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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.

Fossil hominin footprints preserve data on a remarkably short time scale compared to most other fossil evidence, offering snapshots of organisms in their immediate ecological and behavioral contexts. Here, we report on our excavations and analyses of more than 400 Late Pleistocene human footprints from Engare Sero, Tanzania. The site represents the largest assemblage of footprints currently known from the human fossil record in Africa. Speed estimates show that the trackways reflect both walking and running behaviors. Estimates of group composition suggest that these footprints were made by a mixed-sex and mixed-age group, but one that consisted of mostly adult females. One group of similarly-oriented trackways was attributed to 14 adult females who walked together at the same pace, with only two adult males and one juvenile accompanying them. In the context of modern ethnographic data, we suggest that these trackways may capture a unique snapshot of cooperative and sexually divided foraging behavior in Late Pleistocene humans.

A ∼1.52 Ma adult upper limb skeleton of Paranthropus boisei (KNM-ER 47000) recovered from the Koobi Fora Formation, Kenya (FwJj14E, Area 1A) includes most of the distal half of a right humerus (designated KNM-ER 47000B). Natural transverse fractures through the diaphysis of KNM-ER 470000B provide unobstructed views of cortical bone at two sections typically used for analyzing cross-sectional properties of hominids (i.e., 35% and 50% of humerus length from the distal end). Here we assess cross-sectional properties of KNM-ER 47000B and two other P. boisei humeri (OH 80-10, KNM-ER 739). Cross-sectional properties for P. boisei associated with bending/torsional strength (section moduli) and relative cortical thickness (%CA; percent cortical area) are compared to those reported for nonhuman hominids, AL 288-1 (Australopithecus afarensis), and multiple species of fossil and modern Homo. Polar section moduli (Zp) are assessed relative to a mechanically relevant measure of body size (i.e., the product of mass [M] and humerus length [HL]). At both diaphyseal sections, P. boisei exhibits %CA that is high among extant hominids (both human and nonhuman) and similar to that observed among specimens of Pleistocene Homo. High values for Zp relative to size (M × HL) indicate that P. boisei had humeral bending strength greater than that of modern humans and Neanderthals and similar to that of great apes, A. afarensis, and Homo habilis. Such high humeral strength is consistent with other skeletal features of P. boisei (reviewed here) that suggest routine use of powerful upper limbs for arboreal climbing.

Major changes in body shape occurred during human evolution, but questions remain about body shape in australopiths. The present study investigates the specifics of the presumed relationships between limb indices and positional behavior underlying prior work that compared proportions among extant hominids in order to make inferences about extinct hominins. We find that although both intermembral index or ratio of diaphyseal and articular proportions distinguish humans from great apes, neigher correlates well with variation in the degree of arboreality in the locomotor repertoire of extant hominids. Brachial index and a ratio of diaphyseal and articular dimensions from the fore- and hindlimb, however, do correlate with degree of arboreality, and scale slightly positively allometrically within species in all extant taxa. These two observations are taken into consideration in a more nuanced interpretation of a reanalysis of articular-diaphyseal limb proportions in an expanded sample of the Sterkfontein postcrania. This study confirms previous findings that Australopithecus africanus had larger forelimb dimensions in relation to hindlimb dimensions than modern humans and A. afarensis, similar to the patterns seen in extant apes, particularly western gorillas. However, data presented here suggest that interpreting a particular taxon as “human-like” or “ape-like” may be overly simplistic. Instead, while both A. africanus and A. afarensis were almost certainly committed bipeds that incorporated some arboreality into their locomotor repertoire, A. africanus apparently used a set of locomotor behaviors that was more distinct from that of A. afarensis than Pan troglodytes and Gorilla gorilla are from each other.

This chapter presents description and analysis of the metacarpals and manual phalanges from Sterkfontein. Although the morphology is generally similar across the sample where there are duplicates of the same element, there are differences in size that are quite remarkable within the context of all South African hominins. Some very large specimens suggest the presence of individuals at Sterkfontein with much larger hands, and presumably larger body size, at Sterkfontein than those of A. sediba MH2, H. naledi and the Swartkrans hominins. Australopithecus africanus had human-like proportions, but this may be plesiomorphic within the hominoid clade. The potentially less mobile trapezium-Mc1 joint, absence of a fully developed palmar pulp on the distal thumb, more limited pronation of the index finger, and potentially more wedge-shaped trapezoid inferred from the preserved external morphology, is consistent with lower manipulative loading of the thumb than is typical of later Homo. As for other forelimb elements, moderately curved manual phalanges suggests a greater reliance on forelimb-dominated locomotor behaviors and perhaps selection for more frequent use of an arboreal environment in A. africanus than is found in A. afarensis. Thus, within this broader context, the Sterkfontein fossil hominin remains are not unusual. The Sterkfontein hand fossils suggest an overall manipulative and locomotor loading regime that was more similar to that of other South African australopiths and distinct from that of later Homo, but more refined functional interpretations require additional fossil evidence, particularly from associated hand skeletons

Reconstructing function from hominin fossils is complicated by disagreements over how to interpret primitively inherited, ape-like morphology. This has led to considerable research on aspects of skeletal morphology that may be sensitive to activity levels during life. We quantify trabecular bone morphology in three volumes of interest (dorsal, central, and palmar) in the third metacarpal heads of extant primates that differ in hand function: Pan troglodytes, Pongo pygmaeus, Papio anubis, and Homo sapiens. Results show that bone volume within third metacarpal heads generally matches expectations based on differences in function, providing quantitative support to previous studies. Pongo shows significantly low bone volume in the dorsal region of the metacarpal head. Humans show a similar pattern, as manipulative tasks mostly involve flexed and neutral metacarpo-phalangeal joint postures. In contrast, Pan and Papio have relatively high bone volume in dorsal and palmar regions, which are loaded during knuckle-walking/digitigrady and climbing, respectively. Regional variation in degree of anisotropy did not match predictions. Although trabecular morphology may improve behavioral inferences from fossils, more sophisticated quantitative strategies are needed to explore trabecular spatial distributions and their relationships to hand function.

We report on the radioisotopic age, formation, and preservation of a late Pleistocene human footprint site in northern Tanzania on the southern shore of Lake Natron near the village of Engare Sero. Over 400 human footprints, as well as tracks of zebra and bovid, are preserved in a series of volcaniclastic deposits. Based on field mapping along with geochemical and grain-size analyses, we propose that these deposits originated as proximal volcanic material from the nearby active volcano, Oldoinyo L'engai, and were then fluvially transported to the footprint site. Stable isotope results (delta O-18 and delta C-13) suggest that the footprints were originally emplaced on a mudflat saturated by a freshwater spring and were later inundated by the rising alkaline waters of Lake Natron. We employed the Ar-40/Ar-39 and C-14 dating methods to investigate the age of the site and determined that the footprint level is older than 5760 +/- 30 yrs. BP and younger than 19.1 +/- 3.1 ka. These radioisotopic ages are supported by stratigraphic correlations with previously documented debris avalanche deposits and the stable isotope signatures associated with the most recent highstand of Lake Natron, further constraining the age to latest Pleistocene. Since modern humans (Homo sapiens) were present in Africa ca. 200 ka, Engare Sero represents the most abundant and best-preserved footprint site of anatomically modern Homo sapiens currently known in Africa. Fossil footprints are a snapshot in time, recording behavior at a specific moment in history; but the actual duration of time captured by the snapshot is often not well defined. Through analog experiments, we constrain the depositional window in which the prints were made, buried, and ultimately preserved to within a few hours to days or months. (C) 2016 Elsevier B.V. All rights reserved.

Objectives The diet of tufted capuchins (Sapajus) is characterized by annual or seasonal incorporation of mechanically protected foods. Reliance on these foods raises questions about the dietary strategies of young individuals that lack strength and experience to access these resources. Previous research has demonstrated differences between the feeding competencies of adult and juvenile tufted capuchins. Here we test the hypothesis that, compared to adults, juveniles will process foods with lower toughness and elastic moduli. Materials and Methods We present data on variation in the toughness and elastic modulus of food tissues processed by Sapajus libidinosus during the dry season at Fazenda Boa Vista, Brazil. Food mechanical property data were collected using a portable universal mechanical tester. Results Results show that food tissues processed by the capuchins showed significant differences in toughness and stiffness. However, we found no relationship between an individual's age and mean or maximum food toughness or elastic modulus, indicating both juvenile and adult S. libidinosus are able to process foods of comparable properties. Discussion Although it has been suggested that juveniles avoid mechanically protected foods, age-related differences in feeding competence are not solely due to variation in food toughness or stiffness. Other factors related to food type (e.g., learning complex behavioral sequences, achieving manual dexterity, obtaining physical strength to lift stone tools, or recognizing subtle cues about food state) combined with food mechanical properties better explain variation in juvenile feeding competency. Am J Phys Anthropol, 2015. © 2015 Wiley Periodicals, Inc.

A debate has emerged in the last few years over the shape and position of the shoulder in early Homo. That the shoulder joint underwent changes approximately 2 million years ago is not in dispute. A number of newly discovered and relatively complete scapulae show that the orientation of the glenohumeral joint shifted caudally from the more cranial orientation seen in the apes and earlier hominins (Walker and Leakey, 1993; Larson et al., 2007; Lordkipanidze et al., 2007; Haile-Selassie et al., 2010; Green and Alemseged, 2012; Churchill et al., 2013). However, just howmodern human-like this caudally rotated shoulder complex is remains less clear. Larson (2007, 2009) has proposed that early Homo possessed a novel, transitional shoulder morphology in which the shoulder joint faced anteriorly. We have proposed that Homo erectus had an essentially modern human-like shoulder complex with a laterally oriented glenohumeral joint (Roach et al., 2013; Roach and Richmond, 2015). Why does this debate matter? These differing reconstructions of the shoulder have important functional implications for a number of crucial behavioral shifts hypothesized to occur at or near the origins of our genus (e.g., reduced climbing behavior, intensification of tool manufacture and use, endurance running, and high speed throwing). Much of this debate has hinged on the length of the clavicle. As the only bony strut attaching the shoulder complex to the torso

In a broad range of evolutionary studies, an understanding of intraspecific variation is needed in order to contextualize and interpret the meaning of variation between species. However, mechanical analyses of primate crania using experimental or modeling methods typically encounter logistical constraints that force them to rely on data gathered from only one or a few individuals. This results in a lack of knowledge concerning the mechanical significance of intraspecific shape variation that limits our ability to infer the significance of interspecific differences. This study uses geometric morphometric methods (GM) and finite element analysis (FEA) to examine the biomechanical implications of shape variation in chimpanzee crania, thereby providing a comparative context in which to interpret shape-related mechanical variation between hominin species. Six finite element models (FEMs) of chimpanzee crania were constructed from CT scans following shape-space Principal Component Analysis (PCA) of a matrix of 709 Procrustes coordinates (digitized onto 21 specimens) to identify the individuals at the extremes of the first three principal components. The FEMs were assigned the material properties of bone and were loaded and constrained to simulate maximal bites on the P3 and M2. Resulting strains indicate that intraspecific cranial variation in morphology is associated with quantitatively high levels of variation in strain magnitudes, but qualitatively little variation in the distribution of strain concentrations. Thus, interspecific comparisons should include considerations of the spatial patterning of strains rather than focus only on their magnitudes. Anat Rec, 298:122–144, 2015. © 2014 Wiley Periodicals, Inc.

The African Plio-Pleistocene hominins known as australopiths evolved derived craniodental features frequently interpreted as adaptations for feeding on either hard, or compliant/tough foods. Among australopiths, Paranthropus boisei is the most robust form, exhibiting traits traditionally hypothesized to produce high bite forces efficiently and strengthen the face against feeding stresses. However, recent mechanical analyses imply that P. boisei may not have been an efficient producer of bite force and that robust morphology in primates is not necessarily strong. Here we use an engineering method, finite element analysis, to show that the facial skeleton of P. boisei is structurally strong, exhibits a strain pattern different from that in chimpanzees (Pan troglodytes) and Australopithecus africanus, and efficiently produces high bite force. It has been suggested that P. boisei consumed a diet of compliant/tough foods like grass blades and sedge pith. However, the blunt occlusal topography of this and other species suggests that australopiths are adapted to consume hard foods, perhaps including grass and sedge seeds. A consideration of evolutionary trends in morphology relating to feeding mechanics suggests that food processing behaviors in gracile australopiths evidently were disrupted by environmental change, perhaps contributing to the eventual evolution of Homo and Paranthropus.

Paranthropus boisei was first described in 1959 based on fossils from the Olduvai Gorge and now includes many fossils from Ethiopia to Malawi. Knowledge about its postcranial anatomy has remained elusive because, until recently, no postcranial remains could be reliably attributed to this taxon. Here, we report the first associated hand and upper limb skeleton (KNM-ER 47000) of P. boisei from 1.51 to 1.53 Ma sediments at Ileret, Kenya. While the fossils show a combination of primitive and derived traits, the overall anatomy is characterized by primitive traits that resemble those found in Australopithecus, including an oblique scapular spine, relatively long and curved ulna, lack of third metacarpal styloid process, gracile thumb metacarpal, and curved manual phalanges. Very thick cortical bone throughout the upper limb shows that P. boisei had great upper limb strength, supporting hypotheses that this species spent time climbing trees, although probably to a lesser extent than earlier australopiths. Hand anatomy shows that P. boisei, like earlier australopiths, was capable of the manual dexterity needed to create and use stone tools, but lacked the robust thumb of Homo erectus, which arguably reflects adaptations to the intensification of precision grips and tool use. KNM-ER 47000 provides conclusive evidence that early Pleistocene hominins diverged in postcranial and craniodental anatomy, supporting hypotheses of competitive displacement among these contemporaneous hominins.

Multiple hominin species used and produced stone tools, and the archaeological record provides evidence that stone tool behaviors intensified among later members of the genus Homo . This intensification is widely thought to be the product of cognitive and anatomical adaptations that enabled later Homo taxa to produce stone tools more efficiently relative to earlier hominin species. This study builds upon recent investigations of the knapping motions of modern humans to test whether aspects of our upper limb anatomy contribute to accuracy and/or efficiency. Knapping kinematics were captured from eight experienced knappers using a Vicon motion capture system. Each subject produced a series of Oldowan bifacial choppers under two conditions: with normal wrist mobility and while wearing a brace that reduced wrist extension (∼30°–35°), simulating one aspect of the likely primitive hominin condition. Under normal conditions, subjects employed a variant of the proximal-to-distal joint sequence common to throwing activities: subjects initiated down-swing upper limb motion at the shoulder and proceeded distally, increasing peak linear and angular velocities from the shoulder to the elbow to the wrist. At the wrist, subjects utilized the ‘dart-thrower's arc,’ the most stable plane of radiocarpal motion, during which wrist extension is coupled with radial deviation and flexion with ulnar deviation. With an unrestrained wrist, subjects achieved significantly greater target accuracy, wrist angular velocities, and hand linear velocities compared with the braced condition. Additionally, the modern wrist's ability to reach high degrees of extension (≥28.5°) following strike may decrease risk of carpal and ligamentous damage caused by hyperextension. These results suggest that wrist extension in humans contributes significantly to stone tool-making performance.

In a broad range of evolutionary studies, an understanding of intraspecific variation is needed in order to contextualize and interpret the meaning of variation between species. However, mechanical analyses of primate crania using experimental or modeling methods typically encounter logistical constraints that force them to rely on data gathered from only one or a few individuals. This results in a lack of knowledge concerning the mechanical significance of intraspecific shape variation that limits our ability to infer the significance of interspecific differences. This study uses geometric morphometric methods (GM) and finite element analysis (FEA) to examine the biomechanical implications of shape variation in chimpanzee crania, thereby providing a comparative context in which to interpret shape-related mechanical variation between hominin species. Six finite element models (FEMs) of chimpanzee crania were constructed from CT scans following shape-space Principal Component Analysis (PCA) of a matrix of 709 Procrustes coordinates (digitized onto 21 specimens) to identify the individuals at the extremes of the first three principal components. The FEMs were assigned the material properties of bone and were loaded and constrained to simulate maximal bites on the P(3) and M(2) . Resulting strains indicate that intraspecific cranial variation in morphology is associated with quantitatively high levels of variation in strain magnitudes, but qualitatively little variation in the distribution of strain concentrations. Thus, interspecific comparisons should include considerations of the spatial patterning of strains rather than focus only on their magnitudes.

The African Plio-Pleistocene hominins known as australopiths evolved derived craniodental features frequently interpreted as adaptations for feeding on either hard, or compliant/tough foods. Among australopiths, Paranthropus boisei is the most robust form, exhibiting traits traditionally hypothesized to produce high bite forces efficiently and strengthen the face against feeding stresses. However, recent mechanical analyses imply that P. boisei may not have been an efficient producer of bite force and that robust morphology in primates is not necessarily strong. Here we use an engineering method, finite element analysis, to show that the facial skeleton of P. boisei is structurally strong, exhibits a strain pattern different from that in chimpanzees (Pan troglodytes) and Australopithecus africanus, and efficiently produces high bite force. It has been suggested that P. boisei consumed a diet of compliant/tough foods like grass blades and sedge pith. However, the blunt occlusal topography of this and other species suggests that australopiths are adapted to consume hard foods, perhaps including grass and sedge seeds. A consideration of evolutionary trends in morphology relating to feeding mechanics suggests that food processing behaviors in gracile australopiths evidently were disrupted by environmental change, perhaps contributing to the eventual evolution of Homo and Paranthropus.

Fossil footprints preserve the only direct evidence of the external foot morphologies and gaits of extinct hominin taxa. However, their interpretation requires an understanding of the complex interaction among foot anatomy, foot function, and soft sediment mechanics. We applied an experimental approach aimed at understanding how one measure of foot function, the distribution of plantar pressure, influences footprint topography. Thirty-eight habitually unshod and minimally shod Daasanach individuals (19 male, 19 female) walked across a pressure pad and produced footprints in sediment directly excavated from the geological layer that preserves 1.5 Ma fossil footprints at Ileret, Kenya. Calibrated pressure data were collected and three-dimensional models of all footprints were produced using photogrammetry. We found significant correlations (Spearman's rank, p 0.0001) between measurements of plantar pressure distribution and relative footprint depths at ten anatomical regions across the foot. Furthermore, plantar pressure distributions followed a pattern similar to footprint topography, with areas of higher pressure tending to leave deeper impressions. This differs from the results of experimental studies performed in different types of sediment, supporting the hypothesis that sediment type influences the relationship between plantar pressure and footprint topography. Our results also lend support to previous interpretations that the shapes of the Ileret footprints preserve evidence of a medial transfer of plantar pressure during late stance phase, as seen in modern humans. However, the weakness of the correlations indicates that much of the variation in relative depths within footprints is not explained by pressure distributions under the foot when walking on firm ground, using the methods applied here. This warrants caution when interpreting the unique foot anatomies and foot functions of extinct hominins evidenced by their footprint structures. Further research is necessary to clarify how anatomical, functional, and sedimentary variables influence footprint formation and how each can be inferred from footprint morphology.

Recent biomechanical analyses examin- ing the feeding adaptations of early hominins have yielded results consistent with the hypothesis that hard foods exerted a selection pressure that influenced the evolution of australopith morphology. However, this hy- pothesis appears inconsistent with recent reconstructions of early hominin diet based on dental microwear and sta- ble isotopes. Thus, it is likely that either the diets of some australopiths included a high proportion of foods these taxa were poorly adapted to consume (i.e., foods that they would not have processed efficiently), or that aspects of what we thought we knew about the functional morphology of teeth must be wrong. Evaluation of these possibilities requires a recognition that analyses based on microwear, isotopes, finite element modeling, and enamel chips and cracks each test different types of hypotheses and allow different types of inferences. Microwear and isotopic analyses are best suited to recon- structing broad dietary patterns, but are limited in their ability to falsify specific hypotheses about morphological adaptation. Conversely, finite element analysis is a tool for evaluating the mechanical basis of form-function rela- tionships, but says little about the frequency with which specific behaviors were performed or the particular types of food that were consumed. Enamel chip and crack anal- yses are means of both reconstructing diet and examin- ing biomechanics. We suggest that current evidence is consistent with the hypothesis that certain derived aus- tralopith traits are adaptations for consuming hard foods, but that australopiths had generalized diets that could include high proportions of foods that were both compliant and tough. Am J Phys Anthropol 151:339-355

KNM-ER 47000 is a fossil hominin upper limb skeleton from the Koobi Fora Formation, Kenya (FwJj14E, Area 1A) that includes portions of the scapula, humerus, ulna, and hand. Dated to ∼1.52 Ma, the skeleton could potentially belong to one of multiple hominin species that have been documented in the Turkana Basin during this time, including Homo habilis, Homo erectus, and Paranthropus boisei. Although the skeleton lacks associated craniodental material, the partial humerus (described here) preserves anatomical regions (i.e., distal diaphysis, elbow joint) that are informative for taxonomic identification among early Pleistocene hominins. In this study, we analyze distal diaphyseal morphology and the shape of the elbow region to determine whether KNM-ER 47000 can be confidently attributed to a particular species. The morphology of the KNM-ER 47000 humerus (designated KNM-ER 47000B) is compared to that of other early Pleistocene hominin fossil humeri via the application of multivariate ordination techniques to both two-dimensional landmark data (diaphysis) and scale-free linear shape data (elbow). Distance metrics reflecting shape dissimilarity between KNM-ER 47000B and other fossils (and species average shapes) are assessed in the context of intraspecific variation within modern hominid species (Homo sapiens, Pan troglodytes, Gorilla gorilla, Pongo pygmaeus). Our comparative analyses strongly support attribution of KNM-ER 47000 to P. boisei. Compared to four other partial skeletons that have (justifiably or not) been attributed to P. boisei, KNM-ER 47000 provides the most complete picture of upper limb anatomy in a single individual. The taxonomic identification of KNM-ER 47000 makes the skeleton an important resource for testing future hypotheses related to P. boisei upper limb function and the taxonomy of isolated early Pleistocene hominin remains.

Tracks can provide unique, direct records of behaviors of fossil organisms moving across their landscapes millions of years ago. While track discoveries have been rare in the human fossil record, over the last decade our team has uncovered multiple sediment surfaces within the Okote Member of the Koobi Fora Formation near Ileret, Kenya that contain large assemblages of ∼1.5 Ma fossil hominin tracks. Here, we provide detailed information on the context and nature of each of these discoveries, and we outline the specific data that are preserved on the Ileret hominin track surfaces. We analyze previously unpublished data to refine and expand upon earlier hypotheses regarding implications for hominin anatomy and social behavior. While each of the track surfaces discovered at Ileret preserves a different amount of data that must be handled in particular ways, general patterns are evident. Overall, the analyses presented here support earlier interpretations of the ∼1.5 Ma Ileret track assemblages, providing further evidence of large, human-like body sizes and possibly evidence of a group composition that could support the emergence of certain human-like patterns of social behavior. These data, used in concert with other forms of paleontological and archaeological evidence that are deposited on different temporal scales, offer unique windows through which we can broaden our understanding of the paleobiology of hominins living in East Africa at ∼1.5 Ma.

Bipedalism is a key adaptation that shaped human evolution, yet the timing and nature of its evolution remain unclear. Here we use new experimentally based approaches to investigate the locomotor mechanics preserved by the famous Pliocene hominin footprints from Laetoli, Tanzania. We conducted footprint formation experiments with habitually barefoot humans and with chimpanzees to quantitatively compare their footprints to those preserved at Laetoli. Our results show that the Laetoli footprints are morphologically distinct from those of both chimpanzees and habitually barefoot modern humans. By analysing biomechanical data that were collected during the human experiments we, for the first time, directly link differences between the Laetoli and modern human footprints to specific biomechanical variables. We find that the Laetoli hominin probably used a more flexed limb posture at foot strike than modern humans when walking bipedally. The Laetoli footprints provide a clear snapshot of an early hominin bipedal gait that probably involved a limb posture that was slightly but significantly different from our own, and these data support the hypothesis that important evolutionary changes to hominin bipedalism occurred within the past 3.66 Myr.

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.

KNM-ER 47000A is a new 1.52 Ma hominin scapular fossil belonging to an associated partial skeleton from the Koobi Fora Formation, Kenya (FwJj14E, Area 1A). This fossil effectively doubles the record of Early Pleistocene scapulae from East Africa, with KNM-WT 15000 (early African Homo erectus) preserving the only other known scapula to date. KNM-ER 47000A consists of a complete glenoid cavity preserving a portion of the scapular spine and neck, the proximal half of the acromion, and a majority of the axillary border. A sufficient amount of anatomy is preserved to compare KNM-ER 47000A with scapulae of several Australopithecus species, extinct Homo, and living hominoids. The glenohumeral joint of KNM-ER 47000A is more laterally oriented than those of great apes and Australopithecus, aligning it closely with KNM-WT 15000 and modern humans. While this morphology does not imply a strong commitment to arboreality, its scapular spine is obliquely oriented—as in gorillas and some Australopithecus fossils—particularly when compared to the more horizontal orientation seen in KNM-WT 15000 and modern humans. Such a spine orientation suggests a narrow yet long infraspinous region, a feature that has been attributed to suspensory taxa. Accordingly, the morphology of KNM-ER 47000A presents conflicting behavioral implications. Nonetheless, a multivariate consideration of the available scapular traits aligns KNM-ER 47000A and Australopithecus with great apes, whereas KNM-WT 15000 resembles modern humans. The scapular morphology of KNM-ER 47000A is unique among fossil and extant hominoids and its morphological differences from KNM-WT 15000 strengthen the attribution of KNM-ER 47000 to Paranthropus boisei as opposed to early Homo. As the first evidence of scapular morphology in P. boisei, KNM-ER 47000A provides important new information on variation in hominin shoulder and upper limb anatomy from this critical period of hominin evolutionary history.

Reconstructing hominin paleoecology is critical for understanding our ancestors’ diets, social organizations and interactions with other animals. Most paleoecological models lack fine-scale resolution due to fossil hominin scarcity and the time-averaged accumulation of faunal assemblages. Here we present data from 481 fossil tracks from northwestern Kenya, including 97 hominin footprints attributed to Homo erectus. These tracks are found in multiple sedimentary layers spanning approximately 20 thousand years. Taphonomic experiments show that each of these trackways represents minutes to no more than a few days in the lives of the individuals moving across these paleolandscapes. The geology and associated vertebrate fauna place these tracks in a deltaic setting, near a lakeshore bordered by open grasslands. Hominin footprints are disproportionately abundant in this lake margin environment, relative to hominin skeletal fossil frequency in the same deposits. Accounting for preservation bias, this abundance of hominin footprints indicates repeated use of lakeshore habitats by Homo erectus. Clusters of very large prints moving in the same direction further suggest these hominins traversed this lakeshore in multi-male groups. Such reliance on near water environments and possibly aquatic-linked foods, may have influenced hominin foraging behavior and migratory routes across and out of Africa.

Body size is a fundamental biological property of organisms, and documenting body size variation in hominin evolution is an important goal of palaeoanthropology. Estimating body mass appears deceptively simple but is laden with theoretical and pragmatic assumptions about best predictors and the most appropriate reference samples. Modern human training samples with known masses are arguably the ‘best’ for estimating size in early bipedal hominins such as the australopiths and all members of the genus Homo , but it is not clear if they are the most appropriate priors for reconstructing the size of the earliest putative hominins such as Orrorin and Ardipithecus . The trajectory of body size evolution in the early part of the human career is reviewed here and found to be complex and nonlinear. Australopith body size varies enormously across both space and time. The pre- erectus early Homo fossil record from Africa is poor and dominated by relatively small-bodied individuals, implying that the emergence of the genus Homo is probably not linked to an increase in body size or unprecedented increases in size variation. Body size differences alone cannot explain the observed variation in hominin body shape, especially when examined in the context of small fossil hominins and pygmy modern humans. This article is part of the themed issue ‘Major transitions in human evolution’.

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., Dietary adaptations of extinct early humans are often inferred from dental microwear data. Here, the authors employ mechanical analyses to show that Australopithecus sediba had limited ability to consume hard foods.

This book demonstrates how the primate hand combines both primitive and novel morphology, both general function with specialization, and both a remarkable degree of diversity within some clades and yet general similarity across many others. Across the chapters, different authors have addressed a variety of specific questions and provided their perspectives, but all explore the main themes described above to provide an overarching “primitive primate hand” thread to the book. Each chapter provides an in-depth review and critical account of the available literature, a balanced interpretation of the evidence from a variety of perspectives, and prospects for future research questions. In order to make this a useful resource for researchers at all levels, the basic structure of each chapter is the same, so that information can be easily consulted from chapter to chapter. An extensive reference list is provided at the end of each chapter so the reader has additional resources to address more specific questions or to find specific data.

Over the course of early hominin evolution, two fundamental changes in hand function occurred: the loss of a locomotor role and unparalleled intensification of manipulation, tool making, and tool use. In the context of these functional changes, early hominin hand anatomy evolved a number of derived characteristics within an otherwise primitive bauplan. Here we explore the functional significance and evolutionary history of seven major anatomical changes that make our hands distinctive, including hand proportions, thumb robusticity, thumb musculature, distal tuberosities, carpal architecture, wrist mobility, and finger curvature. This chapter highlights many areas that need more research and leads to several major conclusions: the abandonment of arboreal locomotion and rise in manipulative capabilities evolved over long periods of time and in a nonlinear fashion; early hominin taxa likely varied in their locomotor repertoires and manipulative abilities, not unlike differences in behavior seen among closely related species living today; and intensification of manipulation, rather than the origin of stone tool making, was a major driver of human hand evolution. Finally, we propose a new term, hyper-opposable, to describe the derived human ability to produce extensive contact area between the thumb and other digits, and forcefully secure and precision handle objects between the thumb and other digits through pad-to-pad contact.

Mechanical loads play a significant role in determining long bone shape and strength, but less work has explored how these loads influence flat bones like the scapula, which has been shown to vary with locomotor preference among primate taxa. Here, we tested the effects of voluntary running and climbing exercise in mice to examine how the mechanical loads borne from different locomotor patterns influence shoulder morphological development. Ninety-nine female wild-type mice were distributed equally among sedentary control, activity-wheel running, and vertical climbing experimental conditions. Running mice had the lowest body masses, larger intrinsic shoulder muscles, and the most pronounced differences in scapular size and shape relative to the other groups. Climbing mouse scapular morphology also differed significantly from the control individuals, but these differences were not as marked as those between the running and control mice. This might be attributable in part to greater levels of activity in the wheel-runners relative to the climbers. Additionally, climbing mice held their bodies closer to the substrate and maintained more flexed limbs and posterior hand positions compared with the kinematics of running. As a result, climbers differed significantly from both the running and control mice in developing a relatively broader infraspinous region, which is likely related to preferential recruitment of the infraspinatus and teres minor muscles to maintain flexed shoulder postures. The results of this study demonstrate that variation in activity level and type of locomotor regime over a significant portion of the life history influences muscle and bone development in the shoulder.

Recent studies of dental microwear and craniofacial mechanics have yielded contradictory interpretations regarding the feeding ecology and adaptations of Australopithecus africanus. As part of this debate, the methods used in the mechanical studies have been criticized. In particular, it has been claimed that finite element analysis has been poorly applied to this research question. This paper responds to some of these mechanical criticisms, highlights limitations of dental microwear analysis, and identifies avenues of future research.

Bipedalism is a defining feature of the human lineage. Despite evidence that walking on two feet dates back 6–7 Ma, reconstructing hominin gait evolution is complicated by a sparse fossil record and challenges in inferring biomechanical patterns from isolated and fragmentary bones. Similarly, patterns of social behavior that distinguish modern humans from other living primates likely played significant roles in our evolution, but it is exceedingly difficult to understand the social behaviors of fossil hominins directly from fossil data. Footprints preserve direct records of gait biomechanics and behavior but they have been rare in the early human fossil record. Here we present analyses of an unprecedented discovery of 1.5-million-year-old footprint assemblages, produced by 20+ Homo erectus individuals. These footprints provide the oldest direct evidence for modern human-like weight transfer and confirm the presence of an energy-saving longitudinally arched foot in H. erectus. Further, print size analyses suggest that these H. erectus individuals lived and moved in cooperative multi-male groups, offering direct evidence consistent with human-like social behaviors in H. erectus.

Mechanical loads, particularly those generated by skeletal muscle, play a significant role in determining long-bone shape and strength, but it is less clear how these loads influence the morphology of flat bones like the scapula. While scapular morphology has been shown to vary with locomotor mode in mammals, this study seeks to better understand whether genetically modified muscle size can influence scapular shape in the absence of significant locomotor differences. The soft- and hard-tissue morphological characteristics were examined in 11 hypermuscular, mutant (myostatin-deficient), 20 heterozygote, and 15 wild-type mouse shoulders. Body mass did not significantly differ among the genotype groups, but homozygous mutant and heterozygote mice had significantly larger shoulder muscles than wild-type mice. Mutant mice also differed significantly from the wild-type controls in several aspects of scapular size and shape, including glenohumeral joint orientation, total scapular length, superior border length, and supraspinous and infraspinous fossa length. Conversely, several traits describing superoinferior scapular breadth measures (e.g. total breadth and dorsal scapular fossa breadth) did not significantly differ between mutant and wild-type mice. Since the intrinsic musculature of the scapula is oriented in a mediolateral fashion, it follows that mediolaterally configured hard-tissue features like scapular length were most distinct among genotype groups. As had been noted previously with long bones, this study demonstrates that genetically enhanced muscle size has marked effects on the morphological characteristics of the shoulder.

Hypotheses regarding patterns of stress, strain and deformation in the craniofacial skeleton are central to adaptive explanations for the evolution of primate craniofacial form. The complexity of craniofacial skeletal morphology makes it difficult to evaluate these hypotheses with in vivo bone strain data. In this paper, new in vivo bone strain data from the intraorbital surfaces of the supraorbital torus, postorbital bar and postorbital septum, the anterior surface of the postorbital bar, and the anterior root of the zygoma are combined with published data from the supraorbital region and zygomatic arch to evaluate the validity of a finite-element model (FEM) of a macaque cranium during mastication. The behavior of this model is then used to test hypotheses regarding the overall deformation regime in the craniofacial haft of macaques. This FEM constitutes a hypothesis regarding deformation of the facial skeleton during mastication. A simplified verbal description of the deformation regime in the macaque FEM is as follows. Inferior bending and twisting of the zygomatic arches about a rostrocaudal axis exerts inferolaterally directed tensile forces on the lateral orbital wall, bending the wall and the supraorbital torus in frontal planes and bending and shearing the infraorbital region and anterior zygoma root in frontal planes. Similar deformation regimes also characterize the crania of Homo and Gorilla under in vitro loading conditions and may be shared among extant catarrhines. Relatively high strain magnitudes in the anterior root of the zygoma suggest that the morphology of this region may be important for resisting forces generated during feeding.

Past studies have hypothesized that aspects of hominin upper limb morphology are linked to the ability to produce stone tools. However, we lack the data on upper limb motions needed to evaluate the bio- mechanical context of stone tool production. This study seeks to better understand the biomechanics of stone tool-making by investigating upper limb joint kinemat- ics, focusing on the role of the wrist joint, during simple flake production. We test the hypotheses, based on stud- ies of other upper limb activities (e.g., throwing), that upper limb movements will occur in a proximal-to-distal sequence, culminating in rapid wrist flexion just prior to strike. Data were captured from four amateur knappers during simple flake production using a VICON motion analysis system (50 Hz). Results show that subjects uti- lized a proximal-to-distal joint sequence and disassoci- ated the shoulder joint from the elbow and wrist joints, suggesting a shared strategy employed in other contexts (e.g., throwing) to increase target accuracy. The knap- ping strategy included moving the wrist into peak exten- sion (subject peak grand mean 5 47.38) at the beginning of the downswing phase, which facilitated rapid wrist flexion and accelerated the hammerstone toward the nodule. This sequence resulted in the production of sig- nificantly more mechanical work, and therefore greater strike forces, than would otherwise be produced. To- gether these results represent a strategy for increasing knapping efficiency in Homo sapiens and point to aspects of skeletal anatomy that might be examined to assess potential knapping ability and efficiency in fossil hominin taxa. Am J Phys Anthropol 143:134-145, 2010. V

Australopithecus africanus is an early hominin (i.e., human relative) believed to exhibit stress-reducing adaptations in its craniofacial skeleton that may be related to the consumption of resistant food items using its premolar teeth. Finite element analyses simulating molar and premolar biting were used to test the hypothesis that the cranium of A. africanus is structurally more rigid than that of Macaca fascicularis, an Old World monkey that lacks derived australopith facial features. Previously generated finite element models of crania of these species were subjected to isometrically scaled loads, permitting a direct comparison of strain magnitudes. Moreover, strain energy (SE) in the models was compared after results were scaled to account for differences in bone volume and muscle forces. Results indicate that strains in certain skeletal regions below the orbits are higher in M. fascicularis than in A. africanus. Moreover, although premolar bites produce von Mises strains in the rostrum that are elevated relative to those produced by molar biting in both species, rostral strains are much higher in the macaque than in the australopith. These data suggest that at least the midface of A. africanus is more rigid than that of M. fascicularis. Comparisons of SE reveal that the A. africanus cranium is, overall, more rigid than that of M. fascicularis during premolar biting. This is consistent with the hypothesis that this hominin may have periodically consumed large, hard food items. However, the SE data suggest that the A. africanus cranium is marginally less rigid than that of the macaque during molar biting. It is hypothesized that the SE results are being influenced by the allometric scaling of cranial cortical bone thickness.

Hominin footprints offer evidence about gait and foot shape, but their scarcity, combined with an inadequate hominin fossil record, hampers research on the evolution of the human gait. Here, we report hominin footprints in two sedimentary layers dated at 1.51 to 1.53 million years ago (Ma) at Ileret, Kenya, providing the oldest evidence of an essentially modern human–like foot anatomy, with a relatively adducted hallux, medial longitudinal arch, and medial weight transfer before push-off. The size of the Ileret footprints is consistent with stature and body mass estimates for Homo ergaster/erectus , and these prints are also morphologically distinct from the 3.75-million-year-old footprints at Laetoli, Tanzania. The Ileret prints show that by 1.5 Ma, hominins had evolved an essentially modern human foot function and style of bipedal locomotion.

The African Plio-Pleistocene hominins known as australopiths evolved a distinctive craniofacial morphology that traditionally has been viewed as a dietary adaptation for feeding on either small, hard objects or on large volumes of food. A historically influential interpretation of this morphology hypothesizes that loads applied to the premolars during feeding had a profound influence on the evolution of australopith craniofacial form. Here, we test this hypothesis using finite element analysis in conjunction with comparative, imaging, and experimental methods. We find that the facial skeleton of the Australopithecus type species, A. africanus , is well suited to withstand premolar loads. However, we suggest that the mastication of either small objects or large volumes of food is unlikely to fully explain the evolution of facial form in this species. Rather, key aspects of australopith craniofacial morphology are more likely to be related to the ingestion and initial preparation of large, mechanically protected food objects like large nuts and seeds. These foods may have broadened the diet of these hominins, possibly by being critical resources that australopiths relied on during periods when their preferred dietary items were in short supply. Our analysis reconciles apparent discrepancies between dietary reconstructions based on biomechanics, tooth morphology, and dental microwear.

There is considerable debate over the level of size dimorphism and inferred social behavior of Australopithecus afarensis. Most previous studies have analyzed size variation in single variables or multiple variables drawn from single elements. These approaches suffer from small sample sizes, underscoring the need for new techniques that incorporate measurements from multiple unassociated elements, reducing the influence of random sampling on size variation in fossil samples. One such technique, the template method, has recently been proposed but is limited to samples with a template specimen and is sensitive to a number of assumptions. Here we present two new resampling methods that do not require a template specimen, allow measurements from multiple unassociated elements to be included in a single analysis, and allow for significance tests between comparative and fossil multivariate data sets with miss- ing data. Using these new methods, multivariate post- cranial size dimorphism is measured using eight mea- surements of the femur, tibia, humerus, and radius in samples of A. afarensis, modern humans, chimpanzees, gorillas, and orangutans. Postcranial dimorphism in A. afarensis is similar to that of gorillas and orangutans, and significantly greater than in modern humans and chimpanzees. Because studies in living primates have examined the association of behavior with dimorphism in body mass and craniodental measurements, not post- crania, relationships between postcranial dimorphism and social behavior must be established to make robust behavioral inferences for A. afarensis. However, the results of this and past studies strongly suggest behav- ioral and mating strategies differed between A. afarensis and modern humans. Am J Phys Anthropol 135:311-328, 2008. V C 2007 Wiley-Liss, Inc.

Phalangeal curvature has been widely cited in primate functional morphology and is one of the key traits in the ongoing debate about whether the locomotion of early hominins included a significant degree of arboreality. This study examines the biomechanics of phalangeal curvature using data on hand posture, muscle recruitment, and anatomical moment arms to develop a finite element (FE) model of a siamang manual proximal phalanx during suspensory grasping. Strain patterns from experiments on intact cadaver forelimbs validated the model. The strain distribution in the curved siamang phalanx FE model was compared to that in a mathematically straight rendition in order to test the hypotheses that curvature: 1) reduces strain and 2) results in lower bending strains but relatively higher compression. In the suspensory posture, joint reaction forces load the articular ends of the phalanx in compression and dorsally, while muscle forces acting through the flexor sheath pull the mid-shaft palmarly. These forces compress the phalanx dorsally and tense it palmarly, effectively bending it 'open.' Strains in the curved model were roughly half that of the straight model despite equivalent lengths, areas, mechanical properties, and loading conditions in the two models. The curved model also experienced a higher ratio of compressive to tensile strains. Curvature reduces strains during grasping hand postures because the curved bone is more closely aligned with the joint reaction forces. Therefore, phalangeal curvature reduces the strains associated with arboreal, and especially suspensory, activity involving flexed digits. These results offer a biomechanical explanation for the observed association between phalangeal curvature and arboreality.

The zygomatic region and lateral orbital wall are important structural components of the primate craniofacial skeleton, providing attachment for muscles, housing and protecting the eye, and resisti...

Humans are unique, compared with our closest living relatives (chimpanzees) and early fossil hominins, in having an enlarged body size and lower limb joint surfaces in combination with a relatively gracile skeleton (i.e., lower bone mass for our body size). Some analyses have observed that in at least a few anatomical regions modern humans today appear to have relatively low trabecular density, but little is known about how that density varies throughout the human skeleton and across species or how and when the present trabecular patterns emerged over the course of human evolution. Here, we test the hypotheses that (i) recent modern humans have low trabecular density throughout the upper and lower limbs compared with other primate taxa and (ii) the reduction in trabecular density first occurred in early Homo erectus, consistent with the shift toward a modern human locomotor anatomy, or more recently in concert with diaphyseal gracilization in Holocene humans. We used peripheral quantitative CT and microtomography to measure trabecular bone of limb epiphyses (long bone articular ends) in modern humans and chimpanzees and in fossil hominins attributed to Australopithecus africanus, Paranthropus robustus/early Homo from Swartkrans, Homo neanderthalensis, and early Homo sapiens. Results show that only recent modern humans have low trabecular density throughout the limb joints. Extinct hominins, including pre-Holocene Homo sapiens, retain the high levels seen in nonhuman primates. Thus, the low trabecular density of the recent modern human skeleton evolved late in our evolutionary history, potentially resulting from increased sedentism and reliance on technological and cultural innovations.

Afro-Arabian mammalian communities underwent a marked transition near the Oligocene/Miocene boundary at approximately 24 million years (Myr) ago. Although it is well documented that the endemic paenungulate taxa were replaced by migrants from the Northern Hemisphere, the timing and evolutionary dynamics of this transition have long been a mystery because faunas from about 32 to 24 Myr ago are largely unknown. Here we report a late Oligocene fossil assemblage from Ethiopia, which constrains the migration to postdate 27 Myr ago, and yields new insight into the indigenous faunal dynamics that preceded this event. The fauna is composed of large paenungulate herbivores and reveals not only which earlier taxa persisted into the late Oligocene epoch but also demonstrates that one group, the Proboscidea, underwent a marked diversification. When Eurasian immigrants entered Afro-Arabia, a pattern of winners and losers among the endemics emerged: less diverse taxa such as arsinoitheres became extinct, moderately species-rich groups such as hyracoids continued into the Miocene with reduced diversity, whereas the proboscideans successfully carried their adaptive radiation out of Afro-Arabia and across the world.

Using information from primatology, functional morphology, phylogeny, archeology, and paleoanthropology, we argue that before 2.5 mya hominins may have used tools, including unmodified and possibly modified stone tools (Fig. 1). We consider several scenarios to explain why stone tool manufacture and use might not have left archeological traces prior to 2.5 mya and conclude by suggesting means to test our hypotheses.

Recent analyses and new fossil discoveries suggest that the evolution of hominin limb length proportions is complex, with evolutionary reversals and a decoupling of proportions within and between limbs. This study takes into account intraspecific variation to test whether or not the limb proportions of four early hominin associated skeletons (AL 288-1, OH 62, BOU-VP-12/1, and KNM-WT 15000) can be considered to be significantly different from one another. Exact randomization methods were used to compare the differences between pairs of fossil skeletons to the differences observed between all possible pairs of individuals within large samples of Gorilla gorilla, Pan troglodytes, Pongo pygmaeus, and Homo sapiens. Although the difference in humerofemoral proportions between OH 62 and AL 288-1 does not exceed variation in the extant samples, it is rare. When humerofemoral midshaft circumferences are compared, the difference between OH 62 and AL 288-1 is fairly common in extant species. This, in combination with error associated with the limb lengths estimates, suggests that it may be premature to consider H. (or Australopithecus) habilis as having more apelike limb proportions than those in A. afarensis. The humerofemoral index of BOU-VP-12/1 differs significantly from both OH 62 and AL 288-1, but not from KNM-WT 15000. Published length estimates, if correct, suggest that the relative forearm length of BOU-VP-12/1 is unique among hominins, exceeding those of the African apes and resembling the proportions in Pongo. Evidence that A. afarensis exhibited a less apelike upper:lower limb design than A. africanus (and possibly H. habilis) suggests that, if A. afarensis is broadly ancestral to A. africanus, the latter did not simply inherit primitive morphology associated with arboreality, but is derived in this regard. The fact that the limb proportions of OH 62 (and possibly KNM-ER 3735) are no more human like than those of AL 288-1 underscores the primitive body design of H. habilis.

The African Plio-Pleistocene hominins known as australopiths evolved derived craniodental features frequently interpreted as adaptations for feeding on either hard, or compliant/tough foods. Among australopiths, Paranthropus boisei is the most robust form, exhibiting traits traditionally hypothesized to produce high bite forces efficiently and strengthen the face against feeding stresses. However, recent mechanical analyses imply that P. boisei may not have been an efficient producer of bite force and that robust morphology in primates is not necessarily strong. Here we use an engineering method, finite element analysis, to show that the facial skeleton of P. boisei is structurally strong, exhibits a strain pattern different from that in chimpanzees (Pan troglodytes) and Australopithecus africanus, and efficiently produces high bite force. It has been suggested that P. boisei consumed a diet of compliant/tough foods like grass blades and sedge pith. However, the blunt occlusal topography of this and other species suggests that australopiths are adapted to consume hard foods, perhaps including grass and sedge seeds. A consideration of evolutionary trends in morphology relating to feeding mechanics suggests that food processing behaviors in gracile australopiths evidently were disrupted by environmental change, perhaps contributing to the eventual evolution of Homo and Paranthropus.

In a broad range of evolutionary studies, an understanding of intraspecific variation is needed in order to contextualize and interpret the meaning of variation between species. However, mechanical analyses of primate crania using experimental or modeling methods typically encounter logistical constraints that force them to rely on data gathered from only one or a few individuals. This results in a lack of knowledge concerning the mechanical significance of intraspecific shape variation that limits our ability to infer the significance of interspecific differences. This study uses geometric morphometric methods (GM) and finite element analysis (FEA) to examine the biomechanical implications of shape variation in chimpanzee crania, thereby providing a comparative context in which to interpret shape-related mechanical variation between hominin species. Six finite element models (FEMs) of chimpanzee crania were constructed from CT scans following shape-space Principal Component Analysis (PCA) of a matrix of 709 Procrustes coordinates (digitized onto 21 specimens) to identify the individuals at the extremes of the first three principal components. The FEMs were assigned the material properties of bone and were loaded and constrained to simulate maximal bites on the P(3) and M(2) . Resulting strains indicate that intraspecific cranial variation in morphology is associated with quantitatively high levels of variation in strain magnitudes, but qualitatively little variation in the distribution of strain concentrations. Thus, interspecific comparisons should include considerations of the spatial patterning of strains rather than focus only on their magnitudes.