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Acknowledgements IE has received funding from the postdoctoral fellowship programme Beatriu de Pinós (2019 BP00247), funded by the Secretary of Universities and Research (Government of Catalonia) and by the Horizon 2020 programme of research and innovation of the European Union under the Marie Skłodowska-Curie grant agreement No. 801370. Laboratory materials for sample processing at the University of Barcelona were funded by the Ministerio de Ciencia e Innovaci on - PID 2020-119773 GB-I00. Visit to the site for sampling collection for this study was sponsored by a NRF Y-Rated Research Grant (No. 116358) to DS. PdlP was supported by a Poroulis grant through Cambridge University. PdlP thanks the action PID 2019- 1049449 GB-I00 funded by Spanish FEDER/Ministry of Science and Innovation. LB was funded by a National Geographic Explorer grant (NGS-54810R-19), DSI-NRF Centre of Excellence in Palaeosciences (Genus) grant (CEOOP 2020-1), and NRF African Origins Platform grant (No. 98824). Research by FdE is funded by the Research Council of Norway (SFF Centre for Early Sapiens Behaviour- SapienCE-project 262618), the Talents program of the University of Bordeaux (n. 191022_001), and the program “Human Past”, funded by the same university.We are grateful to Amafa for issuing us with the excavation permit (SAH 15/7645), and to SAHRA for an export permit (CaseID 15987).We thank two anonymous reviewers for their helpful comments that significantly improved the manuscript., Supplementary data to this article can be found online at https://doi.org/10.1016/j.quascirev.2022.107898., Border Cave is a well-known South African Middle and Early Later Stone Age site located in KwaZulu-Natal. The site has exceptional plant preservation, unparalleled in the African Middle Stone Age archaeological record. This study focuses on the phytolith and FTIR analysis of two Members (2 BS and 2 WA) of the under-documented post-Howiesons Poort occupations dating to ∼60 ka. These members contain complex successions of vertically overlapping, interdigitating light brown sediments, plant bedding and combustion features of various sizes. The complexity and distinctiveness of these deposits provide an excellent opportunity for the study of plant exploitation strategies and their associated human behaviour. Our taphonomic assessment inferred, through the variability of phytolith properties and minerals composing archaeological layers, that specific occupations suffered more physical weathering than others, for example in the form of trampling. The preservation of fragile and highly soluble phytoliths (eudicot leaf phytoliths) and the high frequencies of articulated phytoliths indicates that some bedding deposits experienced little disturbance after their deposition. Not all bedding layers dating to ⁓60 ka show, from a phytolith perspective, the same plant composition, which could be explained in terms of changes in human preference for the use of plants over time to construct bedding or because distinct types of living floors are represented. Finally, the systematic application of phytoliths and FTIR to the complex archaeological sequence of Border Cave confirm these analyses can be used in the future to identify bedding deposits not visible to the naked eye, and behavioural patterns obscured by diagenetic or biased processes during sampling., Programme Beatriu de Pinós (2019 BP00247), funded by the Secretary of Universities and Research (Government of Catalonia), Horizon 2020 programme of research and innovation of the European Union under the Marie Skłodowska-Curie grant agreement No. 801370, Ministerio de Ciencia e Innovación - PID 2020-119773 GB-I00, NRF Y-Rated Research Grant (No. 116358), Action PID 2019- 1049449 GB-I00 funded by Spanish FEDER/Ministry of Science and Innovation, National Geographic Explorer grant (NGS-54810R-19), DSI-NRF Centre of Excellence in Palaeosciences (Genus) grant (CEOOP 2020-1), NRF African Origins Platform grant (No. 98824), Research Council of Norway (SFF Centre for Early Sapiens Behaviour- SapienCE-project 262618), Talents program of the University of Bordeaux (n. 191022_001), Program “Human Past” University of Bordeaux, SAH 15/7645, CaseID 15987

Sterkfontein is the most prolific single source of Australopithecus fossils, the vast majority of which were recovered from Member 4, a cave breccia now exposed by erosion and weathering at the landscape surface. A few other Australopithecus fossils, including the StW 573 skeleton, come from subterranean deposits [T. C. Partridge et al. , Science 300, 607–612 (2003); R. J. Clarke, K. Kuman, J. Hum. Evol. 134, 102634 (2019)]. Here, we report a cosmogenic nuclide isochron burial date of 3.41 ± 0.11 million years (My) within the lower middle part of Member 4, and simple burial dates of 3.49 ± 0.19 My in the upper middle part of Member 4 and 3.61 ± 0.09 My in Jacovec Cavern. Together with a previously published isochron burial date of 3.67 ± 0.16 My for StW 573 [D. E. Granger et al. , Nature 522, 85–88 (2015)], these results place nearly the entire Australopithecus assemblage at Sterkfontein in the mid-Pliocene, contemporaneous with Australopithecus afarensis in East Africa. Our ages for the fossil-bearing breccia in Member 4 are considerably older than the previous ages of ca. 2.1 to 2.6 My interpreted from flowstones associated with the same deposit. We show that these previously dated flowstones are stratigraphically intrusive within Member 4 and that they therefore underestimate the true age of the fossils.

Surface lithic scatters are relatively common in arid environments worldwide and attest to human occupation and use of desert environmental resources over long (104–106-year) time scales. A recent paper by Hardaker (2020) describes Early to Middle Stone Age lithic scatters in desert environments in Namibia, and deduces their spatial patterns and the relative age of individual lithic artifacts. Here, we challenge the results of Hardaker’s study based on (1) the methodology that was employed to map the scatters and analyse lithic age, and (2) the spatial interpretation applied to the lithic scatters and the derived ages of individual artifacts. This re-evaluation of Hardaker’s study provides a better conceptual basis for documenting and interpreting lithic scatters worldwide, which should apply established geoarchaeological methods and be set in the context of desert geomorphological processes and environments during the Quaternary.

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 origins of Homo , as well as the diversity and biogeographic distribution of early Homo species, remain critical outstanding issues in paleoanthropology. Debates about the recognition of early Homo , first appearance dates, and taxonomic diversity within Homo are particularly important for determining the role that southern African taxa may have played in the origins of the genus. The correct identification of Homo remains also has implications for reconstructing phylogenetic relationships between species of Australopithecus and Paranthropus , and the links between early Homo species and Homo erectus . We use microcomputed tomography and landmark-free deformation-based three-dimensional geometric morphometrics to extract taxonomically informative data from the internal structure of postcanine teeth attributed to Early Pleistocene Homo in the southern African hominin-bearing sites of Sterkfontein, Swartkrans, Drimolen, and Kromdraai B. Our results indicate that, from our sample of 23 specimens, only 4 are unambiguously attributed to Homo , 3 of them coming from Swartkrans member 1 (SK 27, SK 847, and SKX 21204) and 1 from Sterkfontein (Sts 9). Three other specimens from Sterkfontein (StW 80 and 81, SE 1508, and StW 669) approximate the Homo condition in terms of overall enamel–dentine junction shape, but retain Australopithecus -like dental traits, and their generic status remains unclear. The other specimens, including SK 15, present a dominant australopith dental signature. In light of these results, previous dietary and ecological interpretations can be reevaluated, showing that the geochemical signal of one tooth from Kromdraai (KB 5223) and two from Swartkrans (SK 96 and SKX 268) is consistent with that of australopiths.

This contribution focuses on the recently excavated lithic assemblage from Border Cave Members 1 RGBS, 3 BS, 2 WA and 2 BS. These members were attributed by Beaumont to the Howiesons Poort and post-Howiesons Poort Industries of the southern African Middle Stone Age. Here we consider lithics as indicators of cultural behaviour, site formation processes, and occupation intensity. As such, the assemblage is explored in depth through lithic technology attribute analysis, particle size distribution, and spatial analysis. These lines of inquiry follow the new allostratigraphic divisions proposed for the deposits by Stratford and colleagues. Results show that the lower members share a degree of similarity in terms of flaking strategies and raw material selection, whereas the upper members record a dissimilar set of features, with differentiation between them. The technological and spatial analyses suggest the sporadic presence of different groups at the site, each characterized by distinct sets of knapping techniques and methods but employing related knapping strategies and using similar raw materials. The interpretation of these members as a palimpsest of sporadic occupation is supported by the low number of archaeological finds by sediment volume. The evidence at hand does not support a Howiesons Poort attribution for the lowest members. The data show that lithics from part of Member 2 WA and 2 BS are in accordance with those of the recently defined Sibudan technocomplex.

Mozambique is a crucial area for unravelling our evolutionary history, during the early phases of the Quaternary. Indeed,located at the southern end of the Great Rift between Southern and Eastern Africa, this area plays a key‐role in our understanding of the paleobiogeography of fauna documented in the fossil record of East and South Africa. The Human Origins in Mozambique and Malawi Environments – HOMME – project aims to identify evidence of early hominins in this area, which is under‐studied as compared to eastern and southern African regions, as well as to characterize the paleoenvironmental contexts in which the genus Homo emerged. This paper presents the results obtained during our first field season. We explored various karstic cavities located along the Buzi river and the Cheringoma Plateau (west and north off Beira). Our explorations reveal the potential of the Mozambican karst systems for providing new Plio‐Pleistocene paleontological evidence. In particular, we report various endokarstic deposits containing paleontological remains. These promising findings reveal the richness of this area and the fundamental role of the karst in trapping and preserving fossils. Accordingly, this project will certainly fill in a number of gaps in the paleobiogeography and paleobiology of early hominins., Enclavé entre l’Afrique australe et l’Afrique de l’Est, le Mozambique est un secteur essentiel à la compréhension de notre histoire évolutive et ce en particulier pour les phases anciennes du Quaternaire, entre 2,6 et 2 Ma. Le Mozambique, par sa situation géographique comprenant la partie terminale du Grand Rift, apparaît comme la clef des problématiques paléobiogéographiques entre les deux grandes aires est‐ et sud‐africaines. Le projet HOMME – Human Origins in Mozambic and Malawi Environments – vise à identifier, dans ce secteur encore trop peu étudié, les traces du passage des Homininés anciens (Australopithécinés et premiers Homo) ou, tout du moins, à caractériser les paléoenvironements au sein desquels ils ont pu évoluer. Nous présentons ici les premiers résultats de nos premières campagnes de terrain. L’exploration de plusieurs cavités karstiques, à l’ouest (secteur de la Buzi River) et au nord (secteur de Cheringoma Plateau) de la ville de Beira, nous a permis de reconnaître l’importance et les potentialités archéo‐paléontologiques des réseaux karstiques mozambicains. Des reliques de brèches endokarstiques riches en matériel paléontologique (dont porc‐épic, hyène, phacochère) ont d’ailleurs été identifiées. À l’issu de cette campagne, le matériel osseux et la présence de brèches fossilifères laissent entrevoir la richesse du secteur qui nous permettra, à terme, de combler nombre de lacunes concernant la paléobiogéographie des Homininés anciens.

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 Cradle of Humankind World Heritage site in South Africa is renowned for its karstic cave systems, which have yielded extensive fossil and stone tool assemblages dating to the Plio-Pleistocene period. The archaeological record from these caves has provided significant evidence for the evolution of lithic technology in southern Africa. Open-air sites have also been documented in this region, although they rarely receive scientific attention due to their lack of spatial and stratigraphic contexts. However, archaeological research has demonstrated that lithic production strategies were organized through landscape-scale constraints including the availability of raw materials and the energetic costs of their transport. As such, the study of open-air Stone Age sites in the Cradle of Humankind can potentially offer insight into the relationships between raw material procurement and reduction processes in this region. Here, we present a case study from the open-air locality of Elandsdrift Farm, which preserves a palimpsest of Earlier and Middle Stone Age lithic materials. While the chronological and spatial contexts of this locality present significant interpretative challenges, the Elandsdrift Farm lithic materials are not without scientific value. A landscape archaeological approach is employed to explore the influence of raw material transport on tool-making. Trends in core reduction intensity and flaking efficiency suggest that landform ruggedness and distances traveled to procure raw materials significantly influenced lithic production strategies throughout the Early to Middle Pleistocene. Despite the contextual challenges presented by open-air localities, landscape archaeological methods can inform the economic aspects of lithic scatters in this region, which cannot be easily inferred from archaeological assemblages in cave contexts.

Leopards have been the focus of taphonomic research due to their capacity to create bone accumulations, prey on some hominin groups or potentially provide scavenging opportunities to early hominins...

In 2015, which marked 35 years since Beaumont had worked at the site, we renewed excavations at Border Cave. Our primary aims were to reassess the stratigraphic context of the sedimentary and cultural sequence, gain insight into site formation processes, make a detailed study of organic remains, identify long term cultural trends, and characterize expressions of complex behaviour and innovation. This contribution serves as an update on activities conducted in 2018 and 2019 and provides an overview of our research findings to date, placing them in the broader context of the Middle Stone Age in southern Africa. New luminescence ages based on feldspar grains in the sedimentary sequence are in broad agreement with the previous chronology established for the site. Geoarchaeology and faunal taphonomy have started to elucidate site formation processes, showing that the members should not be considered as homogeneous units, and that associated formation interpretations established by Beaumont are simplifications that are not representative of the diverse site formation processes active in the shelter. This finding is supported by lithic analysis of the Member 2 WA assemblage that shows differences in technology between artefacts from the top, middle, and lower part of the same member. In addition, the lithic artefacts from the middle and lower part of Member 2 WA show continuities with the lithics from the underlying Members 3 BS and 1 RGBS, which were attributed by Beaumont to a different industry. Grass mats/bedding layers are preserved throughout the sequence, the oldest of which dates to ∼200 ka. The use of ash and leaves with insecticidal properties in the bedding construction reflects complex cognition, as does the cooking of starchy rhizomes that come from layers dated to 170 ka. In addition to a rich mammal fauna found in all of the deposits, the remains of a new individual, a 3–4-year-old child, were recovered from Member 1 BS.LR C that has an ESR date of 42.6 ka.

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.

Olieboomspoort is one of the few rock shelters in the vast interior of southern Africa documenting pulses of occupation from the Acheulean until the end of the Later Stone Age. Revil Mason excavated the site in 1954 and attributed the large Middle Stone Age (MSA) lithic assemblage to his middle phase of the so-called Pietersburg Industry. Recent work at the site has focused on the Holocene layers, but little is known about the earlier phases of shelter use. Here, we provide some background to the shelter, give a history of past research and present initial results following renewed fieldwork at the site. The MSA deposits contain abundant lithic artefacts and ochre, and we present an initial description of these cultural remains. Palynological analysis reveals limited potential for palaeoenvironmental reconstructions, but some faunal remains indicate open grasslands. We dated two equid teeth that provided highly consistent combined U-series-ESR estimates, resulting in a mean age of 150 ± 14 ka (1σ). Even when considering potential sources of uncertainty such as variations in water-intake, these fossils can securely be dated to Marine Isotopic Stage 6. Our reappraisal of site formation processes highlights the fact that the archaeological assemblage is strongly time-averaged. We discuss these different results in the context of a recently rekindled interest in the so-called Pietersburg Industry.

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.

Funder: DST-NRF Centre of Excellence in Palaeosciences, Funder: Palaeontological Scientific Trust, Numerous aspects of early hominin biology remain debated or simply unknown. However, recent developments in high-resolution imaging techniques have opened new avenues in the field of paleoanthropology. More specifically, X-ray synchrotron-based analytical imaging techniques have the potential to provide crucial details on the ontogeny, physiology, biomechanics, and biological identity of fossil specimens. Here we present preliminary results of our X-ray synchrotron-based investigation of the skull of the 3.67-million-year-old Australopithecus specimen StW 573 (‘Little Foot’) at the I12 beamline of the Diamond Light Source (United Kingdom). Besides showing fine details of the enamel (i.e., hypoplasias) and cementum (i.e., incremental lines), as well as of the cranial bone microarchitecture (e.g., diploic channels), our synchrotron-based investigation reveals for the first time the 3D spatial organization of the Haversian systems in the mandibular symphysis of an early hominin.

Haeusler et al. (1) suggest that our analysis (2) of the distribution of relative bone volume across the articular surface (figure 5) does not justify different taxonomic allocations or locomotor classifications. We agree with their first suggestion, and we did not use these data to make direct arguments for the taxonomic attribution of either specimen in our paper. With regard to locomotor classifications, we note that our results “further [support] the inferred differences in loading between these two specimens as evidenced by their internal BV/TV distribution” (2). The strongest evidence for locomotor differences between StW 522 and StW 311 comes from the internal distribution of trabecular bone, which is not captured by the geometric … [↵][1]1To whom correspondence may be addressed. Email: m.skinner{at}kent.ac.uk. [1]: #xref-corresp-1-1

Early plant use is seldom described in the archaeological record because of poor preservation. We report the discovery of grass bedding used to create comfortable areas for sleeping and working by people who lived in Border Cave at least 200,000 years ago. Sheaves of grass belonging to the broad-leafed Panicoideae subfamily were placed near the back of the cave on ash layers that were often remnants of bedding burned for site maintenance. This strategy is one forerunner of more-complex behavior that is archaeologically discernible from ~100,000 years ago.

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.

Understanding the formation of the Sterkfontein Caves is critical for interpreting the geologic age and patterns of accumulation of the fossils found there. The complex stratigraphy of the caves has been investigated for decades, often yielding conflicting interpretation. Current research at Sterkfontein aims to apply new mutlidisciplinary stratigraphic analyses to new stratigraphically sensitive excavations, thereby providing high-resolution contextual support to recovered assemblages and attempts to increase our understanding of excavated deposit morphology through identification and study of sedimentary and stratigraphic features preserved in the remnants of previous excavations. The Sterkfontein deposits separated into six members, Member 1 to Member 6, of which Member 2, 4, and 5 have yielded hominin remains. Partial collapses and localized erosion of deposits are common, and the spaces created are often filled with sediments from distinct sources, complicating interpretation. This is especially the case with Members 4 and 5. Characterizing taxonomic, morphological, or cultural variability from excavated deposits where no stratigraphy was recognized is difficult but is the subject of ongoing work.

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.

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.

In this paper, we present a revised stratigraphy and results of preliminary analyses of the archaeological material from Mwulu’s Cave. This arises from two excavation campaigns conducted in 2017, 71 years after the site was initially investigated by P.V. Tobias. This cave, located in Limpopo Province (South Africa), preserves one of the few known Middle Stone Age sequences in the northeastern part of the country. Here, we revisit the stratigraphic sequence of the site and provide new analyses of sediments, palynomorphs, phytoliths, ochre and lithics. The renewed excavations and reappraisal of the archaeological material from Mwulu’s Cave form part of a larger research project exploring Middle Stone Age variability in the northeastern part of South Africa, with a specific focus on the so-called Pietersburg industries.

In the present study, we report brown hyena tooth marks on australopiths from Sterkfontein's Plio-Pleistocene-age Member 4 (South Africa). Classic taphonomic analyses and the implementation of new techniques, including Geometric Morphometrics and Machine Learning, are combined to identify the modifying agent and provide the first direct evidence of hyenid scavenging on australopiths. This hypothesis adds a new perspective to the relationships between carnivores and hominins in the Cradle by expanding on previous hypotheses proposing that leopards were the primary predator and bone accumulator of early hominin remains at South African palaeocaves.

The Sterkfontein Caves UNESCO World Heritage site represents one of South Africa’s most valuable cultural heritage resources and is one of the world’s most prolific palaeoanthropological sites with...

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.

Quaternary-age human and animal footprints have been reported from coastal deposits along the South African coast. These footprints and their sedimentary contexts can inform on Quaternary palaeoenvironments and processes stratigraphically associated with the palaeoanthropological record of past human occupation in the coastal zone, and exploitation of its resources. This study examines the palaeoenvironmental context and sedimentology of Marine Isotope Stage 5 aeolianite at Nahoon Point, southeastern coast of South Africa, where preserved human footprints have been previously recorded. This study identifies seven sedimentary facies from 20 logged profiles through the coastal aeolianite in order to develop a landscape-scale stratigraphic framework for coastal palaeoenvironmental reconstruction during the period of the MIS 5 highstand. Two phases of aeolianite deposition are identified, which bracket a single stepped sea-level transgression phase up to +7.82 ± 0.82 m asl, evidenced by intraformational beachrock facies and marine benches. This correlates with the second known global MIS 5e highstand; no evidence for the first highstand reported from the literature was found in the study area. Aeolianite bed geometries and sedimentary structures indicate that wind regimes and rainfall seasonality were similar to present. Newly discovered faunal tracks in aeolianites of the first depositional phase suggest the presence of unidentified bovids of varying sizes. Evidence for human activity is preserved in all three recorded depositional phases, suggesting people were present throughout the early MIS 5 sequence despite changing environmental conditions. This evidence suggests that human occupation of coastal areas, and the exploitation of marine food resources was more widespread than generally thought in southern Africa during early MIS 5.

International audience; Located in north-west Botswana, Drotsky's Cave is a vast cavity hosted within dolomite known for the discovery of archaeological strata dating back to the Late Stone Age. It is home to a colony of bats comprising several ten thousands of individuals who are responsible for the accumulation of a thick deposit of guano, on which we have conducted research in some of the deeper parts of the cavity. A detailed study of the morphologies of the dolomite walls and roofs demonstrates that part of the cave's speleogenesis is probably due to biogenic corrosion. This type of weathering and corrosion process is still poorly described in underground environments. However, it is now accepted that the role played by bats and their droppings on the condition of walls, their weathering, and the enlargement of galleries and chambers by condensation-corrosion processes, should be taken into account when considering the way deep networks evolve. In this article, we discuss the mechanisms of this biogenic corrosion in Drotsky's Cave, as well as their morphogenic impacts (the dissolution and recession of walls, the formation of lateral notches, arches, bell holes, and ceiling spherical cupolas), and the consequences for the wall retreat of conduits. Finally, we propose a new typological classification. The consequences of this biogenic corrosion are important for archaeological questions related to the taphonomy of archaeological remains in the buried deposits and the conservation of cave art. Because of the important impacts of guano on buried biological archaeological remains, such a population would have encouraged accelerated weathering, corrosion and dissolution of the walls, potentially contributing to the total, or partial, disappearance of any trace of human activity on the surrounding rocks.

(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.

In this paper, we present a revised stratigraphy and results of preliminary analyses of the archaeological material from Mwulu’s Cave. This arises from two excavation campaigns conducted in 2017, 71 years after the site was initially investigated by P.V. Tobias. This cave, located in Limpopo Province (South Africa), preserves one of the few known Middle Stone Age sequences in the northeastern part of the country. Here, we revisit the stratigraphic sequence of the site and provide new analyses of sediments, palynomorphs, phytoliths, ochre and lithics. The renewed excavations and reappraisal of the archaeological material from Mwulu’s Cave form part of a larger research project exploring Middle Stone Age variability in the northeastern part of South Africa, with a specific focus on the so-called ‘Pietersburg’ industries.

New excavations at Border Cave use high-resolution techniques, including FT-IR, for sediment samples and thin sections of micromorphology blocks from stratigraphy. These show that sediments have different moisture regimes, both spatially and chronologically. The site preserves desiccated grass bedding in multiple layers and they, along with seeds, rhizomes, and charcoal, provide a profile of palaeo-vegetation through time. A bushveld vegetation community is implied before 100,000 years ago. The density of lithics varies considerably through time, with high frequencies occurring before 100,000 years ago where a putative MSA 1/Pietersburg Industry was recovered. The highest percentage frequencies of blades and blade fragments were found here. In Members 1 BS and 1 WA, called Early Later Stone Age by Beaumont, we recovered large flakes from multifacial cores. Local rhyolite was the most common rock used for making stone tools, but siliceous minerals were popular in the upper members. Fil: Backwell, Lucinda Ruth. University of the Witwatersrand; Sudáfrica. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Tucumán; Argentina Fil: d'Errico, Francesco. Universite de Bordeaux; Francia. University of Bergen; Noruega Fil: Banks, William E.. Universite de Bordeaux; Francia Fil: de la Peña, Paloma. University of the Witwatersrand; Sudáfrica Fil: Sievers, Christine. University of the Witwatersrand; Sudáfrica Fil: Stratford, Dominic. University of the Witwatersrand; Sudáfrica Fil: Lennox, Sandra J.. University of the Witwatersrand; Sudáfrica Fil: Wojcieszak, Marine. University of the Witwatersrand; Sudáfrica Fil: Bordy, Emese M.. University Of Cape Town; Fil: Bradfield, Justin. University of the Witwatersrand; Sudáfrica Fil: Wadley, Lyn. University of the Witwatersrand; Sudáfrica

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.

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.