Cranial Biomechanics Research Articles

Sexual Dimorphism and Divergent Evolutionary Pathways in Primate Cranial Biomechanics: Insights From a Theoretical Morphology Framework.

The mammalian order Primates is known for widespread sexual dimorphism in size and phenotype. Despite repeated speculation that primate sexual size dimorphism either facilitates or is in part driven by functional differences in how males and females interact with their environments, few studies have directly assessed the influence of sexual dimorphism on performance traits. Here, we use a theoretical morphology framework to show that sexual dimorphism in primate crania is associated with divergent biomechanical performance traits. The degree of dimorphism is a significant covariate in biomechanical trait divergence between sexes. Males exhibit less efficient but stiffer cranial shapes and significant evolutionary allometry in biomechanical performance, whereas females maintain performance stability across their size spectrum. Evolutionary rates are elevated for efficiency in females whereas males emphasize size-dependent cranial stiffness. These findings support a hypothesis of sex-linked bifurcation in masticatory system performance: larger male crania and faster size evolution partially compensate for low efficiency and reflect a de-emphasis of mechanical leverage, whereas female crania maintain higher mechanical efficiency overall and evolve more rapidly in molar-based masticatory performance. The evolutionary checks-and-balances between size dimorphism and cranial mechanical performance may be a more important driver of primate phenotypic evolution than has been hitherto appreciated.

Skink systematics inside out: Comparative cranial osteology of the New World Mabuyinae.

The Mabuyinae subfamily exhibits remarkable diversity, encompassing 26 genera and 236 currently recognized species. Traditionally, the entire range of the group was attributed to the single genus Mabuya, which had a wide distribution along tropical regions of the Planet. In recent studies, phylogenetic hypotheses based on molecular data have identified four major groups, which have been further divided into geographically distinct clades. At least two phylogenetically distinct lineages of Mabuyinae are distributed in the Neotropical Region: Trachylepis atlantica and the remaining 16 genera within the Mabuyinae clade from the mainland and the Caribbean islands. Our understanding of Mabuyinae osteology is still quite limited, particularly concerning interspecific variation. This lack of information hinders our ability to make strong contributions to the phylogenetic relationships within this group or even to confirm the existence of certain new taxa considering their relatively conserved external morphology. This work provides a comprehensive anatomical reference for the adult skull of Neotropical Mabuyinae lizards, highlighting osteological features that might be useful for delimiting each genus. This descriptive guide includes illustrations and employs multiple techniques, such as dry preparation, clearing and staining, and high-resolution computerized microtomography. Our results provide additional diagnostic characteristics that include specific cranial bone arrangements, dental patterns, and cranial adaptations, such as dorsoventral head flattening, and their functional implications for bite force and cranial biomechanics. This study reinforces the importance of cranial morphology in understanding the phylogenetic relationships and evolutionary trajectories of New World Mabuyinae lizards, advocating for broader morphological sampling to enrich our understanding of these diverse reptiles.

Beyond CREA: Evolutionary patterns of non-allometric shape variation and divergence in a highly allometric clade of murine rodents.

The shared functions of the skull are thought to result in common evolutionary patterns in mammalian cranial shape. Craniofacial evolutionary allometry (CREA) is a particularly prominent pattern where larger species display proportionally elongate facial skeletons and smaller braincases. It was recently proposed that CREA arises from biomechanical effects of cranial scaling when diets are similar. Thus, deviations from CREA should occur with changes in cranial biomechanics, for example due to dietary change. Here, we test this using 3D geometric morphometric analysis in a dataset of Australian murine crania, which are highly allometric. We contrast allometric and non-allometric variation in the cranium by comparing evolutionary mode, allometry, ordinations, as well as allometry, integration, and modularity in functional modules. We found evidence of stabilising selection in allometry-containing and size-free shape, and substantial non-allometric variation aligned with dietary specialisation in parallel with CREA. Integration among cranial modules was higher, and modularity lower, with size included, but integration between rostrum and cranial vault, which are involved in the CREA pattern, dropped dramatically after size removal. Our results thus support the hypothesis that CREA is a composite arising from selection on cranial function, with substantial non-allometric shape variation occurring alongside CREA where dietary specialisation impacts selection on gnawing function. This emphasises the need to research mammalian cranial evolution in the context of allometric and non-allometric selection on biomechanical function.

Comparing cranial biomechanics between Barbourofelis fricki and Smilodon fatalis: Is there a universal killing-bite among saber-toothed predators?

Saber-tooths, extinct apex predators with long and blade-like upper canines, have appeared iteratively at least five times in the evolutionary history of vertebrates. Although saber-tooths exhibit a relatively diverse range of morphologies, it is widely accepted that all killed their prey using the same predatory behavior. In this study, we CT-scanned the skull of Barbourofelis fricki and compared its cranial mechanics using finite element analysis (FEA) with that of Smilodon fatalis. Our aim was to investigate potential variations in killing behavior between two dirk-toothed sabretooths from the Miocene and Pleistocene of North America. The study revealed that B. fricki had a stoutly-built skull capable of withstanding stress in various prey-killing scenarios, while the skull of S. fatalis appeared less optimized for supporting stress, which highlights the highly derived saber-tooth morphology of the former. The results may indicate that B. fricki was more of a generalist in prey-killing compared to S. fatalis, which experiences lower stresses under stabbing loads. We hypothesize that morphological specialization in saber-tooths does not necessarily indicate ecological specialization. Our results support the notion that morphological convergence among saber-toothed cats may obscure differences in hunting strategies employed to dispatch their prey. Our findings challenge the assumption of the universally assumed canine-shear biting as the prey-killing behavior of all saber-toothed cats. However, further research involving a wider range of dirk and scimitar-toothed forms could provide additional insights into the diversity of cranial biomechanics within this fascinating group of extinct mammalian predators.

Cranial functional specialisation for strength precedes morphological evolution in Oviraptorosauria

Oviraptorosaurians were a theropod dinosaur group that reached high diversity in the Late Cretaceous. Within oviraptorosaurians, the later diverging oviraptorids evolved distinctive crania which were extensively pneumatised, short and tall, and had a robust toothless beak, interpreted as providing a powerful bite for their herbivorous to omnivorous diet. The present study explores the ability of oviraptorid crania to resist large mechanical stresses compared with other theropods and where this adaptation originated within oviraptorosaurians. Digital 3D cranial models were constructed for the earliest diverging oviraptorosaurian, Incisivosaurus gauthieri, and three oviraptorids, Citipati osmolskae, Conchoraptor gracilis, and Khaan mckennai. Finite element analyses indicate oviraptorosaurian crania were stronger than those of other herbivorous theropods (Erlikosaurus and Ornithomimus) and were more comparable to the large, carnivorous Allosaurus. The cranial biomechanics of Incisivosaurus align with oviraptorids, indicating an early establishment of distinctive strengthened cranial biomechanics in Oviraptorosauria, even before the highly modified oviraptorid cranial morphology. Bite modelling, using estimated muscle forces, suggests oviraptorid crania may have functioned closer to structural safety limits. Low mechanical stresses around the beaks of oviraptorids suggest a convergently evolved, functionally distinct rhamphotheca, serving as a cropping/feeding tool rather than for stress reduction, when compared with other herbivorous theropods.

Regional variation of the cortical and trabecular bone material properties in the rabbit skull.

The material properties of some bones are known to vary with anatomical location, orientation and position within the bone (e.g., cortical and trabecular bone). Details of the heterogeneity and anisotropy of bone is an important consideration for biomechanical studies that apply techniques such as finite element analysis, as the outcomes will be influenced by the choice of material properties used. Datasets detailing the regional variation of material properties in the bones of the skull are sparse, leaving many finite element analyses of skulls no choice but to employ homogeneous, isotropic material properties, often using data from a different species to the one under investigation. Due to the growing significance of investigating the cranial biomechanics of the rabbit in basic science and clinical research, this study used nanoindentation to measure the elastic modulus of cortical and trabecular bone throughout the skull. The elastic moduli of cortical bone measured in the mediolateral and ventrodorsal direction were found to decrease posteriorly through the skull, while it was evenly distributed when measured in the anteroposterior direction. Furthermore, statistical tests showed that the variation of elastic moduli between separate regions (anterior, middle and posterior) of the skull were significantly different in cortical bone, but was not in trabecular bone. Elastic moduli measured in different orthotropic planes were also significantly different, with the moduli measured in the mediolateral direction consistently lower than that measured in either the anteroposterior or ventrodorsal direction. These findings demonstrate the significance of regional and directional variation in cortical bone elastic modulus, and therefore material properties in finite element models of the skull, particularly those of the rabbit, should consider the heterogeneous and orthotropic properties of skull bone when possible.

Divergent strategies in cranial biomechanics and feeding ecology of the ankylosaurian dinosaurs

Ankylosaurs were important megaherbivores of Jurassic and Cretaceous ecosystems. Their distinctive craniodental anatomy and mechanics differentiated them from coexisting hadrosaurs and ceratopsians, and morphological evidence suggests dietary niche partitioning between sympatric ankylosaurids and nodosaurids. Here, we investigate the skull biomechanics of ankylosaurs relative to feeding function. First, we compare feeding functional performance between nodosaurids and ankylosaurids applying finite element analysis and lever mechanics to the skulls of Panoplosaurus mirus (Nodosauridae) and Euoplocephalus tutus (Ankylosauridae). We also compare jaw performance across a wider sample of ankylosaurs through lever mechanics and phylogenetic comparative methods. Mandibular stress levels are higher in Euoplocephalus, supporting the view that Panoplosaurus consumed tougher foodstuffs. Bite force and mechanical advantage (MA) estimates indicate that Panoplosaurus had a relatively more forceful and efficient bite than Euoplocephalus. There is little support for a role of the secondary palate in resisting feeding loads in the two ankylosaur clades. Several ankylosaurs converged on similar jaw mechanics, while some nodosaurids specialised towards high MA and some ankylosaurids evolved low MA jaws. Our study supports the hypothesis that ankylosaurs partitioned dietary niches in Late Cretaceous ecosystems and reveals that the two main ankylosaur clades evolved divergent evolutionary pathways in skull biomechanics and feeding habits.

Comparative cranial biomechanics reveal that Late Cretaceous tyrannosaurids exerted relatively greater bite force than in early-diverging tyrannosauroids.

Tyrannosaurus has been an exemplar organism in feeding biomechanical analyses. An adult Tyrannosaurus could exert a bone-splintering bite force, through expanded jaw muscles and a robust skull and teeth. While feeding function of adult Tyrannosaurus has been thoroughly studied, such analyses have yet to expand to other tyrannosauroids, especially early-diverging tyrannosauroids (Dilong, Proceratosaurus, and Yutyrannus). In our analysis, we broadly assessed the cranial and feeding performance of tyrannosauroids at varying body sizes. Our sample size included small (Proceratosaurus and Dilong), medium-sized (Teratophoneus), and large (Tarbosaurus, Daspletosaurus, Gorgosaurus, and Yutyrannus) tyrannosauroids, and incorporation of tyrannosaurines at different ontogenetic stages (small juvenile Tarbosaurus, Raptorex, and mid-sized juvenile Tyrannosaurus). We used jaw muscle force calculations and finite element analysis to comprehend the cranial performance of our tyrannosauroids. Scaled subtemporal fenestrae areas and calculated jaw muscle forces show that broad-skulled tyrannosaurines (Tyrannosaurus, Daspletosaurus, juvenile Tyrannosaurus, and Raptorex) exhibited higher jaw muscle forces than other similarly sized tyrannosauroids (Gorgosaurus, Yutyrannus, and Proceratosaurus). The large proceratosaurid Yutyrannus exhibited lower cranial stress than most adult tyrannosaurids. This suggests that cranial structural adaptations of large tyrannosaurids maintained adequate safety factors at greater bite force, but their robust crania did not notably decrease bone stress. Similarly, juvenile tyrannosaurines experienced greater cranial stress than similarly-sized earlier tyrannosauroids, consistent with greater adductor muscle forces in the juveniles, and with crania no more robust than in their small adult predecessors. As adult tyrannosauroid body size increased, so too did relative jaw muscle forces manifested even in juveniles of giant adults.

Flat or flexible? Evolutionary cranial biomechanics and the origins of modern archosaur skulls

The divergent specializations of living archosaurs, crocodilians and birds, represent two of the great transformations in vertebrate evolution. Despite hailing from common ancestor with a tall skull, braced palate and relatively unspecialized jaw muscles, these two living clades followed disparate paths in cranial biomechanics and feeding functional morphology during the Mesozoic era, resulting in the flat, rigid skulls in crocodylians and the bulbous and beaked, flexible skulls of birds. Here we use new approaches in imaging, 3D modeling, morphometrics, lever mechanics, finite element modeling and phylogenetic comparative methods of living and extinct reptiles to show how morphological and functional changes in the palate, cranial joints and jaw musculature occurred within the fossil record of these lineages that were key features in acquisition of the modern forms. We show how jaw muscles reoriented their resultants and moments about jaw joints during skull flattening, palatocranial suturing and feeding specializations in Jurassic crocodyliforms. These morphological changes resulted in a decrease in mechanical efficiency of the feeding apparatus that was compensated for by an increase in jaw muscle sizes and canalized muscle architecture. Meanwhile, along the line to living birds, the acquisition of enlarged brains in coelurosaur dinosaurs resulted in a shifting of temporal and palatal muscle positions, a canalization of vertically‐oriented of muscle forces and consequently reorientation of their moments about palatocranial joints. These changes in loading environment, along with a concatenate increase in the propulsive and bending properties of the palate, facilitated cranial kinesis in neoavian birds. Together, these examples of evolutionary innovation in reptile cranial biomechanics reveal powerful new approaches to testing biomechanical and evolutionary hypotheses in any cranial or appendicular musculoskeletal system.

Early Origins of Divergent Patterns of Morphological Evolution on the Mammal and Reptile Stem-Lineages.

The origin of amniotes 320 million years ago signaled independence from water in vertebrates and was closely followed by divergences within the mammal and reptile stem lineages (Synapsida and Reptilia). Early members of both groups had highly similar morphologies, being superficially “lizard-like” forms with many plesiomorphies. However, the extent to which they might have exhibited divergent patterns of evolutionary change, with the potential to explain the large biological differences between their living members, is unresolved. We use a new, comprehensive phylogenetic dataset to quantify variation in rates and constraints of morphological evolution among Carboniferous–early Permian amniotes. We find evidence for an early burst of evolutionary rates, resulting in the early origins of morphologically distinctive subgroups that mostly persisted through the Cisuralian. Rates declined substantially through time, especially in reptiles. Early reptile evolution was also more constrained compared with early synapsids, exploring a more limited character state space. Postcranial innovation in particular was important in early synapsids, potentially related to their early origins of large body size. In contrast, early reptiles predominantly varied the temporal region, suggesting disparity in skull and jaw kinematics, and foreshadowing the variability of cranial biomechanics seen in reptiles today. Our results demonstrate that synapsids and reptiles underwent an early divergence of macroevolutionary patterns. This laid the foundation for subsequent evolutionary events and may be critical in understanding the substantial differences between mammals and reptiles today. Potential explanations include an early divergence of developmental processes or of ecological factors, warranting cross-disciplinary investigation. [Amniote; body size; constraint; phylogeny; rate.]

More Challenging Diets Sustain Feeding Performance: Applications Toward the Captive Rearing of Wildlife

SynopsisThe rescue and rehabilitation of young fauna is of substantial importance to conservation. However, it has been suggested that incongruous diets offered in captive environments may alter craniofacial morphology and hinder the success of reintroduced animals. Despite these claims, to what extent dietary variation throughout ontogeny impacts intrapopulation cranial biomechanics has not yet been tested. Here, finite element models were generated from the adult crania of 40 rats (n = 10 per group) that were reared on 4 different diet regimes and stress magnitudes compared during incisor bite simulations. The diets consisted of (1) exclusively hard pellets from weaning, (2) exclusively soft ground pellet meal from weaning, (3) a juvenile switch from pellets to meal, and (4) a juvenile switch from meal to pellets. We hypothesized that a diet of exclusively soft meal would result in the weakest adult skulls, represented by significantly greater stress magnitudes at the muzzle, palate, and zygomatic arch. Our hypothesis was supported at the muzzle and palate, indicating that a diet limited to soft food inhibits bone deposition throughout ontogeny. This finding presents a strong case for a more variable and challenging diet during development. However, rather than the “soft” diet group resulting in the weakest zygomatic arch as predicted, this region instead showed the highest stress among rats that switched as juveniles from hard pellets to soft meal. We attribute this to a potential reduction in number and activity of osteoblasts, as demonstrated in studies of sudden and prolonged disuse of bone. A shift to softer foods in captivity, during rehabilitation after injury in the wild for example, can therefore be detrimental to healthy development of the skull in some growing animals, potentially increasing the risk of injury and impacting the ability to access full ranges of wild foods upon release. We suggest captive diet plans consider not just nutritional requirements but also food mechanical properties when rearing wildlife to adulthood for reintroduction.

The cranial biomechanics and feeding performance of Homo floresiensis.

Homo floresiensis is a small-bodied hominin from Flores, Indonesia, that exhibits plesiomorphic dentognathic features, including large premolars and a robust mandible, aspects of which have been considered australopith-like. However, relative to australopith species, H. floresiensis exhibits reduced molar size and a cranium with diminutive midfacial dimensions similar to those of later Homo, suggesting a reduction in the frequency of forceful biting behaviours. Our study uses finite-element analysis to examine the feeding biomechanics of the H. floresiensis cranium. We simulate premolar (P3) and molar (M2) biting in a finite-element model (FEM) of the H. floresiensis holotype cranium (LB1) and compare the mechanical results with FEMs of chimpanzees, modern humans and a sample of australopiths (MH1, Sts 5, OH5). With few exceptions, strain magnitudes in LB1 resemble elevated levels observed in modern Homo. Our analysis of LB1 suggests that H. floresiensis could produce bite forces with high mechanical efficiency, but was subject to tensile jaw joint reaction forces during molar biting, which perhaps constrained maximum postcanine bite force production. The inferred feeding biomechanics of H. floresiensis closely resemble modern humans, suggesting that this pattern may have been present in the last common ancestor of Homo sapiens and H. floresiensis.

Computational biomechanical modelling of the rabbit cranium during mastication

Although a functional relationship between bone structure and mastication has been shown in some regions of the rabbit skull, the biomechanics of the whole cranium during mastication have yet to be fully explored. In terms of cranial biomechanics, the rabbit is a particularly interesting species due to its uniquely fenestrated rostrum, the mechanical function of which is debated. In addition, the rabbit processes food through incisor and molar biting within a single bite cycle, and the potential influence of these bite modes on skull biomechanics remains unknown. This study combined the in silico methods of multi-body dynamics and finite element analysis to compute musculoskeletal forces associated with a range of incisor and molar biting, and to predict the associated strains. The results show that the majority of the cranium, including the fenestrated rostrum, transmits masticatory strains. The peak strains generated over all bites were found to be attributed to both incisor and molar biting. This could be a consequence of a skull shape adapted to promote an even strain distribution for a combination of infrequent incisor bites and cyclic molar bites. However, some regions, such as the supraorbital process, experienced low peak strain for all masticatory loads considered, suggesting such regions are not designed to resist masticatory forces.

Comparative cranial biomechanics in two lizard species: impact of variation in cranial design.

ABSTRACTCranial morphology in lepidosaurs is highly disparate and characterised by the frequent loss or reduction of bony elements. In varanids and geckos, the loss of the postorbital bar is associated with changes in skull shape, but the mechanical principles underlying this variation remain poorly understood. Here, we sought to determine how the overall cranial architecture and the presence of the postorbital bar relate to the loading and deformation of the cranial bones during biting in lepidosaurs. Using computer-based simulation techniques, we compared cranial biomechanics in the varanid Varanus niloticus and the teiid Salvator merianae, two large, active foragers. The overall strain magnitude and distribution across the cranium were similar in the two species, despite lower strain gradients in V. niloticus. In S. merianae, the postorbital bar is important for resistance of the cranium to feeding loads. The postorbital ligament, which in varanids partially replaces the postorbital bar, does not affect bone strain. Our results suggest that the reduction of the postorbital bar impaired neither biting performance nor the structural resistance of the cranium to feeding loads in V. niloticus. Differences in bone strain between the two species might reflect demands imposed by feeding and non-feeding functions on cranial shape. Beyond variation in cranial bone strain related to species-specific morphological differences, our results reveal that similar mechanical behaviour is shared by lizards with distinct cranial shapes. Contrary to the situation in mammals, the morphology of the circumorbital region, calvaria and palate appears to be important for withstanding high feeding loads in these lizards.

Biomechanical simulations of Leptarctus primus (Leptarctinae, Carnivora), and new evidence for a badger-like feeding capability

ABSTRACTVariations in craniodental morphology have been correlated to feeding adaptations in living organisms and used as proxies for paleodiet reconstruction. Within the mammalian order Carnivora, the Miocene fossil musteloid Leptarctus has been variably interpreted as a carnivore, frugivore, herbivore, omnivore, or insectivore based on morphological comparisons with extant species. Here, we perform the first simulation of cranial biomechanics in Leptarctus primus, aiming to identify a living analogue using biomechanical capability rather than qualitative morphology. Finite element models (FEMs) of 18 extant carnivorans and two extinct outgroup taxa were used to compare known diet-biomechanics relationships with the biomechanical properties of L. primus FEMs within a phylogenetic context. Multivariate analyses of simulated bite efficiency and skull stiffness values indicate that L. primus is most similar overall to Taxidea taxus (American badger) in unilateral bite simulations. Based on biomechanical predictions, we postulate that L. primus resembled the American badger in its feeding ecology more closely than any other taxon tested and thus conclude that L. primus was dominantly a carnivore with an auxiliary feeding capability of omnivory. We also compared the L. primus FEMs with the potentially synonymous Hypsoparia bozemanensis to determine a possible range of feeding capabilities. We observed an increase in mechanical efficiency with a deepening of the zygomae of H. bozemanensis, a trait previously used to differentiate it from L. primus. Ongoing work to expand the database of cranial biomechanical simulation data across Carnivoramorpha should help to further clarify evolutionary patterns of skull biomechanical specializations in musteloids and other carnivorans.

Ontogeny, But Not Sexual Dimorphism, Drives the Intraspecific Variation of Quadrate Morphology in Hemidactylus turcicus (Squamata: Gekkonidae)

Abstract The functional components of the lizard skull are divided into the chondrocranial braincase, protective dermatocranium, lower jaw, and hyobranchial apparatus. These regions are interconnected and become operational through the quadrate, a bone critical for cranial biomechanics and support of the peripheral auditory system. The quadrate is a complex and variable structure in squamates; however, neither the intraspecific nor interspecific variation of this element has been studied in detail. We investigated the intraspecific variation of quadrate morphology within Hemidactylus turcicus with the use of cleared and double-stained specimens and high-resolution x-ray microcomputed tomography. Our objectives were to quantify quadrate shape and the degree of intraspecific variation within this element with the use of 2-D and 3-D geometric morphometric analyses and investigate if this variation is driven by ontogeny and/or sexual dimorphism. Our results demonstrate that ontogeny, but not sexual shape dimo...

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.

Bending Properties of the Jugal Bone in Mallard Ducks and its Significance for Cranial Biomechanics

Few species evolved skulls as derived as ducks and other Anseriform birds, which employ rapid, dexterous movements of the palate and jaws to feed. Despite their exceptional behavior and morphology, little is known about the structure and biomechanics of their jaw muscles, cranial joints, and linkages that facilitate cranial kinesis. In particular, the jugal bone is capable of bending considerably as it links the quadrate and the upper beak. Here we employ a series of methods including microCT imaging, morphometrics, 3D modeling and histology to discover how and why the jugal is capable of such astounding flexibility. Although a narrow suture connects the caudal limb of the jugal with the quadratojugal, flexibility instead appears to occur at narrow regions of the bone. This gross morphology is further complemented by changes to the skeletal tissues that compose the element. Moreover, the orientation of jaw muscle resultants and loading of the skull indicate the jugal experiences a range of loading during different instances of the gape cycle. These findings illustrate the complicated series of adaptations in morphology, histology and biomechanics ducks and other vertebrates employ to engage in derived forms of cranial kinesis and feeding.In birds, cranial kinesis implies the ability to move the upper bill relative to the braincase, contributing to the evolution of the flexible skulls found in birds. The mechanism for elevation of the upper beak consists of a forward movement of the quadrate, exerting force through the jugal and the pterygoid that connect the quadrate to the upper jaw, and elevate the upper beak by bending of the nasofrontal hinge joint and possibly a flexion zone in the jugal. The jugal bone plays a significant role in Mallard feeding as it links the posterior part of the skull with the flexible anterior part of the skull. Interestingly the jugal is quite flexible and when removed from the skull; it is capable of bending 90 degrees while still returning to its original conformation. For the first time, we are studying the biomechanics of birds to determine how their skulls work; while primarily investigating the cause of the flexibility in the jugal bone. We have studied a sample of duck skulls through CT data, 3D modeling, and measurement of cross‐sectional properties of the element. From this data we have found areas of the jugal bone that afford greater consideration in determining the flexibility throughout the jugal bone. This project will create new data about the jugal bone's flexibility, which will contribute to the study of biomechanics of bones and cranial kinesis.

Elastic Properties of Chimpanzee Craniofacial Cortical Bone.

Relatively few assessments of cranial biomechanics formally take into account variation in the material properties of cranial cortical bone. Our aim was to characterize the elastic properties of chimpanzee craniofacial cortical bone and compare these to the elastic properties of dentate human craniofacial cortical bone. From seven cranial regions, 27 cylindrical samples were harvested from each of five chimpanzee crania. Assuming orthotropy, axes of maximum stiffness in the plane of the cortical plate were derived using modified equations of Hooke's law in a Mathcad program. Consistent orientations among individuals were observed in the zygomatic arch and alveolus. The density of cortical bone showed significant regional variation (P < 0.001). The elastic moduli demonstrated significant differences between sites, and a distinct pattern where E3 > E2 > E1 . Shear moduli were significantly different among regions (P < 0.001). The pattern by which chimpanzee cranial cortical bone varies in elastic properties resembled that seen in humans, perhaps suggesting that the elastic properties of craniofacial bone in fossil hominins can be estimated with at least some degree of confidence. Anat Rec, 299:1718-1733, 2016. © 2016 Wiley Periodicals, Inc.

Palate anatomy and morphofunctional aspects of interpterygoid vacuities in temnospondyl cranial evolution.

Temnospondyls were the morphologically and taxonomically most diverse group of early tetrapods with a near-global distribution during the Palaeozoic and Mesozoic. Members of this group occupied a range of different habitats (aquatic, amphibious, terrestrial), reflected by large morphological disparity of the cranium throughout their evolutionary history. A diagnostic feature of temnospondyls is the presence of an open palate with large interpterygoid vacuities, in contrast to the closed palate of most other early tetrapods and their fish-like relatives. Although the function of the interpterygoid vacuities has been discussed in the past, no quantitative studies have been performed to assess their biomechanical significance. Here, we applied finite element analysis, to test the possibility that the interpterygoid vacuities served for stress distribution during contraction of the jaw closing musculature. Different original and theoretical skull models, in which the vacuities differed in size or were completely absent, were compared for their mechanical performance. Our results demonstrate that palatal morphology played a considerable role in cranial biomechanics of temnospondyls. The presence of large cranial vacuities were found to offer the dual benefit of providing additional muscle attachment areas and allowing for more effective force transmission and thus an increase in bite force without compromising cranial stability.Electronic supplementary materialThe online version of this article (doi:10.1007/s00114-016-1402-z) contains supplementary material, which is available to authorized users.