Late Acheulean occupations at Montagu Cave and the pattern of Middle Pleistocene behavioral change in Western Cape, southern Africa.

Patterns of so-called modern human behavior are increasingly well documented in an abundance of Middle Stone Age archaeological sites across southern Africa. Contextualized archives directly preceding the southern African Middle Stone Age, however, remain scarce. Current understanding of the terminal Acheulean in southern Africa derives from a small number of localities that are predominantly in the central and northern interior. Many of these localities are surface and deflated contexts, others were excavated prior to the availability of modern field documentation techniques, and yet other relevant assemblages contain low numbers of characteristic artifacts relative to volume of excavated deposit. The site of Montagu Cave, situated in the diverse ecosystem of the Cape Floral Region, South Africa, contains the rare combination of archaeologically rich, laminated and deeply stratified Acheulean layers followed by a younger Middle Stone Age occupation. Yet little is known about the site owing largely to a lack of contextual information associated with the early excavations. Here we present renewed excavation of Levels 21-22 at Montagu Cave, located in the basal Acheulean sequence, including new data on site formation and ecological context, geochronology, and technological variability. We document intensive occupation of the cave by Acheulean tool-producing hominins, likely at the onset of interglacial conditions in MIS 7. New excavations at Montagu Cave suggest that, while Middle Stone Age technologies were practiced by 300 ka in several other regions of Africa, the classic Acheulean persisted later in the Fynbos Biome of the southwestern Cape. We discuss the implications of this regionalized persistence for the biogeography of African later Middle Pleistocene hominin populations, for the ecological drivers of their technological systems, and for the pattern and pace of behavioral change just prior to the proliferation of the southern African later Middle Stone Age.

The first Miocene fossils from coastal woodlands in the southern East African Rift.

The Miocene was a key time in the evolution of African ecosystems witnessing the origin of the African apes and the isolation of eastern coastal forests through an expanding arid corridor. Until recently, however, Miocene sites from the southeastern regions of the continent were unknown. Here, we report the first Miocene fossil teeth from the shoulders of the Urema Rift in Gorongosa National Park, Mozambique. We provide the first 1) radiometric ages of the Mazamba Formation, 2) reconstructions of paleovegetation in the region based on pedogenic carbonates and fossil wood, and 3) descriptions of fossil teeth. Gorongosa is unique in the East African Rift in combining marine invertebrates, marine vertebrates, reptiles, terrestrial mammals, and fossil woods in coastal paleoenvironments. The Gorongosa fossil sites offer the first evidence of woodlands and forests on the coastal margins of southeastern Africa during the Miocene, and an exceptional assemblage of fossils including new species.

Genomic variation in baboons from central Mozambique unveils complex evolutionary relationships with other Papio species.

BACKGROUND: Gorongosa National Park in Mozambique hosts a large population of baboons, numbering over 200 troops. Gorongosa baboons have been tentatively identified as part of Papio ursinus on the basis of previous limited morphological analysis and a handful of mitochondrial DNA sequences. However, a recent morphological and morphometric analysis of Gorongosa baboons pinpointed the occurrence of several traits intermediate between P. ursinus and P. cynocephalus, leaving open the possibility of past and/or ongoing gene flow in the baboon population of Gorongosa National Park. In order to investigate the evolutionary history of baboons in Gorongosa, we generated high and low coverage whole genome sequence data of Gorongosa baboons and compared it to available Papio genomes. RESULTS: We confirmed that P. ursinus is the species closest to Gorongosa baboons. However, the Gorongosa baboon genomes share more derived alleles with P. cynocephalus than P. ursinus does, but no recent gene flow between P. ursinus and P. cynocephalus was detected when available Papio genomes were analyzed. Our results, based on the analysis of autosomal, mitochondrial and Y chromosome data, suggest complex, possibly male-biased, gene flow between Gorongosa baboons and P. cynocephalus, hinting to direct or indirect contributions from baboons belonging to the "northern" Papio clade, and signal the presence of population structure within P. ursinus. CONCLUSIONS: The analysis of genome data generated from baboon samples collected in central Mozambique highlighted a complex set of evolutionary relationships with other baboons. Our results provided new insights in the population dynamics that have shaped baboon diversity.

Experimental investigation of orangutans' lithic percussive and sharp stone tool behaviours.

Early stone tools, and in particular sharp stone tools, arguably represent one of the most important technological milestones in human evolution. The production and use of sharp stone tools significantly widened the ecological niche of our ancestors, allowing them to exploit novel food resources. However, despite their importance, it is still unclear how these early lithic technologies emerged and which behaviours served as stepping-stones for the development of systematic lithic production in our lineage. One approach to answer this question is to collect comparative data on the stone tool making and using abilities of our closest living relatives, the great apes, to reconstruct the potential stone-related behaviours of early hominins. To this end, we tested both the individual and the social learning abilities of five orangutans to make and use stone tools. Although the orangutans did not make sharp stone tools initially, three individuals spontaneously engaged in lithic percussion, and sharp stone pieces were produced under later experimental conditions. Furthermore, when provided with a human-made sharp stone, one orangutan spontaneously used it as a cutting tool. Contrary to previous experiments, social demonstrations did not considerably improve the stone tool making and using abilities of orangutans. Our study is the first to systematically investigate the stone tool making and using abilities of untrained, unenculturated orangutans showing that two proposed pre-requisites for the emergence of early lithic technologies-lithic percussion and the recognition of sharp-edged stones as cutting tools-are present in this species. We discuss the implications that ours and previous great ape stone tool experiments have for understanding the initial stages of lithic technologies in our lineage.

The Middle to Later Stone Age transition at Panga ya Saidi, in the tropical coastal forest of eastern Africa.

The Middle to Later Stone Age transition is a critical period of human behavioral change that has been variously argued to pertain to the emergence of modern cognition, substantial population growth, and major dispersals of Homo sapiens within and beyond Africa. However, there is little consensus about when the transition occurred, the geographic patterning of its emergence, or even how it is manifested in the stone tool technology that is used to define it. Here, we examine a long sequence of lithic technological change at the cave site of Panga ya Saidi, Kenya, that spans the Middle and Later Stone Age and includes human occupations in each of the last five Marine Isotope Stages. In addition to the stone artifact technology, Panga ya Saidi preserves osseous and shell artifacts, enabling broader considerations of the covariation between different spheres of material culture. Several environmental proxies contextualize the artifactual record of human behavior at Panga ya Saidi. We compare technological change between the Middle and Later Stone Age with on-site paleoenvironmental manifestations of wider climatic fluctuations in the Late Pleistocene. The principal distinguishing feature of Middle from Later Stone Age technology at Panga ya Saidi is the preference for fine-grained stone, coupled with the creation of small flakes (miniaturization). Our review of the Middle to Later Stone Age transition elsewhere in eastern Africa and across the continent suggests that this broader distinction between the two periods is in fact widespread. We suggest that the Later Stone Age represents new short use-life and multicomponent ways of using stone tools, in which edge sharpness was prioritized over durability.

Quantifying differences in hominin flaking technologies with 3D shape analysis.

Genetic and climate-driven estimates of past population dynamics are increasingly influential in broader models of hominin migration and adaptation, yet the contribution of stone artifact variability remains more contentious. Scientists are increasingly recognizing the potential of unretouched stone flakes ('flakes') in exploring existing models of hominin behavioral evolution. This is because flakes (1) were produced by all stone tool manufacturing groups in the past, (2) are abundant from the inception of the archaeological record up into the ethnographic present, and (3) preserve under most conditions. The statistical tools of 3D geometric morphometrics capture detailed approximations of flake form that are challenging to document with conventional artifact analyses. We analyze a collection of 717 3D scans of experimentally produced flakes from 5 production strategies that were practiced by hominins through large parts of the Pleistocene and that scientists have drawn on also to make demographic arguments about past human behavior (n = 45 reduction sequences, n = 3 knappers naive toward the study objectives). First, as a proof of concept, we demonstrate that we can estimate the strategies used to produce these flakes at a high success rate even when flakes from early stages of core reduction are included. We frame the significance of this finding against archaeological classifications from several key Middle Paleolithic assemblages in France (n = 4 sites, n = 28 layers, n = 16,467 flakes). Second, we show that 3D geometric morphometrics captures subtle differences in these strategies that influence flake formation on a flake-by-flake basis and that reflect decisions made by knappers about platform selection, preparation, and core-surface management. We explore the broader potential of our model with a cross-validation approach, and we describe a means of assessing flake form on a continuum wherein variability among assemblages separated by large expanses of space and time can be meaningfully explored.

Introducing platform surface interior angle (PSIA) and its role in flake formation, size and shape.

Four ways archaeologists have tried to gain insights into how flintknapping creates lithic variability are fracture mechanics, controlled experimentation, replication and attribute studies of lithic assemblages. Fracture mechanics has the advantage of drawing more directly on first principles derived from physics and material sciences, but its relevance to controlled experimentation, replication and lithic studies more generally has been limited. Controlled experiments have the advantage of being able to isolate and quantify the contribution of individual variables to knapping outcomes, and the results of these experiments have provided models of flake formation that when applied to the archaeological record of flintknapping have provided insights into past behavior. Here we develop a linkage between fracture mechanics and the results of previous controlled experiments to increase their combined explanatory and predictive power. We do this by documenting the influence of Herztian cone formation, a constant in fracture mechanics, on flake platforms. We find that the platform width is a function of the Hertzian cone constant angle and the geometry of the platform edge. This finding strengthens the foundation of one of the more influential models emerging from the controlled experiments. With additional work, this should make it possible to merge more of the experimental results into a more comprehensive model of flake formation.

Earliest known Oldowan artifacts at >2.58 Ma from Ledi-Geraru, Ethiopia, highlight early technological diversity.

The manufacture of flaked stone artifacts represents a major milestone in the technology of the human lineage. Although the earliest production of primitive stone tools, predating the genus Homo and emphasizing percussive activities, has been reported at 3.3 million years ago (Ma) from Lomekwi, Kenya, the systematic production of sharp-edged stone tools is unknown before the 2.58-2.55 Ma Oldowan assemblages from Gona, Ethiopia. The organized production of Oldowan stone artifacts is part of a suite of characteristics that is often associated with the adaptive grade shift linked to the genus Homo Recent discoveries from Ledi-Geraru (LG), Ethiopia, place the first occurrence of Homo ∼250 thousand years earlier than the Oldowan at Gona. Here, we describe a substantial assemblage of systematically flaked stone tools excavated in situ from a stratigraphically constrained context [Bokol Dora 1, (BD 1) hereafter] at LG bracketed between 2.61 and 2.58 Ma. Although perhaps more primitive in some respects, quantitative analysis suggests the BD 1 assemblage fits more closely with the variability previously described for the Oldowan than with the earlier Lomekwian or with stone tools produced by modern nonhuman primates. These differences suggest that hominin technology is distinctly different from generalized tool use that may be a shared feature of much of the primate lineage. The BD 1 assemblage, near the origin of our genus, provides a link between behavioral adaptations-in the form of flaked stone artifacts-and the biological evolution of our ancestors.

Site fragmentation, hominin mobility and LCT variability reflected in the early Acheulean record of the Okote Member, at Koobi Fora, Kenya.

From its initial appearance at ∼1.7 Ma, the Acheulean was prevalent through a vast chronological span of hominin behavioural evolution that lasted nearly 1.5 million years. The origins and production patterns of large bifacial cutting tools ('LCTs') - the marker of the Acheulean techno-complex - and the systematic changes in this behaviour through time are gaining increasing interest in paleoanthropology. Here we provide a synthesis of early Acheulean LCT variation in a landscape context by analysing assemblages from four different quasi-contemporaneous (∼1.4 Ma) sites from the Koobi Fora Formation. We characterize this variation using both 3D geometric morphometric and descriptive approaches. The expansive lateral exposures of fluvial and lacustrine sediments, as well as the associated tephrostratigraphy of the Koobi Fora Formation provide the landscape context that enables these comparative analyses. Our study demonstrates that when multiple contemporaneous early Acheulean localities are analysed together, a broader picture of LCT variability is elucidated. Four sites at Koobi Fora appear to represent a single system of lithic economy, characterized by a discrete trajectory of changes in LCT size and shape. These sites have ranges of LCT forms which appear to represent different but overlapping stages on a single reduction trajectory. Certain sites exhibit the full reduction trajectory while others exhibit only fragments of this trajectory. Other inter-site lithic proxies further complement these patterns in LCT variability. We explore patterns of site function, mobility and hominin landscape use, all of which may be suggestive of a depth of planning in early Acheulean hominins wherein technological activities were undertaken in substantial anticipation of future needs.

Publisher Correction: 78,000-year-old record of Middle and Later Stone Age innovation in an East African tropical forest.

The originally published version of this Article contained an error in Fig. 3, whereby an additional unrelated graph was overlaid on top of the magnetic susceptibility plot. Furthermore, the Article title contained an error in the capitalisation of 'Stone Age'. Both of these errors have now been corrected in both the PDF and HTML versions of the Article.

78,000-year-old record of Middle and Later stone age innovation in an East African tropical forest.

The Middle to Later Stone Age transition in Africa has been debated as a significant shift in human technological, cultural, and cognitive evolution. However, the majority of research on this transition is currently focused on southern Africa due to a lack of long-term, stratified sites across much of the African continent. Here, we report a 78,000-year-long archeological record from Panga ya Saidi, a cave in the humid coastal forest of Kenya. Following a shift in toolkits ~67,000 years ago, novel symbolic and technological behaviors assemble in a non-unilinear manner. Against a backdrop of a persistent tropical forest-grassland ecotone, localized innovations better characterize the Late Pleistocene of this part of East Africa than alternative emphases on dramatic revolutions or migrations.

What is Still Bay? Human biogeography and bifacial point variability.

'Still Bay' is the name given to a cultural phase within the southern African Middle Stone Age, which remains critical to our understanding of modern human behavioural evolution. Although represented in only a handful of sites, the Still Bay is widespread geographically and, at certain localities, persisted over a substantial period of time. Many studies have focused on tracing the temporal range and geographic reach of the Still Bay, as well as inferring degrees of early modern human demographic connectedness from these parameters. Variation within the Still Bay, relative to the accuracy with which it can be identified, has received considerably less attention. However, demographic models based on the spread of the Still Bay in space and time hinge on the reliability with which it can be recognized in the archaeological record. Here we document patterns of bifacial point shape and size variation in some key Still Bay assemblages, and analyse these patterns using the statistical shape analysis tools of geometric morphometrics. Morphological variation appears to be geographically structured and is driven by the spatial separation between north-eastern and south-western clusters of sites. We argue that allometric variation is labile and reflects environmentally driven differences in point reduction, whereas shape differences unrelated to size more closely reflect technological and cultural fragmentation. Our results suggest that the biogeographic structure of Middle Stone Age populations was complex during the period associated with the Still Bay, and provide little support for heightened levels of cultural interconnectedness between distantly separated groups at this time. We briefly discuss the implications of our findings for tracing classic techno-traditions in the Middle Stone Age record of southern Africa, and for inferring underpinning population dynamics from these patterns.

Diachronic Change within the Still Bay at Blombos Cave, South Africa.

Characteristically shaped bifacial points are stone artefacts with which the Middle Stone Age Still Bay techno-complex in Southern Africa is identified. Traditional approaches such as chaîne opératoire and two-dimensional metrics in combination with attribute analyses have been used to analyse variability within Still Bay point assemblages. Here we develop a protocol to extract and analyse high resolution 3-dimensional geometric morphometric information about Still Bay point morphology. We also investigate ways in which the independent variables of time, raw-material and tool size may be driving patterns of shape variation in the Blombos Cave point assemblage. We demonstrate that at a single, stratified Still Bay site points undergo significant modal changes in tool morphology and standardization. Our results caution against (1) treatment of the Still Bay as a static technological entity and (2) drawing demographic inferences stemming from grouping Still Bay point collections within the same cultural label.

Earliest evidence of dental caries manipulation in the Late Upper Palaeolithic.

Prehistoric dental treatments were extremely rare, and the few documented cases are known from the Neolithic, when the adoption of early farming culture caused an increase of carious lesions. Here we report the earliest evidence of dental caries intervention on a Late Upper Palaeolithic modern human specimen (Villabruna) from a burial in Northern Italy. Using Scanning Electron Microscopy we show the presence of striations deriving from the manipulation of a large occlusal carious cavity of the lower right third molar. The striations have a "V"-shaped transverse section and several parallel micro-scratches at their base, as typically displayed by cutmarks on teeth. Based on in vitro experimental replication and a complete functional reconstruction of the Villabruna dental arches, we confirm that the identified striations and the associated extensive enamel chipping on the mesial wall of the cavity were produced ante-mortem by pointed flint tools during scratching and levering activities. The Villabruna specimen is therefore the oldest known evidence of dental caries intervention, suggesting at least some knowledge of disease treatment well before the Neolithic. This study suggests that primitive forms of carious treatment in human evolution entail an adaptation of the well-known toothpicking for levering and scratching rather than drilling practices.

Documenting Differences between Early Stone Age Flake Production Systems: An Experimental Model and Archaeological Verification.

This study investigates morphological differences between flakes produced via "core and flake" technologies and those resulting from bifacial shaping strategies. We investigate systematic variation between two technological groups of flakes using experimentally produced assemblages, and then apply the experimental model to the Cutting 10 Mid -Pleistocene archaeological collection from Elandsfontein, South Africa. We argue that a specific set of independent variables--and their interactions--including external platform angle, platform depth, measures of thickness variance and flake curvature should distinguish between these two technological groups. The role of these variables in technological group separation was further investigated using the Generalized Linear Model as well as Linear Discriminant Analysis. The Discriminant model was used to classify archaeological flakes from the Cutting 10 locality in terms of their probability of association, within either experimentally developed technological group. The results indicate that the selected independent variables play a central role in separating core and flake from bifacial technologies. Thickness evenness and curvature had the greatest effect sizes in both the Generalized Linear and Discriminant models. Interestingly the interaction between thickness evenness and platform depth was significant and played an important role in influencing technological group membership. The identified interaction emphasizes the complexity in attempting to distinguish flake production strategies based on flake morphological attributes. The results of the discriminant function analysis demonstrate that the majority of flakes at the Cutting 10 locality were not associated with the production of the numerous Large Cutting Tools found at the site, which corresponds with previous suggestions regarding technological behaviors reflected in this assemblage.

Quantifying traces of tool use: a novel morphometric analysis of damage patterns on percussive tools.

Percussive technology continues to play an increasingly important role in understanding the evolution of tool use. Comparing the archaeological record with extractive foraging behaviors in nonhuman primates has focused on percussive implements as a key to investigating the origins of lithic technology. Despite this, archaeological approaches towards percussive tools have been obscured by a lack of standardized methodologies. Central to this issue have been the use of qualitative, non-diagnostic techniques to identify percussive tools from archaeological contexts. Here we describe a new morphometric method for distinguishing anthropogenically-generated damage patterns on percussive tools from naturally damaged river cobbles. We employ a geomatic approach through the use of three-dimensional scanning and geographical information systems software to statistically quantify the identification process in percussive technology research. This will strengthen current technological analyses of percussive tools in archaeological frameworks and open new avenues for translating behavioral inferences of early hominins from percussive damage patterns.

Early Pleistocene aquatic resource use in the Turkana Basin.

Evidence for the acquisition of nutritionally dense food resources by early Pleistocene hominins has implications for both hominin biology and behavior. Aquatic fauna may have comprised a source of highly nutritious resources to hominins in the Turkana Basin at ∼1.95 Ma. Here we employ multiple datasets to examine the issue of aquatic resource use in the early Pleistocene. This study focuses on four components of aquatic faunal assemblages (1) taxonomic diversity, (2) skeletal element proportion, (3) bone fragmentation and (4) bone surface modification. These components are used to identify associations between early Pleistocene aquatic remains and hominin behavior at the site of FwJj20 in the Koobi Fora Fm. (Kenya). We focus on two dominant aquatic species: catfish and turtles. Further we suggest that data on aquatic resource availability as well as ethnographic examples of aquatic resource use complement our observations on the archaeological remains from FwJj20. Aquatic food items provided hominins with a valuable nutritional alternative to an exclusively terrestrial resource base. We argue that specific advantages afforded by an aquatic alternative to terrestrial resources include (1) a probable reduction in required investment of energy relative to economic return in the form of nutritionally dense food items, (2) a decrease in the technological costs of resource acquisition, and (3) a reduced level of inter-specific competition associated with carcass access and an associated reduction of predation risk relative to terrestrial sources of food. The combined evidence from FwJj20 suggests that aquatic resources may have played a substantial role in early Pleistocene diets and these resources may have been overlooked in previous interpretations of hominin behavior.