Mechanical compensation in the evolution of the early hominin feeding apparatus.

Australopiths, a group of hominins from the Plio-Pleistocene of Africa, are characterized by derived traits in their crania hypothesized to strengthen the facial skeleton against feeding loads and increase the efficiency of bite force production. The crania of robust australopiths are further thought to be stronger and more efficient than those of gracile australopiths. Results of prior mechanical analyses have been broadly consistent with this hypothesis, but here we show that the predictions of the hypothesis with respect to mechanical strength are not met: some gracile australopith crania are as strong as that of a robust australopith, and the strength of gracile australopith crania overlaps substantially with that of chimpanzee crania. We hypothesize that the evolution of cranial traits that increased the efficiency of bite force production in australopiths may have simultaneously weakened the face, leading to the compensatory evolution of additional traits that reinforced the facial skeleton. The evolution of facial form in early hominins can therefore be thought of as an interplay between the need to increase the efficiency of bite force production and the need to maintain the structural integrity of the face.

Biomechanics of the mandible of Macaca mulatta during the power stroke of mastication: Loading, deformation, and strain regimes and the impact of food type.

Mandible morphology has yet to yield definitive information on primate diet, probably because of poor understanding of mandibular loading and strain regimes, and overreliance on simple beam models of mandibular mechanics. We used a finite element model of a macaque mandible to test hypotheses about mandibular loading and strain regimes and relate variation in muscle activity during chewing on different foods to variation in strain regimes. The balancing-side corpus is loaded primarily by sagittal shear forces and sagittal bending moments. On the working side, sagittal bending moments, anteroposterior twisting moments, and lateral transverse bending moments all reach similar maxima below the bite point; sagittal shear is the dominant loading regime behind the bite point; and the corpus is twisted such that the mandibular base is inverted. In the symphyseal region, the predominant loading regimes are lateral transverse bending and negative twisting about a mediolateral axis. Compared with grape and dried fruit chewing, nut chewing is associated with larger sagittal and transverse bending moments acting on balancing- and working-side mandibles, larger sagittal shear on the working side, and larger twisting moments about vertical and transverse axes in the symphyseal region. Nut chewing is also associated with higher minimum principal strain magnitudes in the balancing-side posterior ramus; higher sagittal shear strain magnitudes in the working-side buccal alveolar process and the balancing-side oblique line, recessus mandibulae, and endocondylar ridge; and higher transverse shear strains in the symphyseal region, the balancing-side medial prominence, and the balancing-side endocondylar ridge. The largest food-related differences in maximum principal and transverse shear strain magnitudes are in the transverse tori and in the balancing-side medial prominence, extramolar sulcus, oblique line, and endocondylar ridge. Food effects on the strain regime are most salient in areas not traditionally investigated, suggesting that studies seeking dietary effects on mandible morphology might be looking in the wrong places.

Masticatory properties in pre-modern Holocene populations from Northern China.

Recent studies indicate that evolution of the craniofacial skeleton is influenced by dietary behavior, which in turn alters masticatory efficacy and temporomandibular joint (TMJ) stability. In this study the mechanical properties of the masticatory system and the integrity of the TMJ in human populations from Northern China, dated to between 3800 BCE and 100 CE, were assessed. The results demonstrate that the mechanical efficiency is comparable to other modern human populations, though variations are present across different populations. While the ratio of overall weighted muscle efficiency for incisor loading vs. molar loading in pastoral and some recent agricultural groups is similar to early Homo sapiens, the ratio in more ancient agricultural groups is similar to the ratio in populations with heavy anterior paramasticatory activities, such as Neandertals, Inuits, and Native Americans. The TMJ vulnerability negatively correlates with the maxillary dental arch size, and positively with the condylar size. These findings suggest that there are multi-directional strategies in adaptation to heavy anterior teeth loading, such as increasing anterior teeth loading efficiency, increasing facial height, increasing facial breath and facial orthognathy, or decreasing anterior facial length. Furthermore, populations or individuals with a smaller dental arch and high biting efficiency could more easily injure the TMJ during unilateral loadings, which may explain the higher prevalence of TMJ disorders in modern humans, especially in women. These findings further reflect the impact of diachronic changes of the masticatory apparatus and lifestyle and their impact on oral health during recent human history.

Two distalization methods compared in a novel patient-specific finite element analysis.

INTRODUCTION: Orthodontic mini-implants aid in the correction of distocclusions via direct anchorage (pull from mini-implant to teeth) and indirect anchorage (teeth pulled against other teeth anchored by the mini-implant). The aim of this study was to compare stress levels on the periodontal ligament (PDL) of maxillary buccal teeth in direct and indirect distalization against orthodontic mini-implants and accounting for individual variation in maxillary anatomy and biomechanical characteristics of the compact bone. METHODS: A 3D model of the maxilla containing the different components (teeth, PDL, trabecular and cortical bones) was generated from a computed tomographic scan. Cortical bone was divided into several areas according to previously defined zones. Bone stiffness and thickness data, obtained from 11 and 12 cadavers, respectively, were incorporated into the initial model to simulate the individual cortical bone variation at the different locations. Subsequently, a finite element analysis was used to simulate the distalization modalities. RESULTS: Stresses at the buccal, palatal, mesial, and distal surfaces were significantly different between adjacent teeth under stiffness but not thickness variation. In both distalization modalities, low or no significant correlations were found between stress values and corresponding cortical bone thicknesses. High significant and inverted correlations were observed at the first molar between stress amounts and cortical bone stiffness (direct modality: -0.68 < r < -0.72; indirect modality: -0.80 < r < -0.82; P <0.05). CONCLUSIONS: With the use of a novel finite element approach that integrated human data on variations in bone properties, findings suggested that cortical bone stiffness may influence tooth movement more than bone thickness. Significant clinical implications could be related to these findings.

Divided zygoma in Holocene human populations from Northern China.

OBJECTIVES: Divided zygoma (DZ) occurs in contemporaneous human populations, with the highest incidences in people from East Asia and Southern Africa. The present study examines the prevalence and variation of this condition in the Holocene populations of Northern China for the first time. METHODS: In this study, 1145 skulls from various human populations living in Northern China from the Neolithic Age to recent dynasties (5000-300 years BP) were examined. Specifically, cranial measurements and a CT scan were conducted to quantify craniofacial morphology. RESULTS: Fifteen skulls were identified with DZ, revealing an overall prevalence of 1.3% in the collection, while it was determined to be higher in North Asian and Northeast Asian regional groups. In skulls with unilateral DZ, the superior division of the zygoma was generally slender, while the inferior division of the zygoma was more robust. In skulls with bilateral DZ, the maxillae were generally more laterally extended. Moreover, unilateral DZ skulls displayed differences in cortical bone thickness between two sides of the facial skeleton. DISCUSSION: In context, the distribution pattern within these data points toward a greater prevalence of the DZ phenotype in North and Northeast Asian regional groups, suggesting a hypothesis that the DZ trait is more frequent in populations characterized by flat and broad faces. Accordingly, further studies into the DZ condition will deepen our understanding of developments in plasticity, variation, and recent evolution of the human cranium.

Odontogenic abscesses in rhesus macaques (Macaca mulatta) of Cayo Santiago.

OBJECTIVES: Odontogenic abscesses are one of the most common dental diseases causing maxillofacial skeletal lesions. They affect the individual's ability to maintain the dental structures necessary to obtain adequate nutrition for survival and reproduction. In this study, the prevalence and pattern of odontogenic abscesses in relation to age, sex, matriline, and living periods were investigated in adult rhesus macaque skeletons of the free-ranging colony on Cayo Santiago, Puerto Rico. MATERIALS AND METHODS: The skulls used for this study were from the skeletons of 752 adult rhesus macaques, aged 8-31 years, and born between 1951 and 2000. They came from 66 matrilines ranging from 1 to 88 individuals. Fistulae or skeletal lesions caused by odontogenic abscesses drainage, carious lesions, tooth fractures, tooth loss, and alveolar resorption were evaluated visually. RESULTS: Seventy-two specimens (9.57%) had odontogenic abscesses of varying severity. Males had a significantly higher prevalence than females. The prevalence of odontogenic abscesses in several matrilines was significantly higher than in the population as a whole. Animals born between 1950 and 1965 tended to have a higher prevalence of odontogenic abscesses than those born in later periods. DISCUSSION: These results suggest that oral pathologies, such as dental and periodontal abscesses in rhesus macaques are fairly common, which may indicate familial effects interwoven with ecological and social factors. The closeness of the rhesus and human genomes allows insights to understand of the epidemiology of these diseases in the human population. Further assessment of the role played by environmental and familial factors on rhesus oral health and disease are warranted.

VEGF-C promotes the development of lymphatics in bone and bone loss.

Patients with Gorham-Stout disease (GSD) have lymphatic vessels in their bones and their bones gradually disappear. Here, we report that mice that overexpress VEGF-C in bone exhibit a phenotype that resembles GSD. To drive VEGF-C expression in bone, we generated Osx-tTA;TetO-Vegfc double-transgenic mice. In contrast to Osx-tTA mice, Osx-tTA;TetO-Vegfc mice developed lymphatics in their bones. We found that inhibition of VEGFR3, but not VEGFR2, prevented the formation of bone lymphatics in Osx-tTA;TetO-Vegfc mice. Radiological and histological analysis revealed that bones from Osx-tTA;TetO-Vegfc mice were more porous and had more osteoclasts than bones from Osx-tTA mice. Importantly, we found that bone loss in Osx-tTA;TetO-Vegfc mice could be attenuated by an osteoclast inhibitor. We also discovered that the mutant phenotype of Osx-tTA;TetO-Vegfc mice could be reversed by inhibiting the expression of VEGF-C. Taken together, our results indicate that expression of VEGF-C in bone is sufficient to induce the pathologic hallmarks of GSD in mice.

In vivo bone strain and finite element modeling of a rhesus macaque mandible during mastication.

Finite element analysis (FEA) is a commonly used tool in musculoskeletal biomechanics and vertebrate paleontology. The accuracy and precision of finite element models (FEMs) are reliant on accurate data on bone geometry, muscle forces, boundary conditions and tissue material properties. Simplified modeling assumptions, due to lack of in vivo experimental data on material properties and muscle activation patterns, may introduce analytical errors in analyses where quantitative accuracy is critical for obtaining rigorous results. A subject-specific FEM of a rhesus macaque mandible was constructed, loaded and validated using in vivo data from the same animal. In developing the model, we assessed the impact on model behavior of variation in (i) material properties of the mandibular trabecular bone tissue and teeth; (ii) constraints at the temporomandibular joint and bite point; and (iii) the timing of the muscle activity used to estimate the external forces acting on the model. The best match between the FEA simulation and the in vivo experimental data resulted from modeling the trabecular tissue with an isotropic and homogeneous Young's modulus and Poisson's value of 10GPa and 0.3, respectively; constraining translations along X,Y, Z axes in the chewing (left) side temporomandibular joint, the premolars and the m1; constraining the balancing (right) side temporomandibular joint in the anterior-posterior and superior-inferior axes, and using the muscle force estimated at time of maximum strain magnitude in the lower lateral gauge. The relative strain magnitudes in this model were similar to those recorded in vivo for all strain locations. More detailed analyses of mandibular strain patterns during the power stroke at different times in the chewing cycle are needed.

Biomechanical implications of cortical elastic properties of the macaque mandible.

Knowledge of the variation in the elastic properties of mandibular cortical bone is essential for modeling bone function. Our aim was to characterize the elastic properties of rhesus macaque mandibular cortical bone and compare these to the elastic properties from mandibles of dentate humans and baboons. Thirty cylindrical samples were harvested from each of six adult female rhesus monkey mandibles. Assuming orthotropy, axes of maximum stiffness in the plane of the cortical plate were derived from ultrasound velocity measurements. Further velocity measurements with longitudinal and transverse ultrasonic transducers along with measurements of bone density were used to compute three-dimensional cortical elastic properties using equations based on Hooke's law. Results showed regional variations in the elastic properties of macaque mandibular cortical bone that have both similarities and differences with that of humans and baboons. So far, the biological and structural basis of these differences is poorly understood.

Elevation of a full-thickness mucoperiosteal flap alone accelerates orthodontic tooth movement.

INTRODUCTION: Our objective was to determine whether the elevation of a full-thickness mucoperiosteal flap alone, without cortical cuts, decreases the amount of bone around teeth and accelerates mesial tooth movements. METHODS: The mandibular second premolars of 7 beagle dogs were extracted, and on a randomly selected side, a full-thickness mucoperiosteal buccal flap extending from the distal aspect of the third premolar to the mesial aspect of the first premolar was elevated. The other side did not receive flap surgery. The mandibular third premolars were protracted with orthodontic appliances. Tooth movements were analyzed biweekly over an 8-week period with calipers and radiographs. The amount and density of bone were analyzed using microcomputed tomography; bone remodeling was evaluated with histologic sections. RESULTS: Experimental tooth movements measured intraorally between cusp tips were significantly greater (25.3%) than control tooth movements. The approximate center of resistance measured radiographically also moved significantly more (about 31%) on the experimental than on the control side. The experimental premolar tipped more than the control premolar (10.5° vs 8.7°), but the difference was not statistically significant. The medullary bone volume fraction mesial to the third premolar was significantly less (9.1%) and the bone was significantly less dense (9%) on the experimental side than on the control side. Histology showed no apparent side differences in the numbers of osteoclasts and osteoblasts evident in the medullary bone. CONCLUSIONS: Elevation of a full-thickness mucoperiosteal flap alone (ie, without injury to bone) decreases the amount and density of medullary bone surrounding the tooth and accelerates tooth movement. Due to its limited effects, elevation of a flap alone to increase tooth movements may not be justified.

Evolution of the Jugal/Zygomatic Bones.

This issue of the Anatomical Record is the second of a two-volume set on the zygoma (also called the cheek bone, the zygomatic bone, the malar, or the jugal, the latter term being used in vertebrates other than mammals). The zygoma is an important component of the craniofacial skeleton, in which the zygoma is a connection between the midfacial and the cranial skeletons; has a functional role as the origin of one of the masticatory muscles, the masseter muscle, and several facial muscles; has been considered as an essential buttress of the facial skeleton for resisting masticatory forces; and has importance for determining phylogenetic relationships. In humans, the zygoma is also of aesthetic significance for facial appearance, and its restoration following trauma has resulted in a large clinical literature. In this second half of the special issue on the zygoma, a series of papers discuss studies related to evolution of the zygoma and related parts of the craniofacial skeleton throughout the vertebrates, and in particular in human evolution. There are also a series of articles discussing variation of the zygoma in modern humans. This article is an overview in which we discuss the primary findings of these studies and some of their implications. Anat Rec, 300:12-15, 2017. © 2016 Wiley Periodicals, Inc.

The Biomechanics of Bony Facial "Buttresses" in South African Australopiths: An Experimental Study Using Finite Element Analysis.

Australopiths exhibit a number of derived facial features that are thought to strengthen the face against high and/or repetitive loads associated with a diet that included mechanically challenging foods. Here, we use finite element analysis (FEA) to test hypotheses related to the purported strengthening role of the zygomatic root and "anterior pillar" in australopiths. We modified our previously constructed models of Sts 5 (Australopithecus africanus) and MH1 (A. sediba) to differ in the morphology of the zygomatic root, including changes to both the shape and positioning of the zygomatic root complex, in addition to creating variants of Sts 5 lacking anterior pillars. We found that both an expanded zygomatic root and the presence of "anterior pillars" reinforce the face against feeding loads. We also found that strain orientations are most compatible with the hypothesis that the pillar evolved to resist loads associated with premolar loading, and that this morphology has an ancillary effect of strengthening the face during all loading regimes. These results provide support for the functional hypotheses. However, we found that an anteriorly positioned zygomatic root increases strain magnitudes even in models with an inflated/reinforced root complex. These results suggest that an anteriorly placed zygomatic root complex evolved to enhance the efficiency of bite force production while facial reinforcement features, such as the anterior pillar and the expanded zygomatic root, may have been selected for in part to compensate for the weakening effect of this facial configuration. Anat Rec, 300:171-195, 2017. © 2016 Wiley Periodicals, Inc.

Development, Structure, and Function of the Zygomatic Bones: What is New and Why Do We Care?

This issue of The Anatomical Record is the first of a two-volume set on the zygoma (also called the cheek bone, the zygomatic bone, the malar, or the jugal, the latter term being used in vertebrates other than mammals). The zygoma is an important component of the craniofacial skeleton, in which the zygoma is a connection between the midfacial and the cranial skeletons; has a functional role as the origin of one of the masticatory muscles, the masseter muscle, and several facial muscles; has been considered as an essential buttress of the facial skeleton for resisting masticatory forces; and has importance for determining phylogenetic relationships. In humans, the zygoma is also of aesthetic significance for facial appearance, and its restoration following trauma has resulted in a large clinical literature. In this first volume of this Special Issue, a wide ranging series of papers discuss studies related to issues of development, structure, and function of the zygoma and closely related parts of the craniofacial skeleton in mammals, and in particular primates. This Introductory article provides an overview in which we discuss the primary findings of these studies and some of their implications. The second volume, which will be published as the January 2017 issue of The Anatomical Record, will focus on variation and evolution of the zygoma throughout the vertebrates. Anat Rec, 299:1611-1615, 2016. © 2016 Wiley Periodicals, Inc.

Divided Zygomatic Bone in Primates With Implications of Skull Morphology and Biomechanics.

Typically the zygoma is a single bone in the facial skeleton whose shape uniquely copes with loads associated with mastication. Rarely but naturally, the zygoma is divided into two or more parts by supernumerary sutures. These extra intrazygomatic sutures are located at an area of critical morphological and biomechanical importance, yet their impacts have not been studied. In this study, the morphological and possible biomechanical consequences of the divided zygoma (DZ) were investigated in primates including rhesus macaques (Macaca mulatta), orangutans (Pongo abelii and P. pygmaeus), and modern humans (Homo sapiens). Results demonstrated that a unilateral supernumerary suture within the zygoma affected facial symmetry. The superior division of the divided zygoma was normally slender along with the adjacent frontal bone parts; while the inferior division of the divided zygoma was normally more robust, along with stronger temporal and maxillary bones. These were possible biomechanical consequences, in which the stresses incurred during normal masticatory activities were shunted from the upper face to the lower face, especially along the zygomatic arch. These findings revealed that the DZ condition would alter overall morphology of the midface of the affected side, and unfavorably affect the pattern of stress distribution in the loaded side of the face during mastication. The developmental mechanisms for the supernumerary sutures dividing the zygoma were unclear. Further insights into this rare condition may deepen our understanding of craniofacial form, adaptation, developmental plasticity, and evolution, and help to improve therapeutic philosophies in corrective and regenerative medicine. Anat Rec, 299:1801-1829, 2016. © 2016 Wiley Periodicals, Inc.

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.

Internal Bone Architecture in the Zygoma of Human and Pan.

The internal and external anatomy of the primate zygoma is central to orofacial function, health, and disease. The importance of variation in its gross morphology across extinct and extant primate forms has been established using finite element analysis, but its internal structure has yet to be explored. In this study, µCT is used to characterize trabecular bone morphometry in two separate regions of the zygoma of humans and Pan. Trabecular anisotropy and orientation are compared with strain orientations observed in trabecular regions of finite element models of four Pan crania. The results of this study show that trabecular bone morphometry, anisotropy, and orientation are highly compatible with strain orientation and magnitude in the finite element models. Trabecular bone in the zygoma is largely orthotropic (with bone orientation differing in three mutually orthogonal directions), with its primary orientation lying in the mediolateral direction. Trabecular bone in the zygomatic region appears to be highly influenced by the local strain environment, and thus may be closely linked to orofacial function. Anat Rec, 299:1704-1717, 2016. © 2016 Wiley Periodicals, Inc.

Review of In Vivo Bone Strain Studies and Finite Element Models of the Zygomatic Complex in Humans and Nonhuman Primates: Implications for Clinical Research and Practice.

The craniofacial skeleton is often described in the clinical literature as being comprised of vertical bony pillars, which transmit forces from the toothrow to the neurocranium as axial compressive stresses, reinforced transversely by buttresses. Here, we review the literature on bony microarchitecture, in vivo bone strain, and finite-element modeling of the facial skeleton of humans and nonhuman primates to address questions regarding the structural and functional existence of facial pillars and buttresses. Available bone material properties data do not support the existence of pillars and buttresses in humans or Sapajus apella. Deformation regimes in the zygomatic complex emphasize bending and shear, therefore conceptualizing the zygomatic complex of humans or nonhuman primates as a pillar obscures its patterns of stress, strain, and deformation. Human fossil relatives and chimpanzees exhibit strain regimes corroborating the existence of a canine-frontal pillar, but the notion of a zygomatic pillar has no support. The emerging consensus on patterns of strain and deformation in finite element models (FEMs) of the human facial skeleton corroborates hypotheses in the clinical literature regarding zygomatic complex function, and provide new insights into patterns of failure of titanium and resorbable plates in experimental studies. It is suggested that the "pillar and buttress" model of human craniofacial skeleton function be replaced with FEMs that more accurately and precisely represent in vivo function, and which can serve as the basis for future research into implants used in restoration of occlusal function and fracture repair. Anat Rec, 299:1753-1778, 2016. © 2016 Wiley Periodicals, Inc.

Human feeding biomechanics: performance, variation, and functional constraints.

The evolution of the modern human (Homo sapiens) cranium is characterized by a reduction in the size of the feeding system, including reductions in the size of the facial skeleton, postcanine teeth, and the muscles involved in biting and chewing. The conventional view hypothesizes that gracilization of the human feeding system is related to a shift toward eating foods that were less mechanically challenging to consume and/or foods that were processed using tools before being ingested. This hypothesis predicts that human feeding systems should not be well-configured to produce forceful bites and that the cranium should be structurally weak. An alternate hypothesis, based on the observation that humans have mechanically efficient jaw adductors, states that the modern human face is adapted to generate and withstand high biting forces. We used finite element analysis (FEA) to test two opposing mechanical hypotheses: that compared to our closest living relative, chimpanzees (Pan troglodytes), the modern human craniofacial skeleton is (1) less well configured, or (2) better configured to generate and withstand high magnitude bite forces. We considered intraspecific variation in our examination of human feeding biomechanics by examining a sample of geographically diverse crania that differed notably in shape. We found that our biomechanical models of human crania had broadly similar mechanical behavior despite their shape variation and were, on average, less structurally stiff than the crania of chimpanzees during unilateral biting when loaded with physiologically-scaled muscle loads. Our results also show that modern humans are efficient producers of bite force, consistent with previous analyses. However, highly tensile reaction forces were generated at the working (biting) side jaw joint during unilateral molar bites in which the chewing muscles were recruited with bilateral symmetry. In life, such a configuration would have increased the risk of joint dislocation and constrained the maximum recruitment levels of the masticatory muscles on the balancing (non-biting) side of the head. Our results do not necessarily conflict with the hypothesis that anterior tooth (incisors, canines, premolars) biting could have been selectively important in humans, although the reduced size of the premolars in humans has been shown to increase the risk of tooth crown fracture. We interpret our results to suggest that human craniofacial evolution was probably not driven by selection for high magnitude unilateral biting, and that increased masticatory muscle efficiency in humans is likely to be a secondary byproduct of selection for some function unrelated to forceful biting behaviors. These results are consistent with the hypothesis that a shift to softer foods and/or the innovation of pre-oral food processing techniques relaxed selective pressures maintaining craniofacial features that favor forceful biting and chewing behaviors, leading to the characteristically small and gracile faces of modern humans.

Mechanical evidence that Australopithecus sediba was limited in its ability to eat hard foods.

Australopithecus sediba has been hypothesized to be a close relative of the genus Homo. Here we show that MH1, the type specimen of A. sediba, was not optimized to produce high molar bite force and appears to have been limited in its ability to consume foods that were mechanically challenging to eat. Dental microwear data have previously been interpreted as indicating that A. sediba consumed hard foods, so our findings illustrate that mechanical data are essential if one aims to reconstruct a relatively complete picture of feeding adaptations in extinct hominins. An implication of our study is that the key to understanding the origin of Homo lies in understanding how environmental changes disrupted gracile australopith niches. Resulting selection pressures led to changes in diet and dietary adaption that set the stage for the emergence of our genus.