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The earliest South African hominids (humans and their ancestral kin) belong to the genera Australopithecus, Paranthropus, and Homo, with the oldest being a ca. 3.67 million-year-old nearly complete skeleton of Australopithecus (StW 573) from Sterkfontein Caves. This skeleton has provided, for the first time in almost a century of research, the full anatomy of an Australopithecus individual with indisputably associated skull and postcranial bones that give complete limb lengths. The three genera are also found in East Africa, but scholars have disagreed on the taxonomic assignment for some fossils owing to historical preconceptions. Here we focus on the South African representatives to help clarify these debates. The uncovering of the StW 573 skeleton in situ revealed significant clues concerning events that had affected it over time and demonstrated that the associated stalagmite flowstones cannot provide direct dating of the fossil, as they are infillings of voids caused by postdepositional collapse.

The StW 573 skeleton of Australopithecus prometheus from Sterkfontein Member 2 is some 93% complete and thus by far the most complete member of that genus yet found. Firmly dated at 3.67 Ma, it is one of the earliest specimens of its genus. A crucial aspect of interpretation of locomotor behaviour from fossil remains is an understanding of the palaeoenvironment in which the individual lived and the manner in which it would have used it. While the value of this ecomorphological approach is largely accepted, it has not been widely used as a stable framework on which to build evolutionary biomechanical interpretations. Here, we collate the available evidence on StW 573’s anatomy in order, as far as currently possible, to reconstruct what might have been this individual’s realized and potential niche. We explore the concept of a common Australopithecus “bauplan” by comparing the morphology and ecological context of StW 573 to that of paenocontemporaneous australopiths including Australopithecus anamensis and KSD-VP-1/1 Australopithecus afarensis. Each was probably substantially arboreal and woodland-dwelling, relying substantially on arboreal resources. We use a hypothesis-driven approach, tested by: virtual experiments, in the case of extinct species; biomechanical analyses of the locomotor behaviour of living great ape species; and analogical experiments with human subjects. From these, we conclude that the habitual locomotor mode of all australopiths was upright bipedalism, whether on the ground or on branches. Some later australopiths such as Australopithecus sediba undoubtedly became more terrestrial, allowing sacrifice of arboreal stability in favour of manual dexterity. Indeed, modern humans retain arboreal climbing skills but have further sacrificed arboreal effectiveness for enhanced ability to sustain striding terrestrial bipedalism over much greater distances. We compare StW 573’s locomotor adaptations to those of living great apes and protohominins, and agree with those earlier observers who suggest that the common panin-hominin last common ancestor was postcranially more like Gorilla than Pan.

Functional morphology of the atlas reflects multiple aspects of an organism’s biology. More specifically, its shape indicates patterns of head mobility, while the size of its vascular foramina reflects blood flow to the brain. Anatomy and function of the early hominin atlas, and thus, its evolutionary history, are poorly documented because of a paucity of fossilized material. Meticulous excavation, cleaning and high-resolution micro-CT scanning of the StW 573 (‘Little Foot’) skull has revealed the most complete early hominin atlas yet found, having been cemented by breccia in its displaced and flipped over position on the cranial base anterolateral to the foramen magnum. Description and landmark-free morphometric analyses of the StW 573 atlas, along with other less complete hominin atlases from Sterkfontein (StW 679) and Hadar (AL 333-83), confirm the presence of an arboreal component in the positional repertoire of Australopithecus. Finally, assessment of the cross-sectional areas of the transverse foramina of the atlas and the left carotid canal in StW 573 further suggests there may have been lower metabolic costs for cerebral tissues in this hominin than have been attributed to extant humans and may support the idea that blood perfusion of these tissues increased over the course of hominin evolution.

The ca. 3.67 Ma adult skeleton known as ‘Little Foot’ (StW 573), recovered from Sterkfontein Member 2 breccia in the Silberberg Grotto, is remarkable for its morphology and completeness. Preservation of clavicles and scapulae, including essentially complete right-side elements, offers opportunities to assess morphological and functional aspects of a nearly complete Australopithecus pectoral girdle. Here we describe the StW 573 pectoral girdle and offer quantitative comparisons to those of extant hominoids and selected homininans. The StW 573 pectoral girdle combines features intermediate between those of humans and other apes: a long and curved clavicle, suggesting a relatively dorsally positioned scapula; an enlarged and uniquely proportioned supraspinous fossa; a relatively cranially oriented glenoid fossa; and ape-like reinforcement of the axillary margin by a stout ventral bar. StW 573 scapulae are as follows: smaller than those of some homininans (i.e., KSD-VP-1/1 and KNM-ER 47000A), larger than others (i.e., A.L. 288-1, Sts 7, and MH2), and most similar in size to another australopith from Sterkfontein, StW 431. Moreover, StW 573 and StW 431 exhibit similar structural features along their axillary margins and inferior angles. As the StW 573 pectoral girdle (e.g., scapular configuration) has a greater affinity to that of apes—Gorilla in particular—rather than modern humans, we suggest that the StW 573 morphological pattern appears to reflect adaptations to arboreal behaviors, especially those with the hand positioned above the head, more than human-like manipulatory capabilities. When compared with less complete pectoral girdles from middle/late Miocene apes and that of the penecontemporaneous KSD-VP-1/1 (Australopithecus afarensis), and mindful of consensus views on the adaptiveness of arboreal positional behaviors soliciting abducted glenohumeral joints in early Pliocene taxa, we propose that the StW 573 pectoral girdle is a reasonable model for hypothesizing pectoral girdle configuration of the crown hominin last common ancestor.

The Early Pleistocene site of Swartkrans in South Africa's Cradle of Humankind World Heritage Site has been significant for our understanding of the evolution of both early Homo and Paranthropus, as well as the earliest archaeology of southern Africa. Previous attempts to improve a faunal age estimate of the earliest deposit, Member 1, had produced results obtained with uranium-lead dating (U-Pb) on flowstones and cosmogenic burial dating of quartz, which placed the entire member in the range of1.7/1.8 Ma and2.3 Ma. In 2014, two simple burial dates for the Lower Bank, the earliest unit within Member 1, narrowed its age to between ca. 1.8 Ma and 2.2 Ma. A new dating program using the isochron method for burial dating has established an absolute age of 2.22 ± 0.09 Ma for a large portion of the Lower Bank, which can now be identified as containing the earliest Oldowan stone tools and fossils of Paranthropus robustus in South Africa. This date agrees within one sigma with the U-Pb age of 2.25 ± 0.08 Ma previously published for the flowstone underlying the Lower Bank and confirms a relatively rapid rate of accumulation for a large portion of the talus.

International audience; Because of its exceptional degree of preservation and its geological age of ∼3.67 Ma, StW 573 makes an invaluable contribution to our understanding of early hominin evolution and paleobiology. The morphology of the bony labyrinth has the potential to provide information about extinct primate taxonomic diversity, phylogenetic relationships and locomotor behaviour. In this context, we virtually reconstruct and comparatively assess the bony labyrinth morphology in StW 573. As comparative material, we investigate 17 southern African hominin specimens from Sterkfontein, Swartkrans and Makapansgat (plus published data from two specimens from Kromdraai B), attributed to Australopithecus, early Homo or Paranthropus, as well as 10 extant human and 10 extant chimpanzee specimens. We apply a landmark-based geometric morphometric method for quantitatively assessing labyrinthine morphology. Morphology of the inner ear in StW 573 most closely resembles that of another Australopithecus individual from Sterkfontein, StW 578, recovered from the Jacovec Cavern. Within the limits of our sample, we observe a certain degree of morphological variation in the Australopithecus assemblage of Sterkfontein Member 4. Cochlear morphology in StW 573 is similar to that of other Australopithecus as well as to Paranthropus specimens included in this study, but it is substantially different from early Homo. Interestingly, the configuration of semicircular canals in Paranthropus specimens from Swartkrans differs from other fossil hominins, including StW 573. Given the role of the cochlea in the sensory-driven interactions with the surrounding environment, our results offer new perspectives for interpreting early hominin behaviour and ecology. Finally, our study provides additional evidence for discussing the phylogenetic polarity of labyrinthine traits in southern African hominins.

OBJECTIVES We provide descriptions and functional interpretations of 11 >2.0 Ma hominin vertebral and upper limb fossils from Sterkfontein. MATERIALS AND METHODS We employed taphonomic methods to describe postmortem damage observed on the fossils. We used osteometric tools and measurements to generate quantitative descriptions, which were added to qualitative descriptions of the fossils. These observations were then interpreted using published data on the same skeletal elements from extant and extinct hominoid taxa. RESULTS Six of the fossils carry carnivore tooth marks. Two vertebrae show morphologies that are consistent with fully developed lordosis of the lumbar spine, but which are not completely consistent with bipedal loading of the same intensity and/or frequency as reflected in the lumbars of modern humans. A clavicle shows a combination of humanlike and apelike features, the latter of which would have endowed its hominin with good climbing abilities. When combined, analyses of fragmentary radius and ulna fossils yield more ambiguous results. DISCUSSION The new fossil collection presents a mix of bipedal and climbing features. It is unclear whether this mix indicates that all Sterkfontein hominins of >2.0 Ma were terrestrial bipeds who retained adaptations for climbing or whether the collection samples two differently adapted, coeval hominins, Australopithecus africanus and Australopithecus prometheus, both of which are represented at Sterkfontein by skull remains. Regardless, the significant frequency of tooth-marked fossils in the sample might indicate that predation was a selection pressure that maintained climbing adaptations in at least some Sterkfontein hominins of this period.

International audience; Reconstruction of the locomotor repertoire of the australopiths (Australopithecus and Paranthropus) has progressively integrated information from the mechanosensitive internal structure of the appendicular skeleton. Recent investigations showed that the arrangement of the trabecular network at the femoral head center is biomechanically compatible with the pattern of cortical bone distribution across the neck, both suggesting a full commitment to bipedalism in australopiths, but associated with a slightly altered gait kinematics compared to Homo involving more lateral deviation of the body center of mass over the stance limb. To provide a global picture in Paranthropus robustus of the trabecular architecture of the proximal femur across the head, neck and greater trochanter compartments, we applied techniques of virtual imaging to the variably preserved Early Pleistocene specimens SK 82, SK 97, SK 3121, SKW 19 and SWT1/LB-2 from the cave site of Swartkrans, South Africa. We also assessed the coherence between the structural signals from the center of the head and those from the trabecular network of the inferolateral portion of the head and the inferior margin of the neck, sampling the so-called vertical bundle, which in humans represents the principal compressive system of the joint. Our analyses show a functionally related trabecular organization in Pa. robustus that closely resembles the extant human condition, but which also includes some specificities in local textural arrangement. The network of the inferolateral portion of the head shows a humanlike degree of anisotropy and a bone volume fraction intermediate between the extant human and the African ape patterns. These results suggest slight differences in gait kinematics between Pa. robustus and extant humans. The neck portion of the vertical bundle revealed a less biomechanically sensitive signal. Future investigations on the australopith hip joint loading environment should more carefully investigate the trabecular structure of the trochanteric region and possible structural covariation between cortical bone distribution across the neck and site-specific trabecular properties of the arcuate bundle.

We describe late Pliocene and early Pleistocene hominin fossils from Sterkfontein Caves (South Africa), including two femoral specimens, as well as a partial tibia and a partial fibula. The fossils are likely assignable to Australopithecus africanus and/or Australopithecus prometheus and the morphology of each corroborates previous interpretations of Sterkfontein hominins as at least facultative bipeds. Significance: A recent series of papers by our research team describes the morphology of a hominin skeleton from Sterkfontein Caves (South Africa), nicknamed ‘Little Foot’. Based on its unique skull morphology, R.J. Clarke, the skeleton’s discoverer, places it in the species Australopithecus prometheus, as distinct from the better-known and co-occurring Australopithecus africanus. Here we describe additional hominin thigh and leg fossils from Sterkfontein that, when considered in a comparative context, support the hypothesis that there was significant (probably interspecific) variation in South African hominin postcranial morphology during the late Pliocene and early Pleistocene.

The reconstruction of perimortem and postmortem events is of critical importance to criminal investigations. In many cases, the information required for these reconstructions can be accessed through the analysis of skeletal remains. One particular class of skeletal data—trauma to the surfaces of bones, or bone surface modifications (BSMs)—can reveal much about the perimortem and postmortem intervals. While the study of BSMs originated within the fields of paleontology and archeology and was only later integrated into forensic science, a fruitful interdisciplinary exchange of data and methods is now commonplace. BSMs from thermal alteration, sharp-force trauma, terrestrial and aquatic scavengers and predators, bacteria and fungi, insects, weathering, and sediment abrasion can supply investigators with valuable information about the agents and events of a corpse's deposition, including weapon type, local environmental conditions, the postmortem interval, and the presence, temperature(s), and/or length(s) of thermal exposure. Based on a review of this rich body of literature, we argue that (a) all associations between a BSM and its alleged source must rest on observational cause-and-effect studies; (b) secure identifications of BSMs should rely both on the intrinsic features of the modifications themselves and relevant contextual data; (c) the scientific validity of BSM research depends, ultimately, on rigorous blind-testing and the establishment of error rates; and (d) researchers need to make a concerted effort to enhance interanalyst correspondence through objective definitions, measurements, and/or codes of BSM features. The most promising path forward lies in the combination of digital image analysis and multivariate predictive modeling.

Pickering et al. (Nature 2018;565:226–229) utilised calcium carbonate flowstone deposits (i.e. speleothems) from eight Pliocene and Pleistocene South African Cradle of Humankind cave sites to propose that biases were created within the fossil record due to absent clastic sedimentation phases during wet periods, when caves were closed and only speleothems accumulated. Such a scenario has significant implications for our understanding of variability in hominin mobility, resource exploitation, functional repertoires and interactions with competitors in changing environmental and ecological contexts. We find considerable issues with the article. First, Pickering et al.’s contribution omits crucial fossil evidence from various stratigraphic units of the Sterkfontein Caves that indicates conditions were not always arid when the caves were open and sediments were deposited. Second, Pickering et al.’s proposal that clastic and speleothemic deposits (including faunal and floral material) form mutually exclusively is an overly simplified, binary depositional (and in this case environmental) framework that demonstrates an inherent bias in the sampling of cave deposits for dating. This creates the impression that either speleothems or clastic sediments are deposited and does not take into account the full spectrum of sedimentary complexity in karst caves. Third, closure of the caves across the Cradle of Humankind landscape during wet periods is not substantiated geomorphologically or speleologically; identification of the responsible process is critical to the proposed infilling scenario. Significance: We propose that Pickering et al.’s interpretation of the environmental context of the South African early hominin record is problematic in that it omits crucial faunal and floral fossil evidence associating hominins with non-arid climates, is geomorphologically unsupported, and perpetuates biases against temporally and climatically representative clastic sediments due to challenges related to their dating.

Natural selection, as both a process and a scientific concept, is eloquently simple. Unfortunately, this simplicity sometimes belies Darwin’s broader view of evolution as a multifaceted process that proceeds from both ecological pressures and phylogenetic history. Darwin further understood that it is not just physical traits that are transmitted generationally, but also behavioural patterns, both of which are subject to the shaping influences of environment and phylogeny. Chimpanzees, bonobos and humans are the most carnivorous extant primates, an observation that serves as the basis of our extended argument that vertebrate predation is a synapomorphy of these sister taxa. From there, we use archaeological data to trace the inferred polarity of hominin carcass foraging and meat-eating from their first archaeological indications ∼2.6 million years ago (Mya). A review of the early Pleistocene African record demonstrates that taphonomic evidence of a hominin predatory/meat-eating behavioral module cl...

The site of the Sterkfontein Caves, South Africa, is one of the richest early hominin fossil-bearing sites in Africa. Recent excavations in the Milner Hall locality have contributed to the discovery of new hominin specimens, including StW 669, a right permanent maxillary first molar (M1). StW 669 was excavated from the T1 deposits, which consist of a mixture of sediments from Members 2 and 5 of the Sterkfontein Formation. Accordingly, the deposits have the potential to contain remains of Australopithecus, Paranthropus and Homo. In this study, we employed micro-focus X-ray tomography in order to assess dental tissue proportions, enamel thickness distribution and enamel-dentine junction morphology as approaches to investigate the taxonomy of StW 669. We compare our results to those generated on the teeth of Australopithecus africanus, Paranthropus robustus, Homo erectus, Homo antecessor, Homo neanderthalensis and Homo sapiens. Our results suggest that StW 669 shares quantitative and qualitative affinities with M1s of Homo in terms of tissue proportions (i.e. two- and three-dimensional average and relative enamel thickness of 1.2–1.3 mm and 18.4, respectively) and enamel thickness distribution (i.e. thickest enamel on the lingual aspect of the protocone). However, data on the enamel-dentine junction morphology of StW 669 are inconclusive as to the tooth’s taxonomic affinities. Pending additional morphometric analyses, our studies of inner morphology of the crown of StW 669 support its attribution to Homo. Significance: The Sterkfontein Caves have contributed significantly to our understanding of early human evolution and continue to do so. This study highlights the specific value of the Milner Hall locality as a valuable store of hominin fossils. Moreover, we tentatively clarify the enigmatic taxonomic status of StW 669, a right permanent maxillary first molar, excavated from the T1 deposit of Milner Hall. Pending additional morphometric evidence, our preliminary data on tissue proportions, enamel thickness distribution and enamel-dentine junction morphology, suggest an attribution of StW 669 to early Homo. This result is significant given the historical contention concerning the presence of early Homo at Sterkfontein.

International audience; The Sterkfontein Caves is currently the world's richest Australopithecus-bearing site. Included in Sterkfontein's hominin assemblage is StW 573 (‘Little Foot’), a near-complete Australopithecus skeleton discovered in Member 2 in the Silberberg Grotto. Because of its importance to the fossil hominin record, the geological age of StW 573 has been the subject of significant debate. Three main hypotheses have been proposed regarding the formation and age of Member 2 and by association StW 573. The first proposes that Member 2 (as originally defined in the type section in the Silberberg Grotto) started to accumulate at around 2.58 Ma and that the unit is contained within the Silberberg Grotto. The second proposes that Member 2 started forming before 3.67 ± 0.16 Ma and that the deposit extends into the Milner Hall and close to the base of the cave system. The third proposes a ‘two-stage burial scenario’, in which some sediments and StW 573 represent a secondary and mixed-age accumulation reworked from a higher cave. The stratigraphic and sedimentological implications of these hypotheses are tested here through the application of a multiscale investigation of Member 2, with reference to the taphonomy of the StW 573 skeleton. The complete infilling sequence of Member 2 is described across all exposures of the deposit in the Silberberg Grotto and into the Milner Hall. Sediments are generally stratified and conformably deposited in a sequence of silty sands eroded from well-developed lateritic soils on the landscape surface. Voids, clasts and bioclasts are organized consistently across and through Member 2 conforming with the underlying deposit geometry, indicating gradual deposit accretion with no distinct collapse facies evident and only localized intra-unit postdepositional modification. The stratigraphy and sedimentology of Member 2 support a simple single-stage accumulation process of Member 2 and a primary association between the sediments of Member 2 and the StW 573 ‘Little Foot’ skeleton.

It is hypothesized that tool-assisted excavation of plant underground storage organs (USOs) played an adaptive role in hominin evolution and was also once considered a uniquely human behavior. Recent data indicate that savanna chimpanzees also use tools to excavate edible USOs. However, those chimpanzees remain largely unhabituated and we lack direct observations of this behavior in the wild. To fill this gap in our knowledge of hominoid USO extractive foraging, we conducted tool-mediated excavation experiments with captive chimpanzees naive to this behavior. We presented the chimpanzees with the opportunity to use tools in order to excavate artificially-placed underground foods in their naturally forested outdoor enclosure. No guidance or demonstration was given to the chimpanzees at any time. The chimpanzees used tools spontaneously in order to excavate the underground foods. They exhibited six different tool use behaviors in the context of excavation: probe, perforate, dig, pound, enlarge and shovel. However, they still excavated manually more often than they did with tools. Chimpanzees were selective in their choice of tools that we provided, preferring longer tools for excavation. They also obtained their own tools mainly from naturally occurring vegetation and transported them to the excavation site. They reused some tools throughout the study. Our new data provide a direction for the study of variables relevant to modeling USO extractive foraging by early hominins. Funding Agencies|La Caixa Foundation Spain [LCF/BQ/EU15/10350002]; University of Oslo, Department of Biosciences, Norway

The 3.67-million-year-old StW 573 Australopithecus skeleton is important for the light it sheds on the paleobiology of South African species of that genus, including, as discussed here, how the possible pathology of the specimen informs our understanding of Australopithecus behavior. The StW 573 antebrachium exhibits bilateral asymmetry, with significantly more longitudinally curved left forearm bones than right. Arguing from a comparative perspective, we hypothesize that these curvatures resulted from a fall onto a hyperextended, outstretched hand. It is unlikely that the fall was from a significant height and might have occurred when the StW 573 individual was a juvenile. This type of plastic deformation of the forearm bones is well-documented in modern human clinical studies, especially among children between the ages of four and ten years who tumble from bicycles or suffer other common, relatively low-impact accidents. Left untreated, such injuries impinge normal supination and pronation of the hand, a condition that could have had significant behavioral impact on the StW 573 individual.

(300 words)StW 573, from Sterkfontein Member 2, dated ca 3.67 Ma, is by far the most complete skeleton of an australopith to date. Joint morphology is in many cases closely matched in available elements of Australopithecus anamensis (eg. proximal and distal tibial and humeral joint-surfaces) and there are also close similarities to features of the scapula, in particular, of KSD-VP-1/1 A. afarensis from Woranso-Mille. The closest similarities are, however, to the partial skeleton of StW 431 from Sterkfontein Member 4. When considered together, both StW 573 and StW 431 express an hip joint morphology quite distinct from that of A. africanus Sts14, and a proximal femur of a presumed A. africanus from Jacovec Cavern at Sterkfontein, StW 598. This, and other evidence presented herein, suggests there are two pelvic girdle morphs at Sterkfontein, supporting Clarke (2013) in his recognition of a second species, A. prometheus, containing StW 573 and StW 431. StW 573 is the first hominid skeleton where limb proportions are known unequivocally. It demonstrates that some early hominins, at the time of formation of the Laetoli footprints (3.6 Ma), were large-bodied. with hindlimbs longer than forelimbs. Modelling studies on extant primates indicate that the intermembral index (IMI) of StW 573, low for a non-human great ape, would have substantially enhanced economy of bipedal walking over medium-to-long distances, but that it was still too high for effective walking while load-carrying. It would, however, have somewhat reduced the economy of horizontal climbing, but made Gorilla-like embracing of large tree-trunks less possible. Consideration of both ethnographic evidence from modern indigenous arboreal foragers and modern degeneracy theory cautions against prescriptive interpretations of hand- and foot-function, by confirming that both human-like upright bipedalism and functional capabilities of the hand and foot can be effective in short-distance arboreal locomotion.

The Sterkfontein Caves has an 80 year history of yielding remarkable evidence of hominin evolution and is currently the world’s richestAustralopithecus-beafingsite. Included in Sterkfontein’s hominin assemblage is StW 573 (‘Little Foot’). Discovered in the Member 2 deposit in the Silberberg Grotto, StW 573 represents the most completeAustralopithecusskeleton yet found. Because of its importance to the fossil hominin record, the geological age of Little Foot has been the subject of significant debate. Two main hypotheses have been proposed regarding the formation and age of Member 2 and by association StW 573. The first, proposes that Member 2 formed relatively rapidly, starting to accumulate at around 2.8 million years ago and that the unit is isolated to the Silberberg Grotto - the underlying chambers and passages forming later. The second proposes that Member 2 formed slowly, its accumulation starting before 3.67 million years ago and that the deposit extends into the Milner Hall and close to the base of the cave system. Both assume a primary association between StW 573 and Member 2, although which sediments in the Silberberg Grotto are associated with Member 2 has also been questioned. Recently a third alternative hypothesis questioning the association of StW 573 to Member 2 sediments proposed a ‘two-stage burial scenario’ in which sediments associated with StW 573 represent a secondary and mixed-age deposit reworked from a higher cave. The stratigraphic and sedimentological implications of these hypotheses are tested here through the application of a multiscale investigation of Member 2, with reference to the taphonomy of the Little Foot skeleton. The complete infilling sequence of Member 2 is described and depositional units are tracked across all exposures of the deposit in the Silberberg Grotto and into the Milner Hall. Facies development follows patterns characteristic of colluvially accumulated taluses with 30-40° angles of repose developing coarser distal facies. Sediments are generally stratified and conformably deposited in a sequence of silty sands eroded from well-developed lateritic soils on the landscape surface. Voids, clasts and bioclasts are organized consistently across and through Member 2 according to the underlying deposit geometry, indicating a gradual deposit accretion with no distinct collapse facies evident, no successive debris flow accumulation, and only localized intra-unit post- depositional modification. The stratigraphy and sedimentology of Member 2 supports a simple single-stage accumulation process through which Member 2 partially fills the Silberberg Grotto and extends into the deeper chambers and passages of the Sterkfontein Caves. Through this work we demonstrate at multiple scales the primary association between the sediments of Member 2 and the StW 573 ‘Little Foot’ skeleton.

Due to its completeness, the A.L. 288-1 (Lucy) skeleton has long served as the archetypal bipedal Australopithecus. However, there remains considerable debate about its limb proportions. There are three competing, but not necessarily mutually exclusive, explanations for the high humerofemoral index of A.L. 288-1: (1) a retention of proportions from an Ardipithecus-like most recent common ancestor (MRCA); (2) indication of some degree of climbing ability; (3) allometry. Recent discoveries of other partial skeletons of Australopithecus, such as those of A. sediba (MH1 and MH2) and A. afarensis (KSD-VP-1/1 and DIK-1/1), have provided new opportunities to test hypotheses of early hominin body size and limb proportions. Yet, no early hominin is as complete (>90%), as is the 3.67 Ma Little Foot (StW 573) specimen, from Sterkfontein Member 2. Here, we provide the first descriptions of its upper and lower long limb bones, as well as a comparative context of its limb proportions. As to the latter, we found that StW 573 possesses absolutely longer limb lengths than A.L. 288-1, but both skeletons show similar limb proportions. This finding seems to argue against an allometric explanation for the limb proportions of A.L. 288-1. In fact, our multivariate allometric analysis suggests that limb lengths of Australopithecus, as represented by StW 573 and A.L. 288-1, developed along a significantly different (p < 0.001) allometric scale than that which typifies modern humans and African apes. Our analyses also suggest, as have those of others, that hominin limb evolution occurred in two stages with: (1) a modest increase in lower limb length and a concurrent shortening of the antebrachium between Ardipithecus and Australopithecus, followed by (2) considerable lengthening of the lower limb along with a decrease of both upper limb elements occurring between Australopithecus and Homo sapiens.

International audience; Studies of the australopith (Australopithecus and Paranthropus) proximal femur have increasingly integrated information from the local arrangement of the cortical and cancellous bone to allow functional-biomechanical inferences on the locomotor behavioral patterns. In Australopithecus africanus and Paranthropus robustus, the cancellous bone organization at the center of the femoral head shows principal strut orientation similar to that of fossil and recent humans, which indicates that australopiths were human-like in many aspects of their bipedalism. However, by combining outer morphology with superoinferior asymmetry in cortical bone thickness at the base of neck and mid-neck, it has been suggested that, while adapted for terrestrial bipedality, australopiths displayed a slightly altered gait kinematics compared to Homo. We used techniques of 2D and 3D virtual imaging applied to an X-ray microtomographic record to assess cortical bone distribution along the entire femoral neck compartment in four upper femora from Swartkrans, South Africa (SK 82, SK 97, SK 3121, and SWT1/LB-2) and compared the results to the extant human and chimpanzee conditions. Our results support and extend previous evidence for more symmetric superior and inferior femoral neck cortical thicknesses in P. robustus than in modern humans and show that the differences are even greater than previously reported. However, P. robustus and humans still share a trend of lateral-to-medial decrease in asymmetry of the superior/inferior cortical thickness ratio, while this pattern is reversed in chimpanzees. We also identified two features uniquely characterizing P. robustus: an accentuated contrast between the relatively thicker anterior and the thinner posterior walls, and a more marked lateral-to-medial thinning of both cortices compared to extant humans and chimpanzees, which indicate wider interspecific differences among hominids in structural organization of the proximal femur than previously reported. It remains to be ascertained if, and to what extent, these features also characterize the femoral neck of Australopithecus.

OBJECTIVES A prevailing hypothesis in paleoanthropology is that early Pleistocene hominin bones were accumulated in South African caves by carnivores, which used those shelters, and the trees surrounding them, as refuge and feeding sites. We tested this hypothesis at the site of Drimolen, by comparing its hominin age-at-death distribution to that of the nearby and roughly contemporaneous site of Swartkrans. MATERIALS AND METHODS We employed standard dental aging systems in order to categorize the Drimolen hominin teeth into age classes of 5 years each. We then compared the age-at-death distribution for Drimolen with the published data available for the Swartkrans hominins. RESULTS Age-at-death distributions indicate that the age category "young adults" is the best represented age category at Swartkrans and the most poorly represented one at Drimolen. Moreover, Drimolen has a preponderance of infant specimens. Both sites have a low frequency of old adult specimens. CONCLUSIONS Differences observed in frequencies of the age-at-death categories suggest different mechanisms of hominin skeletal accumulation at Drimolen and Swartkrans. Swartkrans' frequency curve reflects mortality in a population subjected to predation and is thus consistent with the carnivore-accumulating hypothesis. In contrast, the Drimolen curve is similar to that of wild populations of living apes. Living primates have been observed exploiting caves as sleeping shelters, for nutritional, security, drinking, and thermoregulatory purposes. We suggest that similar cave use by Pleistocene hominins can explain, in large part, the accumulation of hominin bones at Drimolen. Such a conclusion is another illustration of the growing awareness that a "one-size-fits-all" taphonomic model for South African early Pleistocene hominin sites is probably insufficient.

Humans are set apart from other organisms by the realization of their own mortality. Thus, determining the prehistoric emergence of this capacity is of significant interest to understanding the uniqueness of the human animal. Tracing that capacity chronologically is possible through archaeological investigations that focus on physical markers that reflect “mortality salience.” Among these markers is the deliberate and culturally mediated disposal of corpses. Some Neandertal bone assemblages are among the earliest reasonable claims for the deliberate disposal of hominins, but even these are vigorously debated. More dramatic assertions center on the Middle Pleistocene sites of Sima de los Huesos (SH, Spain) and the Dinaledi Chamber (DC, South Africa), where the remains of multiple hominin individuals were found in deep caves, and under reported taphonomic circumstances that seem to discount the possibility that nonhominin actors and processes contributed to their formation. These claims, with significant implications for charting the evolution of the “human condition,” deserve scrutiny. We test these assertions through machine-learning analyses of hominin skeletal part representation in the SH and DC assemblages. Our results indicate that nonanthropogenic agents and abiotic processes cannot yet be ruled out as significant contributors to the ultimate condition of both collections. This finding does not falsify hypotheses of deliberate disposal for the SH and DC corpses, but does indicate that the data also support partially or completely nonanthropogenic formational histories.

The Sterkfontein Caves site is one of the richest early hominin fossil localities in Africa. More specifically, the fossiliferous deposits within the lower-lying Jacovec Cavern have yielded valuable hominin remains; prominent among them is the Australopithecus partial cranium StW 578. Due to the fragmentary nature of the braincase, the specimen has not yet been formally assigned to a species. In this context, we employ microtomography to quantify cranial thickness and composition of StW 578 in order to assess its taxonomic affinity. As comparative material, we investigate 10 South African hominin cranial specimens from Sterkfontein (StW 505, Sts 5, Sts 25, Sts 71), Swartkrans (SK 46, SK 48, SK 49) and Makapansgat (MLD 1, MLD 10, MLD 37/38), attributed to either Australopithecus or Paranthropus, as well as 10 extant human and 10 extant chimpanzee crania. Thickness variation in and structural arrangement of the inner and outer cortical tables and the diploe are automatically assessed at regular intervals along one parasagittal and one coronal section. Additionally, topographic cranial vault thickness distribution is visualized using color maps. Comparisons highlight an absolutely and relatively thickened condition of the StW 578 cranial vault versus those of other South African Plio-Pleistocene hominins. Moreover, in StW 578, as well as in the Australopithecus specimens Sts 5 and Sts 71 from Sterkfontein, the diploic layer contributes substantially to cumulative vault thickness (i.e., >60%). Within the comparative sample investigated here, StW 505 and Sts 71 from Sterkfontein Member 4, both attributed to Australopithecus, most closely resemble StW 578 in terms of cranial vault thickness values, tissue proportions, and two- and three-dimensional distributions. Including additional Plio-Pleistocene Australopithecus and Paranthropus crania from South and East Africa in future studies would further help establish morphological variability in these hominin taxa.

The oldest recognized artifacts at the Swartkrans cave hominid-bearing site in South Africa have long been known to occur in the Lower Bank of Member 1, now dated with the cosmogenic nuclide burial method to ca. 1.8–2.19 Ma. However, the affinities of this industry have been debated due to small sample size. In this paper we present newly excavated material from the Lower Bank retrieved since 2005 in the Swartkrans Paleoanthropological Research Project. The sample is now large enough to confirm its affinity with the Oldowan industrial complex. The assemblage is highly expedient and core reduction strategies are largely casual. Although freehand flaking is present, the bipolar technique is most significant, even in non-quartz raw materials. The Swartkrans assemblage shows some significant contrasts with the Sterkfontein Oldowan, ca. 2.18 Ma, which can be explained by its closer proximity to raw material sources, its somewhat different geographic context, and its more expedient nature. The Swartkrans Oldowan now provides us with the first good indication of Oldowan variability in southern Africa, where only two sizeable assemblages have thus far been discovered. Comparisons are made with other sites across Africa that help to place this variability within our overall understanding of the Oldowan industrial complex.

There is a long tradition of neotaphonomic research in paleontology and archaeology that is intended for building models of the bone-accumulating and -modifying behaviors of prehistoric biological actors. Building on that tradition, we present new data comparing the ways by which wild and captive leopards ( Panthera pardus ) altered the meat-bearing long limb bones of small bovids on which they fed. Compared to wild leopards, a captive leopard created more than twice as many tooth marks per skeletal element. In addition, the captive leopard imparted much more intensive gross damage to bones than did wild leopards—including complete consumption of some bones or deletion of bone portions (especially epiphyses) and extensive fragmentation of diaphyses. We attribute the stark divergence of taphonomic results in the bone samples formed by wild and captive leopards to environmental variability: the wild samples were modified by leopards operating in systems characterized by low to moderate feeding competition; the captive sample was created by a leopard operating in the complete absence of feeding competition but (unlike the wild leopards) was also subjected to the tedium of confinement—a condition that prompted repeated and prolonged bouts of chewing beyond what was required to simply extract nutrients from bones. The variable condition of the two bone samples also represents opposing endpoints of leopard taphonomic potential—the wild sample reflecting minimal expected alteration and the captive sample reflecting maximal expected alteration. Neither endpoint serves as accurate proxy for expectations of the taphonomic alteration of small bovid bones by leopards operating in a natural setting that is characterized by moderate to high feeding competition, such as Tanzania's modern Serengeti ecosystem or that reconstructed for most African Pliocene and Pleistocene habitats. Our results do, however, reinforce the conclusions of a previous study comparing taphonomic patterns created by wild and captive lions ( P. leo )—that environmental context is a key variable to control if the analyst's goal is to produce the most scientifically realistic and usefully applicable models of the bone-accumulating and -modifying behaviors of prehistoric biological actors.

Based on the cosmogenic nuclide burial dating technique, we present new radiometric age estimates of 2.19 ± 0.08 and 1.80 ± 0.09 million-years-old (Ma) for Member 1, and 0.96 ± 0.09 Ma for Member 3 of the Swartkrans Formation in South Africa. Our data are consistent with, and expand upon, results from previous radiometric dating techniques used at the site. The burial ages of Member 1 are consistent with the uranium–lead (U–Pb) age provided by bracketing flowstones (Pickering et al., 2011), while the age of Member 3 is significantly more precise than the large age bracket provided by U–Pb dating of tooth enamel (Balter et al., 2008) and recently re-evaluated electron spin resonance data (Herries and Adams, 2013). These new dates provide the complete age range for the extinct hominin, Paranthropus robustus , as well as indicate the first appearance of the genus Homo in southern Africa. Our results also indicate: the first, as well as the last, manufacture and use of bone digging tools in South Africa; some of the earliest evidence of stone tool use and large animal butchery in South Africa; and one of the earliest archaeological indications of the domestication of fire in the world.

We describe eleven hominin metacarpals and phalanges recovered from Jacovec Cavern and Member 4 of the Sterkfontein Formation between 1998 and 2003. Collectively, the fossils date in excess of 2.0 Ma, and are probably attributable to Australopithecus africanus and/or Australopithecus prometheus. When combined with results of previous studies on Australopithecus postcranial functional morphology, the new data presented here suggest that at least some late Pliocene and/or early Pleistocene hominins from Sterkfontein were arboreally adept. This finding accords with the reconstruction of the site's >2.0 Ma catchment area as well-vegetated and containing significant woody components. In addition, most of the new specimens described here evince morphologies that indicate the hands from which they derived lacked complete modern humanlike manual dexterity, which is integral to the manufacture and use of intentionally shaped stone tools. The absence of lithic artifacts from both stratigraphic units from which the fossils were excavated is consistent with this conclusion.

The Sterkfontein palaeokarst deposits, the richest repository of Australopithecus fossils in the world, are currently divided into six members (M1–M6) and named the Sterkfontein Formation ( Partridge and Watt, 1991 ). In this system, it was proposed that the passages and chambers deeper than the Silberberg Grotto formed most recently and are filled with comparatively young sediments from newer openings or reworked sediments from older, upper chambers. Here we describe the geomorphology, sedimentology, fauna and archaeological evidence from six deposits excavated near the base of the system in the Milner Hall. The use of a multidisciplinary approach enabled us to clarify the complex stratigraphic history, to track artefact-bearing deposits through multiple phases of deposition, and to identify an early deposit accumulated from the Silberberg Grotto to the base of the system. We propose that the Australopithecus -bearing Member 2 deposit originally extended down to near the base of the Milner Hall. We further propose that the vertical extent of the cave system was developed prior to the opening to the landscape surface, and that at that time the groundwater level was comparable to that seen now. We therefore argue that some of the earliest palaeoanthropological material in South Africa may have been deposited into the deepest chambers of the Sterkfontein Caves.

The Sterkfontein Caves site is one of the richest early hominin localities in Africa. In addition to significant fossil assemblages from Members 2 and 4 of the Sterkfontein Formation, recent excavations have revealed hominin-bearing sedimentary deposits in the lesser-known Milner Hall. We describe two hominin cranial fragments excavated from the Milner Hall in 2015 and present the results of a high-resolution microtomographic-based approach to diagnosing the anatomical and taxonomical origin of these specimens. Based on external morphology, StW 671 and StW 672 are tentatively identified as frontal and occipital fragments, respectively. Our non-invasive bi-dimensional quantitative investigation of the two cranial fragments reveals a mean cranial thickness of 8.8 mm and 5.6 mm for StW 671 and StW 672 respectively and a contribution of the diploic layer to the cumulative cranial thickness that is less than 50%. While the mean cranial thickness of StW 671 falls within the range of Homo, the relative proportion of the diploë in both StW 671 and StW 672 is lower than in Australopithecus (>60%) and extant humans (>50%). Accordingly, in terms of both cranial thickness and inner structural organization, the Milner Hall hominins combine derived and unique traits, consistent with the condition of other postcranial and dental material already described from the deposit. Moreover, our study opens interesting perspectives in terms of analysis of isolated cranial fragments, which are abundant in the hominin fossil record. Significance: The Sterkfontein Caves have widely contributed to our understanding of human evolution. Besides the well-known Members 4 and 2, where the iconic ‘Mrs Ples’ and ‘Little Foot’ have been found, in this study we suggest that the Milner Hall locality represents an additional, stratigraphically associated source of not only fossil hominins, but also Oldowan stone tools. In particular, we describe for the first time two cranial fragments, StW 671 and StW 672, identified as frontal and occipital bones, respectively. Our microtomographic-based analysis of these materials reveals some affinities with Homo combined with unique characters. In this context, our study suggests an intriguing mosaicism consistent with the description of the two fossil hominins found in the Milner Hall.

Dental characters can provide vital clues for understanding intra- and intertaxonomic morphological variation and its underlying genetic and environmental components. However, the unambiguous identification of particular traits and their comparative study is often confounded by lack of consistent terminology in the relevant literature. This difficulty is exacerbated when the etiologies are not completely understood, as is the case with talon cusps. To date, research on talon cusps has focused on modern humans. In many instances, descriptions of talon cusps appear in clinical case studies focusing on their treatment and removal. What is lacking in those discussions, though, is a comparative framework, in which the occurrence of talon cusps in nonhuman primates, and possibly other mammals, is established and understood. Here, we report on a taloned upper central incisor of a wild baboon (Papio hamadryas ursinus) from South Africa. The anomalous incisor of this individual includes an exaggerated accessory cusp diagnosed as a Type II talon. Microcomputed tomographic and radiographic analyses show that the taloned cusp possesses enamel, dentin, and pulp. In addition, we identified an unclassifiable talon cusp on a central maxillary incisor of a baboon skull housed in the Smithsonian Institution's Natural History Museum collection. Our observations of talon cusps on baboon incisors demonstrate that, with regard to this phenomenon, systematic study of nonhuman primates is much needed, along with a consistent use of terminology in the anatomical and anthropological literature. Finally, we present a hypothesis of the formation of talon cusps on mammalian incisors. Anat Rec, 296:1874–1880, 2013. © 2013 Wiley Periodicals, Inc.

We describe 14 hominin teeth and tooth fragments excavated recently from Swartkrans Cave (South Africa). The fossils derive from Members 1 (Lower Bank) and 3, from the Member 2/3 interface and from two deposits not yet assigned to member (the “Talus Cone Deposit” and the “Underground North Excavation” [UNE]) of the Swartkrans Formation, and include the first hominin fossil from the UNE, the two smallest Paranthropus robustus deciduous maxillary second molars in the entire hominin fossil record, and one of the smallest P. robustus permanent maxillary second molars from Swartkrans. The small permanent molar is accompanied by another tooth from a different individual but from the same stratigraphic level of the Swartkrans Formation; this second tooth is among, if not, the largest P. robustus permanent maxillary first molars known from anywhere—lending credence to assertions that degrees of body size sexual dimorphism previously ascribed to this species may be underestimated. It is more equivocal whether this evidence also supports hypotheses proposing that P. robustus assemblages from Swartkrans (as well as those from other South African cave sites) formed through the taphonomically biasing actions of large carnivores.

The world’s first archaeological traces from 2.6 million years ago (Ma) at Gona, in Ethiopia, include sharp-edged cutting tools and cut-marked animal bones, which indicate consumption of skeletal muscle by early hominin butchers. From that point, evidence of hominin meat-eating becomes increasingly more common throughout the Pleistocene archaeological record. Thus, the substantive debate about hominin meat-eating now centers on mode(s) of carcass resource acquisition. Two prominent hypotheses suggest, alternatively, (1) that early Homo hunted ungulate prey by running them to physiological failure and then dispatching them, or (2) that early Homo was relegated to passively scavenging carcass residues abandoned by carnivore predators. Various paleontologically testable predictions can be formulated for both hypotheses. Here we test four predictions concerning age-frequency distributions for bovids that contributed carcass remains to the 1.8 Ma. old FLK 22 Zinjanthropus (FLK Zinj, Olduvai Gorge, Tanzania) fauna, which zooarchaeological and taphonomic data indicate was formed predominantly by early Homo. In all but one case, the bovid mortality data from FLK Zinj violate test predictions of the endurance running-hunting and passive scavenging hypotheses. When combined with other taphonomic data, these results falsify both hypotheses, and lead to the hypothesis that early Homo operated successfully as an ambush predator.

Age profiling of fossil faunal samples relies on the correlation of animal tooth-wear patterns with life history stages, but the criteria used to infer these stages are not necessarily valid. Here we redefine some commonly used prey age classes, such as “juvenile,” “prime-age adult,” and “old adult,” based on the variable characteristics of tooth wear that we have observed in different ungulate size classes, and argue that prey vulnerability to predation is not so clearly predicted by the simplified age classes in widespread use by zooarchaeologists. We recommend instead classifying the youngest animal remains as either young juvenile or subadult juvenile, and adult remains as early prime, late prime or old, and provide specific criteria of dental eruption and occlusal wear for making these determinations. We argue this refined age profiling system, when used in combination with other types of zooarchaeological and taphonomic data, can provide accurate inferences of faunal accumulation processes.

It has been hypothesized that plant underground storage organs (USOs) played key roles in the initial hominin colonization of savanna habitats, the development of the distinctive skull and tooth morphology of the genus Australopithecus , and the evolution of the genus Homo by serving as “fallback foods” exploited during periods of food shortage. These hypotheses have been tested mostly by morphological, isotopic, and microwear analyses of hominin bones and teeth. Archaeological evidence of USO digging technology is equivocal. Until now relevant data from studies of chimpanzees, useful in behavioral models of early hominins because of their phylogenetic proximity and anatomical similarities, have been lacking. Here we report on the first evidence of chimpanzees using tools to dig for USOs, suggesting that exploitation of such resources was within the cognitive and technological reach of the earliest hominins. Consistent with scenarios of hominin adaptation to savannas, these data come from Ugalla (Tanzania), one of the driest, most open and seasonal chimpanzee habitats. USOs are, however, exploited during the rainy season, well after the period of most likely food shortage, contradicting the specific prediction of fallback food hypotheses. The discovery that savanna chimpanzees use tools to obtain USOs contradicts yet another claim of human uniqueness and provides a model for the study of variables influencing USO use among early hominins.

Modern humans are characterized by specialized hand morphology that is associated with advanced manipulative skills. Thus, there is important debate in paleoanthropology about the possible cause–effect relationship of this modern human-like (MHL) hand anatomy, its associated grips and the invention and use of stone tools by early hominins. Here we describe and analyse Olduvai Hominin (OH) 86, a manual proximal phalanx from the recently discovered >1.84-million-year-old (Ma) Philip Tobias Korongo (PTK) site at Olduvai Gorge (Tanzania). OH 86 represents the earliest MHL hand bone in the fossil record, of a size and shape that differs not only from all australopiths, but also from the phalangeal bones of the penecontemporaneous and geographically proximate OH 7 partial hand skeleton (part of the Homo habilis holotype). The discovery of OH 86 suggests that a hominin with a more MHL postcranium co-existed with Paranthropus boisei and Homo habilis at Olduvai during Bed I times., The homin fossil record reveals a complex pattern of hand evolution. Here, the authors describe a phalanx of a >1.84-million-year-old unidentified hominin, which represents the earliest modern human like hand bone in the fossil record.

Some chimpanzees use 2 types of tools to extract underground termites for consumption. Chimpanzees insert thin, flexible probes into tunnels or holes in termite mounds (fishing), and sometimes use stouter, rigid sticks to first puncture the holes and also possibly to fish. Many puncturing sticks have distinctive “brushed” ends. Researchers have hypothesized that chimpanzees create the brushed ends intentionally to increase their affixibility to biting termites (Sugiyama, 1985). The results of our archaeological analysis of a large collection of puncturing sticks used by Central African chimpanzees falsifies this hypothesis, and instead agrees with the recent behavioral observations of Sanz et al. (2004; cf. Bermejo and Illera, 1999) that brushing is a coincidental result of procuring sticks from vegetation sources. The results highlight the positive contribution of an archaeological approach to problems in chimpanzee material culture and emphasize to primatologists the value of curating artifacts.

The common occurrence of hammerstone percussion damage (pits, striae, notches and impact flakes) on the fossil limb bones of ungulates indicates that marrow extraction has been an important component of hominid butchery for over two million years. Beyond this level of basic inference, it would be behaviorally informative if three deeper aspects of marrow harvesting were understood more clearly: (1) whether inter-element patterns of bone fragmentation vary when processing intensity is held constant; (2) whether butcher investment in marrow extraction correlates positively with the number of percussion marks generated; (3) whether taphonomic effectors can be identified based on percussion mark morphology, frequency and placement. Some experimental work has been conducted previously in service of exploring these questions, but we set out here to address them explicitly through the analysis of a large sample of white-tailed deer (Odocoileus virginianus) limb elements fractured by hammerstone percussion. Our results indicate that (1) measures of bone fragmentation, which supposedly reflect processing intensity, are highly contingent on the research question being posed. This stresses the fact that researchers must be explicit in their definition of processing intensity. (2) In addition, hypothesized covariance between number of hammerstone blows and percussion mark frequencies are not met in our sample, corroborating previous conclusions of a lack of covariance between cutting strokes and cutmark frequencies. These results highlight the contingent nature of butchery mark production, and emphasize the need to investigate carcass resource exploitation by posing questions that do not rely on mark frequencies, but instead utilize other zooarchaeological measures. (3) Finally, our results—showing high incidences of impact notches and flakes created by direct anvil contact and “anvil scratches” created by direct hammerstone contact—suggest caution in using specific categories of percussion damage to infer their taphonomic effectors.

Recently, Alcantara Garcia et al. (in press) presented a new method and criteria for distinguishing between fractures imparted by hominid hammerstone percussion and carnivores chewing on ‘green’ limb bones of ungulates. The method uses a combination of fracture plane and fracture angle data that are useful for elucidating the relative role of hominids in the accumulation of prehistoric archaeofaunas, especially when employed in concert with other classes of taphonomic data. We briefly summarise the method and apply it to the ungulate limb bone subassemblage from Swartkrans Member 3, a c. 1.0 million year old site from South Africa that preserves Early Stone Age lithic artefacts, hominid fossils, and an abundant mammalian fauna with cutmarked, hammerstone-percussed and burned bone specimens. Results of the fracture pattern analysis corroborate indications from other lines of taphonomic data that there was minimal carnivore–hominid interdependence in the formation of the fauna, and that carnivores were probably responsible for the majority of the bone collection in Member 3. However, we also document a significant hominid influence on assemblage formation, a finding that expands and refines our understanding of large animal carcass foraging by hominids in southern Africa during the early Pleistocene. Copyright © 2004 John Wiley & Sons, Ltd.