New Blombos Cave evidence supports a multistep evolutionary scenario for the culturalization of the human body.

The emergence of technologies to culturally modify the appearance of the human body is a debated issue, with earliest evidence consisting of perforated marine shells dated between 140 and 60 ka at archaeological sites from Africa and western Asia. In this study, we submit unpublished marine and estuarine gastropods from Blombos Cave Middle Stone Age layers to taxonomic, taphonomic, technological, and use-wear analyses. We show that unperforated and naturally perforated eye-catching shells belonging to the species Semicassis zeylanica, Conus tinianus, and another Conus species, possibly Conus algoensis, were brought to the cave between 100 and 73 ka. At ca. 70 ka, a previously unrecorded marine gastropod, belonging to the species Tritia ovulata, was perforated by pecking and was worn as an ornamental object, isolated or in association with numerous intentionally perforated shells of the species Nassarius kraussianus. Fluctuations in sea level and consequent variations in the site-to-shoreline distances and landscape modifications during the Middle Stone Age have affected the availability of marine shells involved in symbolic practices. During the M3 and M2 Lower phases, with a sea level 50 m lower, the site was approximately 3.5 km away from the coast. In the later M2 Upper and M1 phases, with a sea level at -60 m, the distance increased to about 5.7 km. By the end of the M1 phase, when the site was abandoned, Blombos Cave was situated 18-30 km from the shoreline. We use the new Blombos evidence and a review of the latest findings from Africa and Eurasia to propose a testable ten-step evolutionary scenario for the culturalization of the human body with roots in the deep past.

Microstratigraphic preservation of ancient faunal and hominin DNA in Pleistocene cave sediments.

Ancient DNA recovered from Pleistocene sediments represents a rich resource for the study of past hominin and environmental diversity. However, little is known about how DNA is preserved in sediments and the extent to which it may be translocated between archaeological strata. Here, we investigate DNA preservation in 47 blocks of resin-impregnated archaeological sediment collected over the last four decades for micromorphological analyses at 13 prehistoric sites in Europe, Asia, Africa, and North America and show that such blocks can preserve DNA of hominins and other mammals. Extensive microsampling of sediment blocks from Denisova Cave in the Altai Mountains reveals that the taxonomic composition of mammalian DNA differs drastically at the millimeter-scale and that DNA is concentrated in small particles, especially in fragments of bone and feces (coprolites), suggesting that these are substantial sources of DNA in sediments. Three microsamples taken in close proximity in one of the blocks yielded Neanderthal DNA from at least two male individuals closely related to Denisova 5, a Neanderthal toe bone previously recovered from the same layer. Our work indicates that DNA can remain stably localized in sediments over time and provides a means of linking genetic information to the archaeological and ecological records on a microstratigraphic scale.

Later Stone Age human hair from Vaalkrans Shelter, Cape Floristic Region of South Africa, reveals genetic affinity to Khoe groups.

Previous studies show that the indigenous people of the southern Cape of South Africa were dramatically impacted by the arrival of European colonists starting ~400 years ago and their descendants are today mixed with Europeans and Asians. To gain insight on the occupants of the Vaalkrans Shelter located at the southernmost tip of Africa, we investigated the genetic make-up of an individual who lived there about 200 years ago. We further contextualize the genetic ancestry of this individual among prehistoric and current groups. From a hair sample excavated at the shelter, which was indirectly dated to about 200 years old, we sequenced the genome (1.01 times coverage) of a Later Stone Age individual. We analyzed the Vaalkrans genome together with genetic data from 10 ancient (pre-colonial) individuals from southern Africa spanning the last 2000 years. We show that the individual from Vaalkrans was a man who traced ~80% of his ancestry to local southern San hunter-gatherers and ~20% to a mixed East African-Eurasian source. This genetic make-up is similar to modern-day Khoekhoe individuals from the Northern Cape Province (South Africa) and Namibia, but in the southern Cape, the Vaalkrans man's descendants have likely been assimilated into mixed-ancestry "Coloured" groups. The Vaalkrans man's genome reveals that Khoekhoe pastoralist groups/individuals lived in the southern Cape as late as 200 years ago, without mixing with non-African colonists or Bantu-speaking farmers. Our findings are also consistent with the model of a Holocene pastoralist migration, originating in Eastern Africa, shaping the genomic landscape of historic and current southern African populations.

The Howiesons Poort lithic sequence of Klipdrift Shelter, southern Cape, South Africa.

Howiesons Poort (HP) sites, over the past decades, have provided exceptional access to anthropogenic remains that are enhancing our understanding of early modern human behaviour during the Middle Stone Age in southern Africa. Here, we analyse the technological and typological trends in the lithic record that form part of these behaviours, based on the HP sequence recently excavated at Klipdrift Shelter, located on the southern Cape coast of South Africa. This study contributes to enhance knowledge on the mechanisms of changes that occurred during the transition to the post-HP. Despite patterns of continuity observed, notably for core reduction methods, the seven successive lithic assemblages show significant changes in the typological characteristics and raw material selection but also in the relative importance of blade production over time. However, these changes are not necessarily synchronic and occur either as gradual processes or as abrupt technological shifts. Consequently, we cross-examine the association between the lithic phasing and other anthropogenic remains within the HP sequence at Klipdrift Shelter. We explore the implications of these patterns of changes in terms of cultural behaviours and population dynamics during the HP and we highlight the relationship between the different phases of the HP sequence at Klipdrift Shelter and those from other South African HP sites.

An abstract drawing from the 73,000-year-old levels at Blombos Cave, South Africa.

Depictive and abstract representations produced by drawing-known from Europe, Africa and Southeast Asia after 40,000 years ago-are a prime indicator of modern cognition and behaviour1. Here we report a cross-hatched pattern drawn with an ochre crayon on a ground silcrete flake recovered from approximately 73,000-year-old Middle Stone Age levels at Blombos Cave, South Africa. Our microscopic and chemical analyses of the pattern confirm that red ochre pigment was intentionally applied to the flake with an ochre crayon. The object comes from a level associated with stone tools of the Still Bay techno-complex that has previously yielded shell beads, cross-hatched engravings on ochre pieces and a variety of innovative technologies2-5. This notable discovery pre-dates the earliest previously known abstract and figurative drawings by at least 30,000 years. This drawing demonstrates the ability of early Homo sapiens in southern Africa to produce graphic designs on various media using different techniques.

Identifying early modern human ecological niche expansions and associated cultural dynamics in the South African Middle Stone Age.

The archaeological record shows that typically human cultural traits emerged at different times, in different parts of the world, and among different hominin taxa. This pattern suggests that their emergence is the outcome of complex and nonlinear evolutionary trajectories, influenced by environmental, demographic, and social factors, that need to be understood and traced at regional scales. The application of predictive algorithms using archaeological and paleoenvironmental data allows one to estimate the ecological niches occupied by past human populations and identify niche changes through time, thus providing the possibility of investigating relationships between cultural innovations and possible niche shifts. By using such methods to examine two key southern Africa archaeological cultures, the Still Bay [76-71 thousand years before present (ka)] and the Howiesons Poort (HP; 66-59 ka), we identify a niche shift characterized by a significant expansion in the breadth of the HP ecological niche. This expansion is coincident with aridification occurring across Marine Isotope Stage 4 (ca. 72-60 ka) and especially pronounced at 60 ka. We argue that this niche shift was made possible by the development of a flexible technological system, reliant on composite tools and cultural transmission strategies based more on "product copying" rather than "process copying." These results counter the one niche/one human taxon equation. They indicate that what makes our cultures, and probably the cultures of other members of our lineage, unique is their flexibility and ability to produce innovations that allow a population to shift its ecological niche.

Subsistence strategies during the Late Pleistocene in the southern Cape of South Africa: Comparing the Still Bay of Blombos Cave with the Howiesons Poort of Klipdrift Shelter.

The Still Bay (SB) and Howiesons Poort (HP) were two significant techno-complexes in the Middle Stone Age and key periods in the expression of behavioral complexity. In this study, we compare the recently excavated fauna from the SB layers at Blombos Cave (BBC) with that from the HP levels at Klipdrift Shelter (KDS) in the southern Cape of South Africa. We consider our findings in the framework of recent models for early human subsistence behavior. In particular, we link our study with models involving resource intensification to examine whether foraging strategies in the HP were more or less intensive than those in the SB. Based on our criteria used to assess intensification-the exploitation of low-ranked prey, the processing of low-utility elements, transport decisions, and occupational intensity-intensive subsistence strategies are more evident at KDS than BBC. Our results suggest that low-ranked elements were processed more heavily and diet breath was broader at KDS than at BBC. However, foraging ranges may have been more extensive at BBC than at KDS. Taphonomic data also suggests that the SB at BBC was a low-intensity, sporadically occupied period in contrast to the high-intensity occupations during the HP at KDS. We argue that this may be related to differences in mobility and residential patterns between these techno-complexes.

Early Evidence for the Extensive Heat Treatment of Silcrete in the Howiesons Poort at Klipdrift Shelter (Layer PBD, 65 ka), South Africa.

Heating stone to enhance its flaking qualities is among the multiple innovative adaptations introduced by early modern human groups in southern Africa, in particular during the Middle Stone Age Still Bay and Howiesons Poort traditions. Comparatively little is known about the role and impact of this technology on early modern human behaviors and cultural expressions, due, in part, to the lack of comprehensive studies of archaeological assemblages documenting the heat treatment of stone. We address this issue through an analysis of the procedure used for heating and a technological analysis of a lithic assemblage recovered from one Howiesons Poort assemblage at Klipdrift Shelter (southern Cape, South Africa). The resulting data show extensive silcrete heat treatment, which adds a new dimension to our understanding of fire-related behaviors during the Howiesons Poort, highlighting the important role played by a heat treatment stage in the production of silcrete blades. These results are made possible by our new analytical procedure that relies on the analysis of all silcrete artifacts. It provides direct evidence of a controlled use of fire which took place during an early stage of core exploitation, thereby impacting on all subsequent stages of the lithic chaîne opératoire, which, to date, has no known equivalent in the Middle Stone Age or Middle Paleolithic record outside of southern Africa.

The Small Mammal Sequence from the c. 76 - 72 ka Still Bay Levels at Blombos Cave, South Africa - Taphonomic and Palaeoecological Implications for Human Behaviour.

The Still Bay, c. 76-72 ka, a prominent techno-tradition during the Middle Stone Age of southern Africa, has yielded innovative technologies, symbolic material culture, and shows evidence of expansion of hunting techniques and subsistence strategies. In this paper we present the results of the first systematic, taphonomic and palaeoenvironmental study of micromammals from the Still Bay levels at Blombos Cave. Our taphonomic analysis indicates that the micromammals were accumulated by avian predators occupying the cave. Post-depositional processes affecting the micromammal assemblage include organic waste decomposition and conditions associated with a limestone cave environment. The palaeoenvironmental reconstruction shows that Marine Isotope Stage (MIS) 5a at Blombos Cave had diverse micromammal communities occupying a variety of habitats and with rainfall pattern equal to present. The transition from MIS 5a to 4 is indicated by less diverse micromammal assemblages, increase in grassland and scrub vegetation, shifts in seasonal precipitation, and a decline in shrubs associated with fynbos. The onset of the glacial conditions associated with MIS 4 is visible in the micromammal assemblage. However humans occupying Blombos Cave during this c. 5 ka period showed an ability to cope with changing environmental conditions and were able to adapt and utilise a variety of available resources.

Climate, Environment and Early Human Innovation: Stable Isotope and Faunal Proxy Evidence from Archaeological Sites (98-59ka) in the Southern Cape, South Africa.

The Middle Stone Age (MSA) of southern Africa, and in particular its Still Bay and Howiesons Poort lithic traditions, represents a period of dramatic subsistence, cultural, and technological innovation by our species, Homo sapiens. Climate change has frequently been postulated as a primary driver of the appearance of these innovative behaviours, with researchers invoking either climate instability as a reason for the development of buffering mechanisms, or environmentally stable refugia as providing a stable setting for experimentation. Testing these alternative models has proved intractable, however, as existing regional palaeoclimatic and palaeoenvironmental records remain spatially, stratigraphically, and chronologically disconnected from the archaeological record. Here we report high-resolution records of environmental shifts based on stable carbon and oxygen isotopes in ostrich eggshell (OES) fragments, faunal remains, and shellfish assemblages excavated from two key MSA archaeological sequences, Blombos Cave and Klipdrift Shelter. We compare these records with archaeological material remains in the same strata. The results from both sites, spanning the periods 98-73 ka and 72-59 ka, respectively, show significant changes in vegetation, aridity, rainfall seasonality, and sea temperature in the vicinity of the sites during periods of human occupation. While these changes clearly influenced human subsistence strategies, we find that the remarkable cultural and technological innovations seen in the sites cannot be linked directly to climate shifts. Our results demonstrate the need for scale-appropriate, on-site testing of behavioural-environmental links, rather than broader, regional comparisons.

Intra-Site Variability in the Still Bay Fauna at Blombos Cave: Implications for Explanatory Models of the Middle Stone Age Cultural and Technological Evolution.

To explain cultural and technological innovations in the Middle Stone Age (MSA) of southern Africa, scholars invoke several factors. A major question in this research theme is whether MSA technocomplexes are adapted to a particular set of environmental conditions and subsistence strategies or, on the contrary, to a wide range of different foraging behaviours. While faunal studies provide key information for addressing these factors, most analyses do not assess intra-technocomplex variability of faunal exploitation (i.e. variability within MSA phases). In this study, we assess the spatial variability of the Still Bay fauna in one phase (M1) of the Blombos Cave sequence. Analyses of taxonomic composition, taphonomic alterations and combustion patterns reveal important faunal variability both across space (lateral variation in the post-depositional history of the deposits, spatial organisation of combustion features) and over time (fine-scale diachronic changes throughout a single phase). Our results show how grouping material prior to zooarchaeological interpretations (e.g. by layer or phase) can induce a loss of information. Finally, we discuss how multiple independent subdivisions of archaeological sequences can improve our understanding of both the timing of different changes (for example in technology, culture, subsistence, environment) and how they may be inter-related.

Techno-Cultural Characterization of the MIS 5 (c. 105 - 90 Ka) Lithic Industries at Blombos Cave, Southern Cape, South Africa.

Blombos Cave is well known as an important site for understanding the evolution of symbolically mediated behaviours among Homo sapiens during the Middle Stone Age, and during the Still Bay in particular. The lower part of the archaeological sequence (M3 phase) contains 12 layers dating to MIS 5 with ages ranging from 105 to 90 ka ago (MIS 5c to 5b) that provide new perspectives on the technological behaviour of these early humans. The new data obtained from our extensive technological analysis of the lithic material enriches our currently limited knowledge of this time period in the Cape region. By comparing our results with previously described lithic assemblages from sites south of the Orange River, we draw new insights on the extent of the techno-cultural ties between these sites and the M3 phase at Blombos Cave and highlight the importance of this phase within the Middle Stone Age cultural stratigraphy.

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.

The Still Bay and Howiesons Poort at Sibudu and Blombos: Understanding Middle Stone Age Technologies.

The classification of archaeological assemblages in the Middle Stone Age of South Africa in terms of diversity and temporal continuity has significant implications with respect to recent cultural evolutionary models which propose either gradual accumulation or discontinuous, episodic processes for the emergence and diffusion of cultural traits. We present the results of a systematic technological and typological analysis of the Still Bay assemblages from Sibudu and Blombos. A similar approach is used in the analysis of the Howiesons Poort (HP) assemblages from Sibudu seen in comparison with broadly contemporaneous assemblages from Rose Cottage and Klasies River Cave 1A. Using our own and published data from other sites we report on the diversity between stone artifact assemblages and discuss to what extent they can be grouped into homogeneous lithic sets. The gradual evolution of debitage techniques within the Howiesons Poort sequence with a progressive abandonment of the HP technological style argues against the saltational model for its disappearance while the technological differences between the Sibudu and Blombos Still Bay artifacts considerably weaken an interpretation of similarities between the assemblages and their grouping into the same cultural unit. Limited sampling of a fragmented record may explain why simple models of cultural evolution do not seem to apply to a complex reality.

Nutritional values of tortoises relative to ungulates from the Middle Stone Age levels at Blombos Cave, South Africa: implications for foraging and social behaviour.

The site of Blombos Cave (BBC), Western Cape, South Africa has been a strong contributor to establishing the antiquity of important aspects of modern human behaviour, such as early symbolism and technological complexity. However, many linkages between Middle Stone Age (MSA) behaviour and the subsistence record remain to be investigated. Understanding the contribution of small fauna such as tortoises to the human diet is necessary for identifying shifts in overall foraging strategies as well as the collecting and processing behaviour of individuals unable to participate in large-game hunting. This study uses published data to estimate the number of calories present in tortoises as well as ungulates of different body size classes common at South African sites. A single tortoise (Chersina angulata) provides approximately 3332 kJ (796 kcal) of calories in its edible tissues, which is between 20 and 30% of the daily energetic requirements for an active adult (estimated between 9360 kJ [3327 kcal] and 14,580 kJ [3485 kcal] per day). Because they are easy to process, this would have made tortoises a highly-ranked resource, but their slow growth and reproduction makes them susceptible to over-exploitation. Zooarchaeological abundance data show that during the ca. 75 ka (thousands of years) upper Still Bay M1 phase at BBC, tortoises contributed twice as many calories to the diet relative to ungulates than they did during the ca. 100 ka lower M3 phase. However, in spite of the abundance of their fossils, their absolute caloric contribution relative to ungulates remained modest in both phases. At the end of the site's MSA occupation history, human subsistence strategies shifted to emphasise high-return large hunted mammals, which likely precipitated changes in the social roles of hunters and gatherers during the Still Bay.

Thinking strings: additional evidence for personal ornament use in the Middle Stone Age at Blombos Cave, South Africa.

Here we report on newly identified beads recovered from four Middle Stone Age levels at Blombos Cave and, in particular, on a cluster of 24 perforated Nassarius kraussianus shells that probably originate from a single beadwork. Contextual information, morphometric, technological and use-wear analysis of the 68 published beads and those recently found, coupled with experimental reproduction of wear patterns, allow us to reconstruct the most probable way in which the N. kraussianus shells were strung. The results reveal unexpected regularities but also variability through the various levels that we interpret as resulting from changes in beadwork manufacture and design over time. The Blombos Cave beads may document one of the first examples of changes in social norms affecting the production and design of symbolic material culture.

A 100,000-year-old ochre-processing workshop at Blombos Cave, South Africa.

The conceptual ability to source, combine, and store substances that enhance technology or social practices represents a benchmark in the evolution of complex human cognition. Excavations in 2008 at Blombos Cave, South Africa, revealed a processing workshop where a liquefied ochre-rich mixture was produced and stored in two Haliotis midae (abalone) shells 100,000 years ago. Ochre, bone, charcoal, grindstones, and hammerstones form a composite part of this production toolkit. The application of the mixture is unknown, but possibilities include decoration and skin protection.

Taphonomic analysis of the Middle Stone Age larger mammal faunal assemblage from Blombos Cave, southern Cape, South Africa.

A detailed taphonomic analysis is reported for a sample of the larger mammalian faunal assemblage (>4.5 kg live body weight) from Blombos Cave. The analysis provides an assessment of human involvement in the accumulation and modification of the faunal assemblage, and precedes equally detailed analyses and separate reports of Middle Stone Age (MSA) butchery, transport, and hunting behaviour. At Blombos, there are clear differences in the relative abundances of ungulate body size classes, with the lower MSA phases (upper/lower M2 and M3) showing a high representation of size 1 ungulates relative to the most recent MSA phase (M1). The bones from the earliest MSA phase (M3) have not undergone much post-depositional fragmentation, in contrast to fragments from more recent phases (M1 and upper M2). Much of this variability can be attributed to more burning activity and trampling during M1 and upper M2, which could indicate more intensive occupation. Bone surfaces are variably preserved, with high levels of exfoliation in the most recent two phases. Surface modification analyses revealed high proportions of human modification throughout the sequence, indicating that MSA humans were responsible for accumulating most of the larger mammals. After discard, the bones were modified by scavenging carnivores, leading to a moderate amount of density-mediated destruction and tooth-marking. Carnivores independently accumulated some of the smaller ungulates, mainly in the form of partially-digested remains. Raptorial birds are not implicated as major faunal accumulators. The results from Blombos are directly comparable with analogous datasets from two other sites in the Western Cape (Pinnacle Point Cave 13B and Die Kelders Cave 1). Such comparisons demonstrate that MSA faunal assemblages from nearby coastal sites have complex and different taphonomic histories both within and between sites. Because the human occupants were a major part of these processes, MSA subsistence behaviour and site use was also quite variable over time and space.

Early use of pressure flaking on lithic artifacts at Blombos Cave, South Africa.

Pressure flaking has been considered to be an Upper Paleolithic innovation dating to ~20,000 years ago (20 ka). Replication experiments show that pressure flaking best explains the morphology of lithic artifacts recovered from the ~75-ka Middle Stone Age levels at Blombos Cave, South Africa. The technique was used during the final shaping of Still Bay bifacial points made on heat-treated silcrete. Application of this innovative technique allowed for a high degree of control during the detachment of individual flakes, resulting in thinner, narrower, and sharper tips on bifacial points. This technology may have been first invented and used sporadically in Africa before its later widespread adoption.