Inferences regarding the diet of extinct hominins: structural and functional trends in dental and mandibular morphology within the hominin clade

This contribution investigates the evolution of diet in the Pan-Homo and hominin clades. It does this by focusing on 12 variables (nine dental and three mandibular) for which data are available about extant chimpanzees, modern humans and most extinct hominins. Previous analyses of this type have approached the interpretation of dental and gnathic function by focusing on the identification of the food consumed (i.e. fruits, leaves, etc.) rather than on the physical properties (i.e. hardness, toughness, etc.) of those foods, and they have not specifically addressed the role that the physical properties of foods play in determining dental adaptations. We take the available evidence for the 12 variables, and set out what the expression of each of those variables is in extant chimpanzees, the earliest hominins, archaic hominins, megadont archaic hominins, and an inclusive grouping made up of transitional hominins and pre-modern Homo. We then present hypotheses about what the states of these variables would be in the last common ancestor of the Pan-Homo clade and in the stem hominin. We review the physical properties of food and suggest how these physical properties can be used to investigate the functional morphology of the dentition. We show what aspects of anterior tooth morphology are critical for food preparation (e.g. peeling fruit) prior to its ingestion, which features of the postcanine dentition (e.g. overall and relative size of the crowns) are related to the reduction in the particle size of food, and how information about the macrostructure (e.g. enamel thickness) and microstructure (e.g. extent and location of enamel prism decussation) of the enamel cap might be used to make predictions about the types of foods consumed by extinct hominins. Specifically, we show how thick enamel can protect against the generation and propagation of cracks in the enamel that begin at the enamel-dentine junction and move towards the outer enamel surface.     

Dental and skeletal growth in early fossil hominins

Early fossil hominins have often been assigned a chronological age on the basis of modern human data for tooth eruption. Better data and more sophisticated methods are now available to estimate their chronological age from modern human standards for stages of mineralization of individual teeth developing within the jaws. However, while comparisons with modern human dentitions are interesting, they can also be misleading as early hominin teeth and dentitions did not grow like modern human teeth. Chronological age can also be estimated using the microanatomy of tooth enamel and root dentine. Counts of incremental markings in enamel predict much younger ages at death for early fossil hominins than those based on modern human radiographic standards of dental development. Comparative evidence from the skeleton suggests that a greater proportion of adult body mass and stature was achieved earlier in the growth period of fossil hominins than it is in modern humans. The combined skeleto-dental evidence provides the basis for a hypothesis that the earliest hominins grew more like modern great apes, but that Homo erectus had a slightly more prolonged period of growth, and which was still not totally modern human-like in its pattern or timing.     

On the scaling relationship between enamel prism length and enamel thickness in primate molars: a comment

As part of a larger study we developed a computer programme which allows the recreation of the complex 3-dimensional arrangement of prisms. Data presented in these earlier publications are re-analyzed to assess the relationship between projected prism length (i.e., enamel thickness) and the true prism length. Across primates, proportional prism deviation increases as the enamel becomes thicker. This supports suggestions that prism decussation may be particularly marked in large-bodied and thick-enameled species. There are differences, however, in scaling relationships between species, which correspond to the species' dietary adaptations. Finally, the findings highlight the importance of employing species-specific correction factors for the calculation of prism length (i.e., life-span of the ameloblast) for life history enquiry.     

Megadontia, striae periodicity and patterns of enamel secretion in Plio-Pleistocene fossil hominins

Early hominins formed large and thick-enamelled cheek-teeth within relatively short growth periods as compared with modern humans. To understand better the developmental basis of this process, we measured daily enamel increments, or cross striations, in 17 molars of Plio-Pleistocene hominins representing seven different species, including specimens attributed to early Homo. Our results show considerable variation across species, although all specimens conformed to the known pattern characterised by greater values in outer than inner enamel, and greater cuspal than cervical values. We then compared our results with the megadontia index, which represents tooth size in relation to body mass, for each species to assess the effect of daily growth rates on tooth size. Our results indicate that larger toothed (megadont) taxa display higher rates or faster forming enamel than smaller toothed hominins. By forming enamel quickly, large tooth crowns were able to develop within the constraints of shorter growth periods. Besides daily increments, many animals express long-period markings (striae of Retzius) in their enamel. We report periodicity values (number of cross striations between adjacent striae) in 14 new specimens of Australopithecus afarensis, Paranthropus aethiopicus, Paranthropus boisei, Homo habilis, Homo rudolfensis and Homo erectus, and show that long-period striae express a strong association with male and average male–female body mass. Our results for Plio-Pleistocene hominins show that the biological rhythms that give rise to long-period striae are encompassed within the range of variation known for modern humans, but show a lower mean and modal value of 7 days in australopithecines. In our sample of early Homo, mean and modal periodicity values were 8 days, and therefore similar to modern humans. These new data on daily rates of enamel formation and periodicity provide a better framework to interpret surface manifestations of internal growth markings on fossil hominin tooth crowns. Importantly, our data on early hominin cross striation variation may now contribute towards solving difficult taxonomic diagnoses where much may depend on fragmentary molar remains and enamel structure.     

Comparison between deciduous and permanent incisor teeth in morphology of bovine enamel.

Functional incisor teeth (deciduous and permanent teeth) from Bovidae (14 species) were prepared for scanning electron microscopic observation. Ultrastructural patterns of the enamel layer of deciduous and permanent incisor teeth varied (ex. prisms, arrangement pattern of matrices, and in thickness of enamel layer) in each species. The ultrastructures of prisms in longitudinal sections were classified into three types; A, radial, B, tangential, and C, mix of A and B arrangement enamel; modified Koenigswald's method (1982) in examined species. Type A was found in a large part of permanent and a small part of deciduous incisor teeth, while types B and C were mainly found in the deciduous teeth. These morphological features show the remarkable correlation between permanent and deciduous teeth.     

Role of crystal arrangement on the mechanical performance of enamel

The superior mechanical properties of enamel, such as excellent penetration and crack resistance, are believed to be related to the unique microscopic structure. In this study, the effects of hydroxyapatite (HAP) crystallite orientation on the mechanical behavior of enamel have been investigated through a series of multiscale numerical simulations. A micromechanical model, which considers the HAP crystal arrangement in enamel prisms, the hierarchical structure of HAP crystals and the inelastic mechanical behavior of protein, has been developed. Numerical simulations revealed that, under compressive loading, plastic deformation progression took place in enamel prisms, which is responsible for the experimentally observed post-yield strain hardening. By comparing the mechanical responses for the uniform and non-uniform arrangement of HAP crystals within enamel prisms, it was found that the stiffness for the two cases was identical, while much greater energy dissipation was observed in the enamel with the non-uniform arrangement. Based on these results, we propose an important mechanism whereby the non-uniform arrangement of crystals in enamel rods enhances energy dissipation while maintaining sufficient stiffness to promote fracture toughness, mitigation of fracture and resistance to penetration deformation. Further simulations indicated that the non-uniform arrangement of the HAP crystals is a key factor responsible for the unique mechanical behavior of enamel, while the change in the nanostructure of nanocomposites could dictate the Young's modulus and yield strength of the biocomposite.     

Three-dimensional changes in direction and interrelationships among enamel prisms in the dog tooth.

This study describes the three-dimensional features of enamel prisms and their arrangement in dog teeth. Tangential semithin sections of demineralized tooth germ were serially cut from the enamel surface to the enamel-dentin junction. Straight rows of enamel prisms parallel or perpendicular to the meridian were selected at the enamel-dentin junction; these prisms were reconstructed from micrographs with a personal computer. Near the enamel-dentin junction, the arrangement of enamel prisms appeared regular. Viewed from the enamel surface, the cut-ends of the enamel prisms that were parallel to the meridian at the enamel-dentin junction appeared as a sine curve, with 16 enamel prisms forming one period. The enamel prisms in a row perpendicular to the meridian were parallel to each other and deflected to the left or right from the enamel-dentin junction. Away from the enamel-dentin junction, the periodicity of the prisms gradually disappeared. The sine curve formed by the cut-ends of prisms in a row parallel to the meridian became irregular, and prisms in rows perpendicular to the meridian crossed each other. The semithin sections showed belt-like zones arranged perpendicular to the meridian. Each belt-like zone consisted of enamel prisms oriented in the same direction, those in neighboring zones being oriented in opposite directions. The disappearance of the regular arrangement of prisms was related to changes in their location in the belt-like zones.     

Locomotion and posture from the common hominoid ancestor to fully modern hominins, with special reference to the last common panin/hominin ancestor

Based on our knowledge of locomotor biomechanics and ecology we predict the locomotion and posture of the last common ancestors of (a) great and lesser apes and their close fossil relatives (hominoids); (b) chimpanzees, bonobos and modern humans (hominines); and (c) modern humans and their fossil relatives (hominins). We evaluate our propositions against the fossil record in the context of a broader review of evolution of the locomotor system from the earliest hominoids of modern aspect (crown hominoids) to early modern Homo sapiens. While some early East African stem hominoids were pronograde, it appears that the adaptations which best characterize the crown hominoids are orthogrady and an ability to abduct the arm above the shoulder - rather than, as is often thought, manual suspension sensu stricto. At 7-9 Ma (not much earlier than the likely 4-8 Ma divergence date for panins and hominins, see Bradley, 2008) there were crown hominoids in southern Europe which were adapted to moving in an orthograde posture, supported primarily on the hindlimb, in an arboreal, and possibly for Oreopithecus, a terrestrial context. By 7 Ma, Sahelanthropus provides evidence of a Central African hominin, panin or possibly gorilline adapted to orthogrady, and both orthogrady and habitually highly extended postures of the hip are evident in the arboreal East African protohominin Orrorin at 6 Ma. If the traditional idea that hominins passed through a terrestrial 'knuckle-walking' phase is correct, not only does it have to be explained how a quadrupedal gait typified by flexed postures of the hindlimb could have preadapted the body for the hominin acquisition of straight-legged erect bipedality, but we would have to accept a transition from stem-hominoid pronogrady to crown hominoid orthogrady, back again to pronogrady in the African apes and then back to orthogrady in hominins. Hand-assisted arboreal bipedality, which is part of a continuum of orthograde behaviours, is used by modern orangutans to forage among the small branches at the periphery of trees where the core hominoid dietary resource, ripe fruit, is most often to be found. Derivation of habitual terrestrial bipedality from arboreal hand-assisted bipedality requires fewer transitions, and is also kinematically and kinetically more parsimonious.     

Assessment of the accuracy of dental enamel thickness measurements using microfocal X-ray computed tomography

Tooth enamel thickness has long been an important character in studies of primate and especially hominin phylogeny, taxonomy, and adaptation. Current methods for accurately assessing enamel thickness involve the physical sectioning of teeth, because measurements of enamel thickness using some radiographic techniques are unreliable. However, because destructive methods limit sample sizes and access to important fossil specimens, it is desirable that they be replaced with nondestructive techniques. Although microfocal X-ray computed tomography (mCT) has been used recently in studies of enamel thickness, the accuracy of this technique has yet to be established. The present research compares physical sections to computer-generated mCT sections of teeth from a variety of primate and nonprimate, recent and fossil taxa to examine whether enamel thickness, tooth size, and diagenetic remineralization (fossilization) impact the ability of mCT to measure enamel thickness accurately. Results indicate that recent teeth of varying size and thickness are clearly and accurately depicted in mCT scans, with measurements from nearly identical planes in physical and mCT sections differing by 3-5%. A fossil papionin molar (ca. 2 Myr) was also accurately measured using mCT scans, although thinner enamel in much older therapsid (ca. 263-241 Myr) teeth could not be distinguished from dentine. mCT is thus an accurate technique for measuring enamel thickness in recent taxa, although heavily mineralized teeth pose an obstacle to the ability of mCT to distinguish dental tissues. Moreover, absolutely thin enamel (less than approximately 0.10 mm) is difficult to resolve adequately in raw mCT images based on pixel values alone. Therefore, caution must be exercised in the application of mCT to the study of fossilized teeth.     

Femoral mechanics in the lesser bushbaby (Galago senegalensis): Structural adaptations to leaping in primates

One method used to examine the relationship between behavioral strategies and anatomical adaptation is to study the results of mechanical stress associated with a given behavior and compare this with skeletal adaptations to other behaviors. This comparative approach is appropriate for highlighting combinations of features that are specializations to specific types of behavior. The purpose of this paper is to compare femoral mechanics in Galago senegalensis with previously collected data for macaques and humans as a basis for discussing structural adaptations in the primate hindlimb to leaping. The stiffness and load carrying capabilities of the femoral diaphyses of 27 G. senegalensis were analyzed using the SCADS computer program. The data suggest that the galago femur is well adapted to sustain large sagittal plane compressive loads rather than large bending loads. The straightness of the femoral shaft and large midshaft area moments of inertia prevent buckling from these large compressive loads. Calculations indicate that the ratio of critical buckling load to body weight in galago is 31 times that in macaques and 55 times that in humans. The femur of this saltatory primate is morphologically adapted to resist buckling when subjected to large compressive loads, while those of macaques and humans are better adapted to resist bending moments caused by ground reaction forces acting on the extended limb. The differences between galago on the one hand and macaques and humans on the other suggest that relatively smaller moments about the hip and relatively larger moments about the knee accompany more quadrupedal and bipedal walking, while habitual leaping is associated with relatively larger moments about the hip. These data reinforce the apparent similarity of the mechanical effects of quadrupedal and bipedal locomotion on the femur and dissimilarity with femoral mechanics in habitually saltatory primates.     

The Sima de los Huesos Middle Pleistocene hominin site (Burgos,Spain). Estimation of the number of individuals

After 34 years of research and findings in the Middle Pleistocene site of the Sima de los Huesos (SH) of the Sierra de Atapuerca (Burgos, Spain), we present an update of the estimation of the number of individuals (ENI) identified in the SH hominin assemblage. The last ENI, published in 2004, was 28. Although the number of specimens recovered has almost doubled since then and more complete information is now available, this new analysis suggests that the ENI is 29. Some individuals are still represented by only one or a few teeth and the upper and lower jaws of various individuals have not been completed. We suggest that the amateur cavers, who accessed the SH site for years looking for bear fossils, destroyed a significant number of hominin fossils. Despite this, we have a good picture of the group of hominins represented in the SH assemblage. Because complete corpses were accumulated in SH and it is a closed cavity, it is expected that a significant number of hominin fossils remains to be discovered. According to the previous estimates, the representation of females is greater than that of males. However, the observed sex ratio is not significantly different from the 1:1 ratio. With the exception of the possible presence of a child individual, most of the remaining 28 individuals are teenagers or young adults (24/28 = 0.857). The ages of death of SH hominins appear to conform to a catastrophic profile.     

A fourth hominin skull from Dmanisi, Georgia

Newly discovered Homo remains, stone artifacts, and animal fossils from Dmanisi, Republic of Georgia, provide a basis for better understanding patterns of hominin evolution and behavior in Eurasia ca. 1.77 million years ago. Here we describe a fourth skull that is nearly complete, lacking all but one of its teeth at the time of death. Both the maxillae and the mandible exhibit extensive bone loss due to resorption. This individual is similar to others from the site but supplies information about variation in brain size and craniofacial anatomy within the Dmanisi paleodeme. Although this assemblage presents numerous primitive characters, the Dmanisi skulls are best accommodated within the species H. erectus. On anatomical grounds, it is argued that the relatively small-brained and lightly built Dmanisi hominins may be ancestral to African and Far Eastern branches of H. erectus showing more derived morphology.     

The development of enamel structure in rat incisors as compared to the teeth of monkey and man

The rat incisor is an excellent model system in which to study amelgenesis. However, the information obtained has not been extrapolated to the human because of alleged structural differences between the teeth. The obvious differences include continuous eruption in rat incisors and an enamel rod pattern in rats which seemingly differs from the keyhole pattern of human enamel. A comprehensive analysis was made of those features of enamel structure considered fundamental to the understanding of its formation. This was done by applying the knowledge of amelogenesis obtained in rat incisors to the teeth of monkey and man. The following points of basic similarity were established between these species: (1) Interrod enamel is secreted first. It forms the side walls of cavities which are initially occupied by Tomes' processes. (2) The formation of interrod cavities is followed by deposition of enamel rods within these spaces. (3) The rods conform to the shape of the cavities and are secreted from one surface of Tomes' process. (4) At the initial site of rod deposition its enamel is continuous with the interrod enamel wall. (5) Growth of the rod compresses the process to one side of the cavity resulting in an arcade-shaped "space" between the rod and the remaining interrod walls. This study demonstrates that it is no longer necessary to postulate a keyhole structure for primate enamel, and it has established that a fundamental similarity exists in the basic structure and in the mode of formation of enamel in all three species.     

Structure–function relations of primate lower incisors: a study of the deformation of Macaca mulatta dentition using electronic speckle pattern interferometry (ESPI)

Teeth adopt a variety of different morphologies, each of which is presumably optimized for performing specific functions during feeding. It is generally agreed that the enamel cap is a crucial element in controlling the mechanical behavior of mammalian teeth under load. Incisors are particularly interesting in terms of structure–function relations, as their role in feeding is that of the ‘first bite’. However, little is known how incisor cap morphology is related to tooth deformation. In the present paper we examine the mechanical behavior of mandibular central incisors in the cercopithecine primate Macaca mulatta under loads similar to those encountered during ingestion. We map three‐dimensional displacements on the labial surface of the crown as it is compressed, using electronic speckle pattern interferometry (ESPI), an optical metrology method. In addition, micro‐computed tomography is used to obtain data regarding the morphology of the enamel cap, which in the M. mulatta lower incisors exhibits missing or very little enamel on the lingual face. The results showed that although compressed along a longitudinal axis, deformation in the incisors mostly occurred in the lingual direction and orthogonal to the direction of the applied load. Both isolated, embedded teeth and teeth in the mandible showed considerable lingual deformation. Incisor deformation in the mandible was generally greater, reflecting the additional freedom of movement enabled by the supporting structures. We show that the association with adjacent teeth in the arch is significant for the behavior of the tooth under load. Finally, loading two teeth simultaneously in the mandible showed that they work as one functional unit. We suggest that these results demonstrate the importance of enamel cap morphology in directing deformation behavior; an ability stemming from the stiffness of the enamel cap overlying the more pliable dentin.     

Correlation of the arrangement pattern of enamel rods and secretory ameloblasts in pig and monkey teeth: a possible role of the terminal webs in ameloblast movement during secretion.

Enamel rod architecture and ameloblast arrangement were examined in pig and monkey teeth using light microscopy and scanning and transmission electron microscopy. Enamel rods in the pig teeth were arranged in longitudinal straight rows in the initial enamel layer, in longitudinal wavy rows in the inner enamel layer, and in a staggered pattern in the outer enamel layer. Rod decussation was seen only in the inner layer. Cross-sectioned enamel rods in the pig were arcade-shaped in the initial and inner layers, and mostly round in shape with circular boundaries in the outer layer. Arrangement of secretory ameloblasts at the level of the distal terminal web and Tomes' processes, and shape of Tomes' processes, corresponded to those of the enamel rod in the enamel layers. Distal terminal webs were well developed between straight rows of the ameloblasts forming the initial layer and between wavy rows of the ameloblasts forming the inner layer, and less developed within a row. The filament bundles in the distal terminal webs were also oriented along the rows. However, in the ameloblasts forming the outer layer, which lost their row pattern, distal terminal web filaments were distributed uniformly at the cell periphery. A similar arrangement of wavy rows of ameloblasts at the level of distal terminal web and Tomes' processes was also seen in monkey teeth.     

Correlation of the arrangement pattern of enamel rods and secretory ameloblasts in pig and monkey teeth: A possible role of the terminal webs in ameloblast movement during secretion

Enamel rod architecture and ameloblast arrangement were examined in pig and monkey teeth using light microscopy and scanning and transmission electron microscopy. Enamel rods in the pig teeth were arranged in longitudinal straight rows in the initial enamel layer, in longitudinal wavy rows in the inner enamel layer, and in a staggered pattern in the outer enamel layer. Rod decussation was seen only in the inner layer. Cross-sectined enamel rods in the pig were arcade-shaped in the initial and inner layers, and mostly round in shape with circular boundaries in the outer layer. Arrangement of secretory ameloblasts at the level of the distal terminal web and Tomes' processes, and shape of Tomes' processes, corresponded to those of the enamel rod in the enamel layers. Distal terminal webs were well developed between straight rows of the ameloblasts forming the initial layer and between wavy rows of the ameloblasts forming the inner layer, and less developed within a row. The filament bundles in the distal terminal webs were also oriented along the rows. However, in the ameloblasts forming the outer layer, which lost their row pattern, distal terminal web filaments were distributed uniformly at the cell periphery. A similar arrangement of wavy rows of ameloblasts at the level of distal terminal web and Tomes' processes was also seen in monkey teeth.     

The evolutionary history of the hominin hand since the last common ancestor of Pan and Homo

Molecular evidence indicates that the last common ancestor of the genus Pan and the hominin clade existed between 8 and 4 million years ago (Ma). The current fossil record indicates the Pan-Homo last common ancestor existed at least 5 Ma and most likely between 6 and 7 Ma. Together, the molecular and fossil evidence has important consequences for interpreting the evolutionary history of the hand within the tribe Hominini (hominins). Firstly, parsimony supports the hypothesis that the hand of the last common ancestor most likely resembled that of an extant great ape overall (Pan, Gorilla, and Pongo), and that of an African ape in particular. Second, it provides a context for interpreting the derived changes to the hand that have evolved in various hominins. For example, the Australopithecus afarensis hand is likely derived in comparison with that of the Pan-Homo last common ancestor in having shorter fingers relative to thumb length and more proximo-distally oriented joints between its capitate, second metacarpal, and trapezium. This evidence suggests that these derived features evolved prior to the intensification of stone tool-related hominin behaviors beginning around 2.5 Ma. However, a majority of primitive features most likely present in the Pan-Homo last common ancestor are retained in the hands of Australopithecus, Paranthropus/early Homo, and Homo floresiensis. This evidence suggests that further derived changes to the hands of other hominins such as modern humans and Neandertals did not evolve until after 2.5 Ma and possibly even later than 1.5 Ma, which is currently the earliest evidence of Acheulian technology. The derived hands of modern humans and Neandertals may indicate a morphological commitment to tool-related manipulative behaviors beyond that observed in other hominins, including those (e.g. H. floresiensis) which may be descended from earlier tool-making species.     

The initial process of enamel prism arrangement and its relation to the Hunter-Schreger bands in dog teeth

The three-dimensional architecture of enamel prisms at early stages of enamel formation and its spatial relationship to the Hunter-Schreger bands were examined in canine tooth germs by light and electron microscopy. In serial semithin sections of demineralized tooth germs tangential to the enamel-dentin junction, a straight row of enamel prisms was depicted along the longitudinal tooth axis at the level of the enamel-dentin junction and then their three-dimensional arrangement was reconstructed using computer software. The spatial arrangement of the groups of enamel rods oriented in specific sideward directions was also reconstructed in deep layers of the enamel. Initially, all enamel prisms were parallel to perpendicular toward the enamel-dentin junction, but at 10μm from the enamel-dentin junction, some small specks, or groups of enamel prisms--tilting to the right or the left--emerged as small islands. In each speck of enamel prism, the inclined prisms were uniformly oriented in a sideward direction and gradually expanded their boundary until merging with the neighboring specks inclined in the same direction. Consequently, at 50μm from the enamel-dentin junction, the group of enamel prisms oriented either to the right or the left formed alternately arranged horizontal belt-like zones, corresponding to the parazone or the diazone of the Hunter-Schreger bands. Reversed images of scanning electron-micrographs of the exposed surfaces of the developing enamel revealed round and bulb-like profiles of Tomes' processes at early amelogenesis and its changes into a characteristic structure combined with flat secretory and enclosing nonsecretory faces that dictated the orientation of corresponding enamel prisms. The results suggest that the groups of enamel prisms oriented in sideward directions first appear as small island-like specks near the enamel-dentin junction, which later merge and form alternating horizontal belt-like zones as a consequence of morphological changes of the Tomes' processes. However, the mechanisms whereby the functional grouping of secretory ameloblasts with similarly oriented Tomes' processes is induced are yet to be determined.     

The Effective Mechanical Advantage of A.L. 129‐1a for Knee Extension

The functional significance of shape differences between modern human and australopithecine distal femora remains unclear. Here, we examine the morphological component of the effective mechanical advantage (EMA) of the quadriceps muscle group in a sample of hominins that includes the fossil A.L. 129‐1a (Australopithecus afarensis) and modern humans. Quadriceps muscle moment arms were calculated from three‐dimensional computer models of specimens through a range of knee flexion. All hominins were compared using the same limb positions to allow us to examine, in isolation, the morphological component of the lengths of the pertinent moment arms. After taking into account the differences in bicondylar angle, the morphological component of the EMA was calculated as the ratio of the quadriceps muscle and ground reaction force moment arms. Our analyses reveal that A.L. 129‐1a would have possessed a morphological component of the quadriceps muscle EMA expected for a hominin of its body mass. Anat Rec, 2011. © 2011 Wiley‐Liss, Inc.