extinct organisms. Novel
techniques allow fossils to be characterized in three
dimensions and in unprecedented detail. This has enabled
paleontologists to gain important insights into
their anatomy, development, and preservation. New
protocols allow more objective reconstructions of fossil
organisms, including soft tissues, from incomplete
remains. The resulting digital reconstructions can be
used in functional analyses, rigorously testing longstanding
hypotheses regarding the paleobiology of extinct
organisms. These approaches are transforming our
understanding of long-studied fossil groups, and of the
narratives of organismal and ecological evolution that
have been built upon them.

Maniraptoriformes, the speciose group of derived theropod dinosaurs that ultimately gave rise to modern birds, display a diverse and remarkable suite of skeletal adaptations. Apart from the evolution of flight, a large-scale change in dietary behavior appears to have been one of the main triggers for specializations in the bauplan of these derived theropods. Among the different skeletal specializations, partial or even complete edentulism and the development of keratinous beaks form a recurring and persistent trend in from the evolution of derived nonavian dinosaurs. Therizinosauria is an enigmatic maniraptoriform clade, whose members display these and other osteological characters thought to be correlated with the shift from carnivory to herbivory. This makes therizinosaurians prime candidates to assess the functional significance of these morphological characters. Based on a highly detailed biomechanical model of Erlikosaurus andrewsi, a therizinosaurid from the Upper Cretaceous of Mongolia, different morphological configurations incorporating soft-tissue structures, such as a keratinous rhamphotheca, are evaluated for their biomechanical performance. Our results indicate that the development of beaks and the presence of a keratinous rhamphotheca would have helped to dissipate stress and strain, making the rostral part of the skull less susceptible to bending and displacement, and this benefit may extend to other vertebrate clades that possess rhamphothecae. Keratinous beaks, paralleled by edentulism, thus represent an evolutionary innovation developed early in derived theropods to enhance cranial stability, distinct to postulated mass-saving benefits associated with the origin of flight.

High megaherbivore species richness is documented in both fossil and contemporary ecosystems despite their high individual energy requirements. An extreme example of this is the Late Jurassic Morrison Formation, which was dominated by sauropod dinosaurs, the largest known terrestrial vertebrates. High sauropod diversity within the resource-limited Morrison is paradoxical, but might be explicable through sophisticated resource partitioning. This hypothesis was tested through finite-element analysis of the crania of the Morrison taxa Camarasaurus and Diplodocus. Results demonstrate divergent specialization, with Camarasaurus capable of exerting and accommodating greater bite forces than Diplodocus, permitting consumption of harder food items. Analysis of craniodental biomechanical characters taken from 35 sauropod taxa demonstrates a functional dichotomy in terms of bite force, cranial robustness and occlusal relationships yielding two polyphyletic functional ‘grades’. Morrison taxa are widely distributed within and between these two morphotypes, reflecting distinctive foraging specializations that formed a biomechanical basis for niche partitioning between them. This partitioning, coupled with benefits associated with large body size, would have enabled the high sauropod diversities present in the Morrison Formation. Further, this provides insight into the mechanisms responsible for supporting the high diversities of large megaherbivores observed in other Mesozoic and Cenozoic communities, particularly those occurring in resource-limited environments.

Making the link between evolutionary processes and development in extinct organisms is usually hampered by the lack of preservation of ontogenetic stages in the fossil record. Planktic foraminifers, which grow by adding chambers, are an ideal target organism for such studies as their test incorporates all prior developmental stages. Previously, studies of development in these organisms were limited by the small size of their early chambers. Here, we describe the application of synchrotron radiation X-ray tomographic microscopy (SRXTM) to document the ontogenetic history of the foraminifers Globigerinoides sacculifer and Globorotalia menardii. Our SRXTM scans permit resolution at submicrometre scale, thereby displaying additional internal structures such as pores, dissolution patterns and complexity of the wall growth. Our methods provide a powerful tool to pick apart the  developmental history of these microfossils and subsequently assist in inferring phylogenetic relationships and evolutionary processes.

A number of extant and extinct archosaurs evolved an elongate, narrow rostrum. This longirostrine condition has been
associated with a diet comprising a higher proportion of fish and smaller prey items compared to taxa with broader, more
robust snouts. The evolution of longirostrine morphology and a bulbous anterior rosette of premaxillary teeth also occurs in
the spinosaurid theropod dinosaurs, leading to suggestions that at least some members of this clade also had a diet
comprising a notable proportion of fish or other small vertebrates. Here we compare the rostral biomechanics of the
spinosaurs Baryonyx walkeri and Spinosaurus c.f. S. aegyptiacus to three extant crocodilians: two longistrine taxa, the African slender-snouted crocodile Mecistops cataphractus and the Indian gharial Gavialis gangeticus; and the American alligator
Alligator mississippiensis. Using computed tomography (CT) data, the second moments of area and moments of inertia at
successive transverse slices along the rostrum were calculated for each of the species. Size-independent results tested the
biomechanical benefits of material distribution within the rostra. The two spinosaur rostra were both digitally reconstructed from CT data and compared against all three crocodilians. Results show that African slender-snouted crocodile skulls are
more resistant to bending than an equivalent sized gharial. The alligator has the highest resistances to bending and torsion
of the crocodiles for its size and greater than that of the spinosaurs. The spinosaur rostra possess similar resistance to
bending and torsion despite their different morphologies. When size is accounted for, B. walkeri performs mechanically
differently from the gharial, contradicting previous studies whereas Spinosaurus does not. Biomechanical data support
known feeding ecology for both African slender-snouted crocodile and alligator, and suggest that the spinosaurs were not obligate piscivores with diet being determined by individual animal size.

Conodont elements are the earliest vertebrate dental structures. The dental tools on elements responsible for food fracture—cusps and denticles—are usually composed of lamellar crown tissue (a putative enamel homologue) and the enigmatic tissue known as ‘white matter’. White matter is unique to conodonts and has been hypothesized to be a functional adaptation for the use of elements as teeth. We test this quantitatively using finite-element analysis. Our results indicate that white matter allowed cusps and denticles to withstand greater tensile stresses than do cusps comprised solely of lamellar crown tissue. Microstructural variation is demonstrably associated with dietary and loading differences in teeth, so secondary loss of white matter through conodont phylogeny may reflect changes in diet and element occlusal kinematics. The presence, development and distribution of white matter could thus provide constraints on function in the first vertebrate dental structures.

Background Therizinosauria is one of the most enigmatic and peculiar clades among theropod dinosaurs, exhibiting an unusual suite of characters, such as lanceolate teeth, a rostral rhamphotheca, long manual claws, and a wide, opisthopubic pelvis. This specialized anatomy has been associated with a shift in dietary preferences and an adaptation to herbivory. Despite a large number of discoveries in recent years, the fossil record for Therizinosauria is still relatively poor, and cranial remains are particularly rare.

Pedal claw geometry can be used to predict behaviour in extant tetrapods and has frequently been used as an indicator of lifestyle and ecology in Mesozoic birds and other fossil reptiles, sometimes without acknowledgement of the caveat that data from other aspects of morphology and proportions also need to be considered. Variation in styles of measurement (both inner and outer claw curvature angles) has made it difficult to compare results across studies, as have over-simplified ecological categories.

Computational models such as finite-element analysis offer biologists a means of exploring the structural mechanics of biological systems that cannot be directly observed. Validated against experimental data, a model can be manipulated to perform virtual experiments, testing variables that are hard to control in physical experiments. The relationship between tooth form and the ability to break down prey is key to understanding the evolution of dentition. Recent experimental work has quantified how tooth shape promotes fracture in biological materials. We present a validated finite-element model derived from physical compression experiments. The model shows close agreement with strain patterns observed in photoelastic test materials and reaction forces measured during these experiments. We use the model to measure strain energy within the test material when different tooth shapes are used. Results show that notched blades deform materials for less strain energy cost than straight blades, giving insights into the energetic relationship between tooth form and prey materials. We identify a hypothetical 'optimal' blade angle that minimizes strain energy costs and test alternative prey materials via virtual experiments. Using experimental data and computational models offers an integrative approach to understand the mechanics of tooth morphology.

Sauropod dinosaurs were the largest terrestrial herbivores and pushed at the limits of vertebrate biomechanics and physiology. Sauropods exhibit high craniodental diversity in ecosystems where numerous species coexisted, leading to the hypothesis that this biodiversity is linked to niche subdivision driven by ecological specialisation.mHere, we quantitatively investigate feeding behaviour hypotheses for the iconic sauropod Diplodocus. Biomechanical modelling, using finite element analysis, was used to examine the performance of the Diplodocus skull. Three feeding behaviours were modelled: muscle-driven static biting, branch stripping and bark stripping. The skull was found to be ‘over engineered’ for static biting, overall experiencing lowstress with only the dentition enduring high stress. When branch stripping, the skull, similarly, is under low stress, with little appreciable difference between those models. When simulated for bark stripping, the skull experiences far greater stresses, especially in the teeth and at the jaw joint. Therefore, we refute the bark-stripping hypothesis, while the hypotheses of branch stripping and/or precision biting are both consistent with our findings, showing that branch stripping is a biomechanically plausible feeding behaviour for diplodocids. Interestingly, in all simulations, peak stress is observed in the premaxillary–maxillary ‘lateral plates’, supporting the hypothesis that these structures evolved to dissipate stress induced while feeding. These results lead us to conclude that the aberrant craniodental form of Diplodocus was adapted for food procurement
rather than resisting high bite forces.

Conodonts have been considered the earliest skeletonizing vertebrates and their mineralized feeding apparatus interpreted as having performed a tooth function. However, the absence of jaws in conodonts and the small size of their oropharyngeal musculature limits the force available for fracturing food items, presenting a challenge to this interpretation. We address this issue quantitatively using engineering approaches previously applied to mammalian dentitions. We show that the morphology of conodont food-processing elements was adapted to overcome size limitations through developing dental tools of unparalleled sharpness that maximize applied pressure. Combined with observations of wear, we also show how this morphology was employed, demonstrating how Wurmiella excavata used rotational kinematics similar to other conodonts, suggesting that this occlusal style is typical for the clade. Our work places conodont elements within a broader dental framework, providing a phylogenetically independent system for examining convergence and scaling in dental tools.

The furcula displays enormous morphological and structural diversity. Acting as an important origin for flight muscles involved in the downstroke, the form of this element has been shown to vary with flight mode. This study seeks to clarify the strength of this form-function relationship through the use of eigenshape morphometric analysis coupled with recently developed phylogenetic comparative methods (PCMs), including phylogenetic Flexible Discriminant Analysis (pFDA). Additionally, the morphospace derived from the furculae of extant birds is used to shed light on possible flight adaptations of Mesozoic fossil taxa. While broad conclusions of earlier work are supported (U-shaped furculae are associated with soaring, strong anteroposterior curvature with wing-propelled diving), correlations between form and function do not appear to be so clear-cut, likely due to the significantly larger dataset and wider spectrum of flight modes sampled here. Interclavicular angle is an even more powerful discriminator of flight mode than curvature, and is positively correlated with body size. With the exception of the close relatives of modern birds, the ornithuromorphs, Mesozoic taxa tend to occupy unique regions of morphospace, and thus may have either evolved unfamiliar flight styles or have arrived at similar styles through divergent musculoskeletal configurations.

The masticatory musculature of rodents has evolved to enable both gnawing at the incisors and chewing at the molars. In particular, the masseter muscle is highly specialised, having extended anteriorly to originate from the rostrum. All living rodents have achieved this masseteric expansion in one of three ways, known as the sciuromorph, hystricomorph and myomorph conditions. Here, we used finite element analysis (FEA) to investigate the biomechanical implications of these three morphologies, in a squirrel, guinea pig and rat. In particular, we wished to determine whether each of the three morphologies is better adapted for either gnawing or chewing. Results show that squirrels are more efficient at muscle-bite force transmission during incisor gnawing than guinea pigs, and that guinea pigs are more efficient at molar chewing than squirrels. This matches the known diet of nuts and seeds that squirrels gnaw, and of grasses that guinea pigs grind down with their molars. Surprisingly, results also indicate that rats are more efficient as well as more versatile feeders than both the squirrel and guinea pig. There seems to be no compromise in biting efficiency to accommodate the wider range of foodstuffs and the more general feeding behaviour adopted by rats. Our results show that the morphology of the skull and masticatory muscles have allowed squirrels to specialise as gnawers and guinea pigs as chewers, but that rats are high-performance generalists, which helps explain their overwhelming success as a group.

Models are a principal tool of modern science. By definition, and in practice, models are not literal representations of reality but provide simplifications or substitutes of the events, scenarios or behaviours that are being studied or predicted. All models make assumptions, and palaeontological models in particular require additional assumptions to study unobservable events in deep time. In the case of functional analysis, the degree of missing data associated with reconstructing musculoskeletal anatomy and neuronal control in extinct organisms has, in the eyes of some scientists, rendered detailed functional analysis of fossils intractable. Such a prognosis may indeed be realized if palaeontologists attempt to recreate elaborate biomechanical models based on missing data and loosely justified assumptions. Yet multiple enabling methodologies and techniques now exist: tools for bracketing boundaries of reality; more rigorous consideration of soft tissues and missing data and methods drawing on physical principles that all organisms must adhere to. As with many aspects of science, the utility of such biomechanical models depends on the questions they seek to address, and the accuracy and validity of the models themselves.

More than 99 per cent of the roughly 58,000 living vertebrate species have jaws1. This major clade, whose members are collectively known as gnathostomes (‘jawed mouths’), made its earliest definitive appearance in the Silurian period, 444–416 million years (Myr) ago, with both the origin of the modern (crown-group) radiation and the presumptive invasion of land occurring by the end of the Devonian period2 (359 Myr ago). These events coincided with a major faunal shift that remains apparent today: the transition from Silurian ecosystems dominated by jawless fishes (agnathans) to younger assemblages composed almost exclusively of gnathostomes2, 3. This pattern has inspired several qualitative descriptions of the trophic radiation and ecological ascendance of the earliest jawed vertebrates3, 4, 5, 6, 7. Here we present a quantitative analysis of functional variation in early gnathostome mandibular elements, placing constraints on our understanding of evolutionary patterns during this critical interval. We document an initial increase in functional disparity in the Silurian that stabilized by the first stage of the Devonian, before the occurrence of an Emsian (~400 Myr ago) oxygenation event implicated in the trophic radiation of vertebrates8. Subsequent taxonomic diversification during the Devonian did not result in increased functional variation; instead, new taxa revisited and elaborated on established mandibular designs. Devonian functional space is dominated by lobe-finned fishes and ‘placoderms’; high disparity within the latter implies considerable trophic innovation among jaw-bearing stem gnathostomes. By contrast, the major groups of living vertebrates—ray-finned fishes and tetrapods—show surprisingly conservative mandibular morphologies with little indication of functional diversification or innovation. Devonian gnathostomes reached a point where they ceased to accrue further mandibular functional disparity before becoming taxonomic dominants relative to ‘ostracoderm’-grade jawless fishes, providing a new perspective on classic adaptive hypotheses concerning this fundamental shift in vertebrate biodiversity.

Rodents are defined by a uniquely specialized dentition and a highly complex arrangement of jaw-closing muscles. Finite element analysis (FEA) is an ideal technique to investigate the biomechanical implications of these specializations, but it is essential to understand fully the degree of influence of the different input parameters of the FE model to have confidence in the model's predictions. This study evaluates the sensitivity of FE models of rodent crania to elastic properties of the materials, loading direction, and the location and orientation of the models' constraints. Three FE models were constructed of squirrel, guinea pig and rat skulls. Each was loaded to simulate biting on the incisors, and the first and the third molars, with the angle of the incisal bite varied over a range of 45 degrees. The Young's moduli of the bone and teeth components were varied between limits defined by findings from our own and previously published tests of material properties. Geometric morphometrics (GMM) was used to analyse the resulting skull deformations. Bone stiffness was found to have the strongest influence on the results in all three rodents, followed by bite position, and then bite angle and muscle orientation. Tooth material properties were shown to have little effect on the deformation of the skull. The effect of bite position varied between species, with the mesiodistal position of the biting tooth being most important in squirrels and guinea pigs, whereas bilateral vs. unilateral biting had the greatest influence in rats. A GMM analysis of isolated incisor deformations showed that, for all rodents, bite angle is the most important parameter, followed by elastic properties of the tooth. The results here elucidate which input parameters are most important when defining the FE models, but also provide interesting glimpses of the biomechanical differences between the three skulls, which will be fully explored in future publications.

How theropod dinosaur skulls experience stress and strain during the application of adductor muscle loads provides a unique insight into their feeding behaviour and principles of skeletal construction and scaling. Of particular interest are unusual cranial morphologies, such as those seen in the spinosaurid theropods, Baryonyx walkeri, Suchomimus tenerensis, Irritator challengeri and Spinosaurus aegyptiacus. This study uses the engineering technique finite element analysis to reconstruct feeding-related stress and strain in the skulls of seven theropod dinosaurs: five non-neotetanurans (Afrovenator, Dubreuillosaurus, Monolophosaurus, Spinosaurus and Suchomimus) and two basal neotetanurans (Acrocanthosaurus, and Carcharodontosaurus). Two-dimensional finite element models are created, and simulated adductor muscle loads are applied in proportion to the lateral surface area of the skull, thereby removing the influence of size and testing the efficiency of shape at resisting relative loads. Results show a significant size-related trend, with large taxa experiencing greater stresses than smaller taxa. Whilst Suchomimus scales with other theropods, Spinosaurus is a notable outlier and experiences much higher magnitudes of cranial stress than would be predicted. It may be that when realistic loading parameters are considered, larger theropods mitigate potential cranial weakness through concomitnat scaling of adductor muscle and bite force or through modifications to feeding ecology, especially in taxa such as Spinosaurus. Give the 2D nature of these models, results and interpretations should be treated with caution, and are best considered predictors of biomechanical performance and feeding ecology, to be tested in the future with more appropriate 3D finite element models.

A finite element (FE) validation and sensitivity study was undertaken on a modern domestic pig cranium. Bone strain data were collected ex vivo from strain gauges, and compared with results from specimen-specific FE models. An isotropic, homogeneous model was created, then input parameters were altered to investigate model sensitivity. Heterogeneous, isotropic models investigated the effects of a constant-thickness, stiffer outer layer (representing cortical bone) atop a more compliant interior (representing cancellous bone). Loading direction and placement of strain gauges were also varied, and the use of 2D membrane elements at strain gauge locations as a method of projecting 3D model strains into the plane of the gauge was investigated. The models correctly estimate the loading conditions of the experiment, yet at some locations fail to reproduce correct principal strain magnitudes, and hence strain ratios. Principal strain orientations are predicted well. The initial model was too stiff by approximately an order of magnitude. Introducing a compliant interior reported strain magnitudes more similar to the ex vivo results without notably affecting strain orientations, ratios or contour patterns, suggesting that this simple heterogeneity was the equivalent of reducing the overall stiffness of the model. Models were generally insensitive to moderate changes in loading direction or strain gauge placement, except in the squamosal portion of the zygomatic arch. The use of membrane elements made negligible differences to the reported strains. The models therefore seem most sensitive to changes in material properties, and suggest that failure to model local heterogeneity in material properties and structure of the bone may be responsible for discrepancies between the experimental and model results. This is partially attributable to a lack of resolution in the CT scans from which the model was built, and partially due to an absence of detailed material properties data for pig cranial bone. Thus, caution is advised when using FE models to estimate absolute numerical values of breaking stress and bite force unless detailed input parameters are available. However, if the objective is to compare relative differences between models, the fact that the strain environment is replicated well means that such investigations can be robust.

Finite element (FE) models provide discrete solutions to continuous problems. Therefore, to arrive at the correct solution, it is vital to ensure that FE models contain a sufficient number of elements to fully resolve all the detail encountered in a continuum structure. Mesh convergence testing is the process of comparing successively finer meshes to identify the point of diminishing returns; where increasing resolution has marginal effects on results and further detail would become costly and unnecessary. Historically, convergence has not been considered in most CT-based biomechanical reconstructions involving complex geometries like the skull, as generating such models has been prohibitively time-consuming. To assess how mesh convergence influences results, 18 increasingly refined CT-based models of a domestic pig skull were compared to identify the point of convergence for strain and displacement, using both linear and quadratic tetrahedral elements. Not all regions of the skull converged at the same rate, and unexpectedly, areas of high strain converged faster than low-strain regions. Linear models were slightly stiffer than their quadratic counterparts, but did not converge less rapidly. As expected, insufficiently dense models underestimated strain and displacement, and failed to resolve strain ‘‘hot-spots’’ notable in contour plots. In addition to quantitative differences, visual assessments of such plots often inform conclusions drawn in many comparative studies, highlighting that mesh convergence should be performed on all finite element models before further analysis takes place.