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Africa’s Middle Stone Age preserves sporadic evidence for novel behaviours among early modern humans, prompting a range of questions about the influence of social and environmental factors on patterns of human behavioural evolution. Here we document a suite of novel adaptations dating approximately 92–80 thousand years before the present at the archaeological site Varsche Rivier 003 (VR003), located in southern Africa’s arid Succulent Karoo biome. Distinctive innovations include the production of ostrich eggshell artefacts, long-distance transportation of marine molluscs and systematic use of heat shatter in stone tool production, none of which occur in coeval assemblages at sites in more humid, well-studied regions immediately to the south. The appearance of these novelties at VR003 corresponds with a period of reduced regional wind strength and enhanced summer rainfall, and all of them disappear with increasing winter rainfall dominance after 80 thousand years before the present, following which a pattern of technological similarity emerges at sites throughout the broader region. The results indicate complex and environmentally contingent processes of innovation and cultural transmission in southern Africa during the Middle Stone Age. acceptedVersion

Research interest in patterns of cultural transmission in the southern African Palaeolithic is complicated by poor resolution of site occupation, technological behaviour, and underlying environmental conditions. Here, we describe the Pleistocene Later Stone Age sequence from Klipfonteinrand rock shelter, dating 22.3–13.4 ka, in light of local and regional data. Analysis of chronology, stratigraphy, wood charcoal, fauna, and lithics suggest an early termination for the Robberg technocomplex, with a locally adaptive response to declining humidity emerging by ~ 16 ka. Subsequent Oakhurst-aligned systems are in place by ~ 14.4 ka, associated with diminished intensity of site use and drier conditions. Technological responses in the post-Robberg interval appear to differ between sites in coastal and interior areas, despite evidence for interaction between the two in this period. The results suggest a complex mix of local adaptation and inter-regional transmission through MIS 2.

The African Middle Pleistocene (781–126 ka) is a key period for human evolution, witnessing both the origin of the modern human lineage and the lithic turnover from Earlier Stone Age (ESA) Acheulean bifacial tools to Middle Stone Age (MSA) prepared core and point technologies. This ESA/MSA transition is interpreted as representing changing landscape use with greater foraging distances and more active hunting strategies. So far, these behavioral inferences are mainly based on the extensive stone tool record, with only a minor role for site-based and regional faunal studies. To provide additional insights into these behavioral changes, this paper details a pan-African metastudy of 63 Middle Pleistocene faunal assemblages from 40 sites. A hierarchical classification system identified 26 well-contextualized assemblages with quantitative paleontological and/or zooarcheological data available for detailed comparative analyses and generalized linear mixed modeling. Modeling of ungulate body size classes structured around three dimensions (context, antiquity and technology) illustrates no one-to-one correlation between changes in lithic technology (Acheulean vs. MSA) and changes in prey representation. All assessed faunal assemblages are dominated by medium-sized bovids, and variations between smaller and larger body size classes are linked to site context (cave vs. open-air), with an increase in cave sites during the Middle Pleistocene. Current data do not signal a broadening of the hominin dietary niche during the Middle Pleistocene; no meaningful variation was visible in the exploitation of smaller-sized bovids or dangerous game, with coastal resources exploited when available. Proportions of anthropogenic bone surface modifications, and hence carcass processing intensity, do increase over time although more zooarcheological data is crucial before making behavioral inferences. Overall, this paper illustrates the potential of broad scale comparative faunal analyses to provide additional insights into processes of human behavioral evolution and the mechanisms underlying patterns of technological, chronological and contextual change.

Evidence for complex behaviours appears sporadically through the Middle Stone Age of Africa, leaving unclear the major factors shaping the evolution of human behaviour. Here we present evidence for a novel suite of adaptations in the arid Knersvlake region of southern Africa that were deployed during a specific set of environmental conditions dating 90-80 000 years before present, at the archaeological site Varsche Rivier 003 (VR003). This includes the earliest production of artefacts from ostrich eggshell, long-distance transportation of economic marine molluscs, and stone tool technologies focussed on manufacture of small flakes and blades from deliberately heat fractured blocks of silcrete. Similar evidence is absent from coeval assemblages at sites in less arid areas immediately to the south, and subsequently disappears from VR003 with the onset of more humid conditions. The results indicate complex cultural dynamics in southern Africa during the Middle Stone Age, and suggest that marginal subsistence conditions fostered distinctive technological and subsistence innovations.

Africa's Middle Stone Age preserves sporadic evidence for novel behaviours among early modern humans, prompting a range of questions about the influence of social and environmental factors on patterns of human behavioural evolution. Here we document a suite of novel adaptations dating approximately 92-80 thousand years before the present at the archaeological site Varsche Rivier 003 (VR003), located in southern Africa's arid Succulent Karoo biome. Distinctive innovations include the production of ostrich eggshell artefacts, long-distance transportation of marine molluscs and systematic use of heat shatter in stone tool production, none of which occur in coeval assemblages at sites in more humid, well-studied regions immediately to the south. The appearance of these novelties at VR003 corresponds with a period of reduced regional wind strength and enhanced summer rainfall, and all of them disappear with increasing winter rainfall dominance after 80 thousand years before the present, following which a pattern of technological similarity emerges at sites throughout the broader region. The results indicate complex and environmentally contingent processes of innovation and cultural transmission in southern Africa during the Middle Stone Age.

Exploring the role of changing climates in human evolution is currently impeded by a scarcity of climatic information at the same temporal scale as the human behaviors documented in archaeological sites. This is mainly caused by high uncertainties in the chronometric dates used to correlate long-term climatic records with archaeological deposits. One solution is to generate climatic data directly from archaeological materials representing human behavior. Here we use oxygen isotope measurements of Bos/Bison tooth enamel to reconstruct summer and winter temperatures in the Late Pleistocene when Neandertals were using the site of La Ferrassie. Our results indicate that, despite the generally cold conditions of the broader period and despite direct evidence for cold features in certain sediments at the site, Neandertals used the site predominantly when climatic conditions were mild, similar to conditions in modern day France. We suggest that due to millennial scale climate variability, the periods of human activity and their climatic characteristics may not be representative of average conditions inferred from chronological correlations with long-term climatic records. These results highlight the importance of using direct routes, such as the high-resolution archives in tooth enamel from anthropogenically accumulated faunal assemblages, to establish climatic conditions at a human scale.

International audience; Five nearly identical fragments of specialized bone tools, interpreted as lissoirs (French for "smoothers"), have been found at two Middle Paleolithic sites in southwest France. The finds span three separate archaeological deposits, suggesting continuity in the behavior of late Neandertals. Using standard morphological assessments, we determined that the lissoirs were produced on ribs of medium-sized ungulates. However, since these bones are highly fragmented and anthropogenically modified, species determinations were challenging. Also, conservative curation policy recommends minimizing destructive sampling of rare, fragile, or small artifacts for molecular identification methods. To better understand raw material selection for these five lissoirs, we reassess their taxonomy using a non-destructive ZooMS methodology based on triboelectric capture of collagen. We sampled four storage containers and obtained identifiable MALDI-TOF MS collagen fingerprints, all indicative of the same taxonomic clade, which includes aurochs and bison (Bos sp. and Bison sp.). The fifth specimen, which was stored in a plastic bag, provided no useful MALDI-TOF MS spectra. We show that the choice of large bovid ribs in an archaeological layer dominated by reindeer (Rangifer tarandus) demonstrates strategic selection by these Neandertals. Furthermore, our results highlight the value of a promising technique for the non-destructive analysis of bone artifacts.

Increasingly researchers have employed confocal microscopy and 3D surface texture analysis to assess bone surface modifications in an effort to understand ancient behavior. However, quantitative comparisons between the surfaces of purported archaeological bone tools and experimentally manufactured and used bones are complicated by taphonomic processes affecting ancient bone. Nonetheless, it may be reasonable to assume that bones within the same deposits are altered similarly and thus these alterations are quantifiable. Here we show how unworked bones can be used to quantify the taphonomic effect on bone surfaces and how this effect can then be controlled for and incorporated into an analysis for evaluating the modified surfaces of purported bone tools. To assess the baseline taphonomy of Middle Paleolithic archaeological deposits associated with typologically identified bone artifacts, specifically lissoirs, we directly compare the surface textures of ancient and modern unworked ribs. We then compare the ancient unworked ribs and lissoirs to assess their differences and predict the ancient artifacts’ original surface state using a multilevel multivariate Bayesian model. Our findings demonstrate that three of five tested surface texture parameters (Sa, Spc, and IsT) are useful for distinguishing surface type. Our model predictions show that lissoirs tend to be less rough, have more rounded surface peaks, and exhibit more directionally oriented surfaces. These characteristics are likely due to anthropogenic modifications and would have been more pronounced at deposition. Quantifying taphonomic alterations moves us one step closer to accurately assessing how bone artifacts were made and used in the ancient past.

The grand abri at La Ferrassie (France) has been a key site for Palaeolithic research since the early part of the 20th century. It became the eponymous site for one variant of Middle Palaeolithic stone tools, and its sequence was used to define stages of the Aurignacian, an early phase of the Upper Palaeolithic. Several Neanderthal remains, including two relatively intact skeletons, make it one of the most important sites for the study of Neanderthal morphology and one of the more important data sets when discussing the Neanderthal treatment of the dead. However, the site has remained essentially undated. Our goal here is to provide a robust chronological framework of the La Ferrassie sequence to be used for broad regional models about human behaviour during the late Middle to Upper Palaeolithic periods. To achieve this goal, we used a combination of modern excavation methods, extensive geoarchaeological analyses, and radiocarbon dating. If we accept that Neanderthals were responsible for the Châtelperronian, then our results suggest an overlap of ca. 1600 years with the newly arrived Homo sapiens found elsewhere in France., Open access funding enabled and organized by Projekt DEAL.

Ostrich ( Struthio spp.) eggshells are present in archaeological assemblages in many regions of Africa and Asia. However, and unlike other artifacts and ecofacts, there are no standardized guidelines for observing and recording non-ornamental ostrich eggshell. Here, we review prior research that focuses on facets of the taphonomic history of ostrich eggshell assemblages, and we document results from our actualistic studies of the changes in color that occur when ostrich eggshells are heated. We further propose some guidelines for recording ostrich eggshell in archaeological contexts, which include burning categories and quantification methods. These guidelines are intended to facilitate the development of large, comparative, and standardized ostrich eggshell data sets that will contribute to our understanding of site taphonomy and past human behavior.

Accumulating genomic, fossil and archaeological data from Africa have led to a renewed interest in models of modern human origins. However, such discussions are often discipline-specific, with limited integration of evidence across the different fields. Further, geneticists typically require explicit specification of parameters to test competing demographic models, but these have been poorly outlined for some scenarios. Here, we describe four possible models for the origins of Homo sapiens in Africa based on published literature from paleoanthropology and human genetics. We briefly outline expectations for data patterns under each model, with a special focus on genetic data. Additionally, we present schematics for each model, doing our best to qualitatively describe demographic histories for which genetic parameters can be specifically attached. Finally, it is our hope that this perspective provides context for discussions of human origins in other manuscripts presented in this special issue.

International audience; "Use-wear analysis provides a means of studying traces produced on animal bone during manufacture and use in an effort to reconstruct these processes. Often, these analyses are qualitative and based on experience and expertise. Previous studies have focused on interpreting final traces, but little is known about how these traces develop and change over time. We propose the use of an innovative quantitative method for studying bone surface traces that aims to reduce any unreliable or non-replicable results that can confound more traditional qualitative analyses. We seek to understand the basics of use-wear formation over Time by taking incremental molds of bone specimens subjected to a controlled, mechanical experiment. This study assesses how bone wears during extended use on three Material types (fresh skin, processed leather, or dry bark), from three initial Manufacturing states (unworked, ground with sandstone, or scraped with flint). With data obtained from a confocal disc-scanning microscope, we then apply 3D surface texture analysis using ISO 25178 parameters: surface roughness [Sa], autocorrelation length [Sal], peak curvature [Spc], and upper material ratio [Smr1]. We employ a multilevel multivariate Bayesian model to explain parameter variation under experimental conditions. Our findings show how duration of use strongly affects the transformation of the bone’s surface. Unworked bone is completely distinguishable from bone used for long time intervals and those modified by scraping. Interestingly, material wear does not often produce type-specific traces, but does affect the rate of bone alteration and how it is transformed. Specifically, fresh skin transforms bone at a faster rate than other materials. This novel quantitative and experimental approach enhances our understanding of the use of bone as a raw material for making and using tools and provides a foundation for future exploration of archaeological materials and questions." (source éditeur)

We report on excavations of a small rock shelter — Putslaagte 8 (PL8) — located on the arid interior fringe of South Africa’s Fynbos biome. The shelter preserves a long sequence of Holocene and late Pleistocene occupation dating back beyond 75,000 years BP. This paper presents data on the technological, faunal and chronological sequence. Occupation is markedly pulsed and includes three late Pleistocene Later Stone Age (LSA) units (macrolithic, Robberg and early LSA), as well as several distinct Middle Stone Age (MSA) components from Marine Isotope Stage (MIS) 3–5. Pulsing may reflect the arid and possibly marginal environments in which the shelter is situated, and to that end some elements of the sequence contrast with occupational patterns towards the coast. Viewed in a regional setting PL8 suggests: 1) complementarity of resource movements between the coast and interior in terminal MIS 2; 2) distinctions in material selection, and possibly technology, between the coast and interior in earlier MIS 2; 3) an MSA lasting to at least 40,000 years before present; 4) a weak Howiesons Poort and post-Howiesons Poort in the interior; 5) possibly distinct periods of denticulate manufacture within the MIS 5 MSA; 6) highly localised patterns of material acquisition in the earlier MSA.

Humans and animals share a universal and intimate relationship that stretches from the earliest appearance of our lineage, through our prehistory, and into history and the modern era. Consequently, zooarchaeology -- the analysis of animal remains from archaeological sites -- transcends all cultural, temporal, and geographic boundaries, and therefore can be used to address questions spanning the breadth of archaeology. Over the past decades, the field has grown tremendously, from a small group of specialized practitioners to a large sub-discipline that is integrated with the larger field's research directions. Zooarchaeological analyses have addressed questions ranging from forager ecology to the processes of domestication to how animals function in the social realm. In each instance, methodological developments have been crucial in allowing existing questions to be tackled and new questions to be posed. One of the largest changes in zooarchaeology is a shift from considering how the environment has shaped human societies to considering how humans have altered the environment, a process that began deep in our prehistory. Applied zooarchaeology looks towards using zooarchaeological data to address questions relevant to conservation biology, two fields that are becoming increasingly integrated. Standardization of data collection and presentation, open access to these data, and global access remain challenges for the future.

In a paper based on recent excavations and analysis of recovered material at the French Middle Paleolithic site of La Chapelle-aux-Saints, Rendu et al. (2014) concluded that there is sufficient evidence to support the long-held interpretation that the Neandertal remains found in the bouffia Bonneval locality represents an intentional Neandertal burial. This paper critically examines their data and arguments in relation to criteria that can be used to provide an objective evaluation of such evidence. In each case, the evidence from La Chapelle-aux-Saints either fails to meet these criteria or supports other interpretations equally well. As a result, this site fails to provide unequivocal evidence in support of the notion that Neandertals intentionally interred their dead, whether in any ritualistic or symbolic context or not.

The later Middle Stone Age of southern Africa witnesses a number of important changes in lithic technology including the early appearance of bifacial point and microlithic systems. Though radiometric ages for these changes remain contested, they can in places be reconciled with elements of climatic variation. This paper examines the organization of provisioning systems through four successive industrial phases – the Still Bay, early Howiesons Poort, later Howiesons Poort, and post-Howiesons Poort – using data from rock-shelters and open sites. We show that the dominant means of technological delivery shifted from individual to place provisioning and back to individual provisioning through these industries, likely reflecting variation in humidity through the last glacial. We also show that the apparent abundance of sites and richness of assemblages in industries such as the Howiesons Poort may be in part a consequence of a research focus on rock-shelters; we found limited evidence for Howiesons Poort sites in the open. This is in contrast to the other industries, most clearly the Still Bay, for which we identified several large open-air gearing-up locations. Finally, we suggest that while interrogation of the archaeological record at the industry level is viable, it also masks behaviorally meaningful variation that should serve to encourage finer-scaled analyses.

International audience; The timing and location of the emergence of our species and of associated behavioural changes are crucial for our understanding of human evolution. The earliest fossil attributed to a modern form of Homo sapiens comes from eastern Africa and is approximately 195 thousand years old, therefore the emergence of modern human biology is commonly placed at around 200 thousand years ago. The earliest Middle Stone Age assemblages come from eastern and southern Africa but date much earlier. Here we report the ages, determined by thermoluminescence dating, of fire-heated flint artefacts obtained from new excavations at the Middle Stone Age site of Jebel Irhoud, Morocco, which are directly associated with newly discovered remains of H. sapiens. A weighted average age places these Middle Stone Age artefacts and fossils at 315 ± 34 thousand years ago. Support is obtained through the recalculated uranium series with electron spin resonance date of 286 ± 32 thousand years ago for a tooth from the Irhoud 3 hominin mandible. These ages are also consistent with the faunal and microfaunal assemblages and almost double the previous age estimates for the lower part of the deposits. The north African site of Jebel Irhoud contains one of the earliest directly dated Middle Stone Age assemblages, and its associated human remains are the oldest reported for H. sapiens. The emergence of our species and of the Middle Stone Age appear to be close in time, and these data suggest a larger scale, potentially pan-African, origin for both.

The faunal sample from the Middle Stone Age (MSA) and overlying Later Stone Age (LSA) deposits of Diepkloof Rock Shelter (Western Cape Province, South Africa) includes at least 40 taxa, mostly mammals, but also tortoises, snakes, birds (especially ostrich represented by eggshell), and intertidal mollusks. The LSA sample contains only species that occurred nearby historically, including domestic sheep, which LSA people introduced to the region by 1800 years ago. In contrast, like other Western Cape MSA faunas, the Diepkloof MSA sample has more species and it is especially notable for five large extralimital grazing species. These imply a greater-than-historic role for grasses in the local vegetation, particularly in the post-Howiesons Poort (latest MSA) interval where the grazers appear most abundant. Extreme fragmentation and dark-staining impedes analysis of the MSA bones, but cut-marks, abundant burning, and numerous associated artifacts suggest that people were the main accumulators. Rare coprolites imply that carnivores could have contributed some bones, and concentrations of small mammal bones, particularly near the bottom of the MSA sequence, suggest a role for raptors. Tortoise bones are common throughout the sequence, and the MSA specimens tend to be especially large, as in other MSA assemblages. The LSA specimens are smaller, probably because LSA human populations were denser and preyed on tortoises more intensively. The most surprising aspect of the Diepkloof assemblage is its marine component. The coast is currently 14 km away and it would have been even more distant during much of the MSA when sea levels were often lower. Intertidal mollusks, particularly black mussels and granite limpets, are concentrated in the LSA and in the Late and Post-Howiesons Poort layers. Only LSA shells are complete enough for measurement, and the limpets are small as at other LSA sites. The implication is again for more intense LSA collection by relatively dense human populations. Both the LSA and MSA deposits also contain bones of shorebirds and Cape fur seals. Whale barnacles and occasional dolphin bones indicate that MSA people scavenged beached cetaceans.

Approximately 50 ka, one or more subgroups of modern humans expanded from Africa to populate the rest of the world. Significant behavioral change accompanied this expansion, and archaeologists commonly seek its roots in the African Middle Stone Age (MSA; ∼200 to ∼50 ka). Easily recognizable art objects and “jewelry” become common only in sites that postdate the MSA in Africa and Eurasia, but some MSA sites contain possible precursors, especially including abstractly incised fragments of ocher and perforated shells interpreted as beads. These proposed art objects have convinced most specialists that MSA people were behaviorally (cognitively) modern, and many argue that population growth explains the appearance of art in the MSA and its post-MSA florescence. The average size of rocky intertidal gastropod species in MSA and later coastal middens allows a test of this idea, because smaller size implies more intense collection, and more intense collection is most readily attributed to growth in the number of human collectors. Here we demonstrate that economically important Cape turban shells and limpets from MSA layers along the south and west coasts of South Africa are consistently and significantly larger than turban shells and limpets in succeeding Later Stone Age (LSA) layers that formed under equivalent environmental conditions. We conclude that whatever cognitive capacity precocious MSA artifacts imply, it was not associated with human population growth. MSA populations remained consistently small by LSA standards, and a substantial increase in population size is obvious only near the MSA/LSA transition, when it is dramatically reflected in the Out-of-Africa expansion.

Archaeological sites in northern Africa provide a rich record of increasing importance for the origins of modern human behaviour and for understanding human dispersal out of Africa. However, the timing and nature of Palaeolithic human behaviour and dispersal across north-western Africa (the Maghreb), and their relationship to local environmental conditions, remain poorly understood. The cave of Rhafas (northeast Morocco) provides valuable chronological information about cultural changes in the Maghreb during the Palaeolithic due to its long stratified archaeological sequence comprising Middle Stone Age (MSA), Later Stone Age (LSA) and Neolithic occupation layers. In this study, we apply optically stimulated luminescence (OSL) dating on sand-sized quartz grains to the cave deposits of Rhafas, as well as to a recently excavated section on the terrace in front of the cave entrance. We hereby provide a revised chronostratigraphy for the archaeological sequence at the site. We combine these results with geological and sedimentological multi-proxy investigations to gain insights into site formation processes and the palaeoenvironmental record of the region. The older sedimentological units at Rhafas were deposited between 135 ka and 57 ka (MIS 6 –MIS 3) and are associated with the MSA technocomplex. Tanged pieces start to occur in the archaeological layers around 109 ka, which is consistent with previously published chronological data from the Maghreb. A well indurated duricrust indicates favourable climatic conditions for the pedogenic cementation by carbonates of sediment layers at the site after 57 ka. Overlying deposits attributed to the LSA technocomplex yield ages of ~21 ka and ~15 ka, corresponding to the last glacial period, and fall well within the previously established occupation phase in the Maghreb. The last occupation phase at Rhafas took place during the Neolithic and is dated to ~7.8 ka.

National audience

Teeth are the basis for the best methods for estimating the age-at-death of archaeological and paleontological faunal remains, because they change by eruption and wear throughout an individual’s life and because they preserve well. However, age-at-death can be difficult to estimate when teeth are isolated or when no known-age reference sample is available. For these reasons, researchers developed the Quadratic Crown Height Method (QCHM), a set of quadratic formulae that can be used to predict age-atdeath from tooth crown height, when unworn crown height and the ages when the tooth erupts and when its crown height should reach zero can be estimated. Previous tests of the QCHM suggest that modified equations could improve the method. Here, we use crown height measurements on a sample of 226 known-age Rocky Mountain elk (Cervus elaphus) to perform such modifications. We adjust the age at which each tooth type’s crown height reaches zero from the species’ potential ecological longevity or average maximum life span to an age that we empirically estimate for each tooth type. We also empirically assess whether for different elk teeth the exponent in the QCHM formula is actually equal to 2; it is for M1, but for P4 it is about 1, indicating a roughly linear relationship. The exponents for M2 and M3 are intermediate, being closer to 1.5. Because different teeth wear at different rates and wear completely away at different ages, we recommend that researchers use the modified equations provided here to estimate age-at-death for samples of Cervus elaphus.

The construction of mortality profiles to investigate age-at-death patterns is a typical component of most faunal analyses. While many methods exist for constructing and comparing mortality profiles, plotting the percentages of juvenile, prime, and old individuals on a triangular graph, or ternary diagram, remains a popular method for making comparisons. Typically, these comparisons are made visually, but because sample sizes are often small, any differences may be the product of sampling rather than meaningful contrasts in depositional history. To overcome this problem, here we present a likelihood-based method for making statistical comparisons of mortality profiles on a triangular graph, and we make available a cross-platform computer program that implements the method. Although we developed the method with mortality profiles in mind, in principle, it can be used to analyze any kind of artifact for which there are three distinct categories.

In order to understand the behaviours and subsistence choices of Palaeolithic hunter-gatherers, it is essential to understand the behavioural ecology of their prey. Here, we present strontium isotope data from sequentially-sampled enamel from three reindeer (Rangifer tarandus ssp.) and a single bison (Bison cf. priscus) from the late Middle Palaeolithic site of Jonzac (Chez-Pinaud), France. The results are used to investigate the ranging and migratory behaviours of these important prey species. We found that the bison had isotope values most consistent with a local range, while the three reindeer had values indicating a seasonal migration pattern. Due to the similarity of the patterning of two of the three reindeer and in conjunction with zooarchaeological results, we suggest that they may have been from the same herd, were likely killed around the same point during their seasonal round and may therefore be the product of a single hunting event or a small number of successive hunting events. The isotope analyses complement the zooarchaeological data and have allowed greater insight into the palaeoecology of these species, the palaeoenvironment, and Neanderthal site use and hunting strategies.

Klein (1979, 1994) reported a contrast in ungulate species frequencies between the Middle Stone Age (MSA) layers of Klasies River Main (KRM) and the Later Stone Age (LSA) layers of nearby Nelson Bay Cave (NBC), South Africa. Early European travelers noted that the distinctive subtropical broadleaf forest surrounding both coastal sites housed exceptional numbers of buffalo (Syncerus caffer) and bushpig (Potamochoerus larvatus; Skead, 1980). They failed to mention eland, suggesting that eland were locally absent or rare, in keeping with this species’ tendency to avoid forest (Skinner and Chimimba, 2005). At both KRM and NBC the deposits formed under similar climatic conditions (Last Interglacial and the Holocene Interglacial, respectively), yet only the NBC LSA fauna anticipated the historic forest fauna in the abundance of buffalo and bushpig and the rarity of eland. The unanticipated abundance of eland in the KRM MSA fauna led Klein to suggest that the KRM people were less competent hunters. Unlike buffalo and bushpig, which commonly counterattack predators, eland tend to flee, and associated artifacts suggest that only the NBC LSA hunters had projectile weapons that would have allowed them to hunt dangerous species from a relatively safe distance. The NBC LSA people could then have obtained buffalo and bushpig more in proportion to their live abundance. Lacking projectiles, the KRM MSA people would have tended to capture more of the less threatening eland, which they could have driven over nearby cliffs. Driving could explain why the KRM eland were mainly prime-age adults, whereas the KRM buffalo were mostly very young or old individuals, as would be expected if the hunters focused on the most vulnerable buffalo or acquired them by scavenging. Conceivably, culture was irrelevant, and topography or some other strictly local factor explains the abundance of eland at KRM. However, a test for this would require additional fossiliferous MSA sites in the same forest, and KRM remains the only one. The KRM/ NBC contrast is roughly replicated between Die Kelders Cave 1 (DK1) and nearby Byneskranskop 1 (BNK1), where eland dominate buffalo and bushpig in the DK1MSA layers and buffalo and bushpig dominate eland in the BNK1 LSA layers (Klein and Cruz-Uribe, 1996). However, DK1 and BNK1 are located 400e500 km east of KRM and NBC (Fig. 1) in a scrub/thicket mosaic where buffalo and bushpig may not have outnumbered eland historically. In addition, the DK1 and BNK1 faunas accumulated under differing climatic conditions (mainly glacial and interglacial, respectively), and a paleoenvironmental difference could thus explain the faunal contrast. The DK1/BNK1 contrast underscores the need to control for environment before invoking cultural or human behavioral explanations for differences between sites. With Klein’s interpretation of the KRM/NBC contrast in mind, Faith (2008) undertook log-transformed regressions of eland and buffaloþ pig abundance on the total number of ungulates in 95 LSA samples and 27 MSA samples from Lesotho and South Africa. He concluded that “eland, buffalo, andwild pig are equally abundant in the MSA and LSA” (p. 24) and therefore that MSA and LSA people were equally adept hunters. We show below that Faith’s results are based on inappropriate use of regression, but even if we were to accept them, they do not show that MSA and LSA people were behaving in the same way. Eland, buffalo, and wild pigs could be equally abundant, on average, in MSA and LSA sites only if their live proportions (relative numbers) were usually the same nearby, or if MSA and LSA people, faced with different proportions at different times and places, managed to equalize the proportions in their sites. Faith’s result could mean, for example, that buffalo and pigs were generally rarer near LSA sites but that more effective LSA hunting overcame the environmental difference. Our point here is not to argue that MSA people were inferior hunters. That issue remains debatable. Rather, we are concerned with how to appropriately use the zooarchaeological record to * Corresponding author. E-mail address: rklein@stanford.edu (R.G. Klein).

Principal Components Analysis (PCA) of measurements on twelve metacarpal and ten metatarsal dimensions indicates that metapodials differ much more in size than in shape among the twenty-three genera of extant Sub-Saharan bovids. The only conspicuous shape variation is in metapodial length, which sometimes differs significantly among genera that are similar in all other dimensions. It follows that Discriminant Analysis (DA) intended to identify metapodials to genus will depend primarily on size, and except for the occasional like-sized genera that differ in metapodial length, DA will often confuse genera of similar size. DA of the same metapodials subjected to PCA supports this prediction. PCA further shows that bovid species that favor the same habitat often differ in metapodial size, shape, or both, and species that favor different habitats often overlap substantially in their metapodial morphometrics. DA intended to assign metapodials to habitat will thus produce different results depending on the species used to characterize each habitat type and on the number of specimens by which each species is represented. Habitat reconstruction founded in DA is therefore taxon dependent rather than taxon-free, and it cannot supplant reconstruction based on taxonomic identifications below the family level.

The cultural history and past ecology of the northern Rukwa Valley, southwestern Tanzania, are largely unknown. We report here the results of excavations conducted at Kibaoni, a Pimbwe village located near Katavi National Park, which, based on ceramic typologies, displays a long sequence of occupation dating to as early as the fourth century AD. The faunal remains at the site indicate opportunistic hunting practices similar to those seen in Rukwa today, despite the numerous political and demographic changes of the past century. Our analysis contributes to the construction of a cultural history for the region and indicates that further study will aid in understanding the interplay between humans and the environment in late precolonial, interior East Africa.

A new interdisciplinary project was initiated to excavate a portion of the Palaeolithic site of La Ferrassie left intact by earlier excavations. One of the aims of this project was to provide chronological information on the succession of Middle and Upper Palaeolithic layers, as well as on the skeletons unearthed by Capitan and Peyrony in the early 1900's. We report here preliminary results on the lithics, faunal remains, site formation processes, and on the stratigraphic context of the La Ferrassie 1 and 2 skeletons that were found adjacent to our excavations. Finally, results from luminescence dating of the sediments and a preliminary set of radiocarbon ages are presented. Quartz OSL, both at the multi-grain and single-grain levels of analysis, and post-IR IRSL of feldspar at various stimulation temperatures are compared. The quartz/feldspar comparison revealed a bleaching problem for the quartz OSL (and the feldspar pIRIR signals) from Layer 2; as a consequence, the age of this Layer was determined using a minimum age model. A Mousterian industry with bifaces, at the base of the sequence, has been dated between 91 ± 9 and 44 ± 3 ka. The Ferrassie Mousterian layers are attributed to MIS 3, between 54 ± 3 and 40 ± 2 ka, and thus appear very late in the final Middle Palaeolithic of the region; furthermore, these ages constrain the chronology of the La Ferrassie 1 and 2 skeletons, which have been attributed to one of these Ferrassie Mousterian layers. The Châtelperronian layer is dated to 42 ± 3 ka and the Aurignacian to 37 ± 2 ka. Implications of the ages for the La Ferrassie 1 and 2 skeletons, and for the variability of late Mousterian, are discussed.

In this study, I examine three methods that are currently used for comparing mortality profiles from zooarchaeological and palaeontological samples: (1) histograms with 10% of life-span age classes; (2) boxplots showing tooth crown height medians; and (3) triangular plots of the proportions of young, prime and old animals. I assess the advantages and disadvantages of each method using data collected on two samples of Northern Yellowstone elk (Cervus elaphus nelsoni) with known, or cementum annuli-determined, ages at death. One sample was hunted by wolves (n = 96), and the other was hunted by recent humans using rifles (n = 226). I tested each method with the known or cementum annuli age distributions and with age estimation techniques appropriate for archaeological assemblages. Histograms are best used when the relationship between dental eruption/attrition and age is well established so that individuals can be confidently assigned into 10% of life-span groups, and when more than 30 or 40 individuals are present in the assemblage. Boxplots employ raw crown heights, thus removing the error introduced by assigning specimens to age classes, and therefore they allow the analysis of species where the relationship between dental eruption/attrition and age is unknown. Confidence intervals around the medians allow samples to be statistically compared. Triangular plots are easy to use and allow multiple samples and species to be considered simultaneously, but samples cannot be statistically compared. Modified triangular plots bootstrap samples to provide 95% confidence ellipses, allowing for statistical comparisons between samples. When possible, samples should be examined using multiple methods to increase confidence in the results. Copyright © 2005 John Wiley & Sons, Ltd.

Prey age-at-death is often used to assess hunting strategies, but the age profiles of animals represented in Middle Palaeolithic ungulate assemblages vary through time and across space. Understanding this diversity is important for reconstructing human behavioural evolution. To explore this issue, I studied one species, red deer (Cervus elaphus), in late Acheulean and Middle Palaeolithic assemblages from France (Combe-Grenal, Lazaret, and Piegu) and northern Spain (Gabasa 1 and Labeko Koba). I reconstructed age distributions using three methods: histograms, boxplots of median crown heights, and modified triangular plots, and two tooth combinations: dp4/p4 and dp4/m1. Two modern samples served as comparative baselines: a sample of C. elaphus killed by modern hunters in western Montana, USA and a sample of C. elaphus hunted by wolves reintroduced into Yellowstone National Park, Wyoming, USA. The mortality distributions of the modern samples differ dramatically. The wolf-hunted sample contains more juvenile and old individuals, while the human-hunted sample has more young adults. The fossil age structures fall into two clusters: 1) Gabasa 1 and Labeko Koba with a high proportion of juvenile red deer and few prime-aged adults, and 2) Combe-Grenal, Lazaret, and Piegu with equal abundances of juvenile and prime-aged red deer. I consider possible explanations for this pattern. More sites and more detailed analyses are needed to determine conclusively that the difference is related ancient human behaviour and not other pre- and post-depositional processes. However, the assemblage from Gabasa Cave is important for demonstrating that deciduous teeth can be well preserved in the Late Pleistocene fossil record. Copyright © 2004 John Wiley & Sons, Ltd.

Human fossils and the genetics of extant human populations indicate that living people derive primarily from an African population that lived within the last 200,000 years. Yet it was only ≈50,000 years ago that the descendants of this population spread to Eurasia, where they swamped or replaced the Neanderthals and other nonmodern Eurasians. Based on archaeological observations, the most plausible hypothesis for the delay is that Africans and Eurasians were behaviorally similar until 50,000 years ago, and it was only at this time that Africans developed a behavioral advantage. The archaeological findings come primarily from South Africa, where they suggest that the advantage involved much more effective use of coastal resources. Until now, the evidence has come mostly from deeply stratified caves on the south (Indian Ocean) coast. Here, we summarize results from recent excavations at Ysterfontein 1, a deeply stratified shelter in a contrasting environment on the west (Atlantic) coast. The Ysterfontein 1 samples of human food debris must be enlarged for a full comparison to samples from other relevant sites, but they already corroborate two inferences drawn from south coast sites: ( i ) coastal foragers before 50,000 years ago did not fish routinely, probably for lack of appropriate technology, and ( ii ) they collected tortoises and shellfish less intensively than later people, probably because their populations were smaller.

Mortality profiles can be used to investigate modes of accumulation of fossil assemblages, including predation by ancient nonhuman and human (Homo) hunters. Prey age at death reflects hunting decisions and opportunities in terms of calories returned for energy spent and risks taken. Nonhuman carnivores commonly harvest vulnerable individuals (young and old), whereas humans consistently can take prime adults. As an example of how mortality profiles can be used to illuminate predator behavior, this article examines when the ability to select prime-age animals emerged during human behavioral evolution. Although it would be expected that the appearance of this behavior coincided with the appearance of morphologically and archaeologically modern humans, analyses of archaeological assemblages from western Europe and South Africa suggest that this trait was already present in archaic people who preceded modern humans. Studying age distributions in fossil assemblages is not without its limitations, including the difficulty of estimating age at death in fossil specimens, pre- and postdepositional biases, and reconstructing the age structure and behavior of prey herds. Nevertheless, valuable behavioral information can be gained by using controlled comparisons of many assemblages.

Plotting the percentages of juvenile, prime, and old individuals on a triangular graph has become a popular method for analysing mortality profiles or age structures found in archaeological faunal samples. This method allows easy comparisons of multiple samples or multiple species and appears to work even with small sample sizes. However, the utility of triangular graphs is compromised for two reasons: (1) samples cannot be statistically compared and (2) the points on the graph are based on percentages, and, therefore, they do not provide information about sample size. The modified triangular graph described here offers a method for approximating 95% confidence intervals around the data points by using bootstrapping. Samples with non-overlapping contours are likely to have had different pre- or post-depositional histories. The 95% density contours reflect sample size since they shrink as samples get larger. Thus, the modified triangular graph allows more confident comparisons of three age class data.

The Archaeology of the West Coast of South Africa. Edited by Antonieta Jerardino, Antonia Malan, and David Braun. Cambridge Monographs in African Archaeology 84, BAR International Series 2526. Arch...

Modern humans replaced Neandertals ∼40,000 y ago. Close to the time of replacement, Neandertals show behaviors similar to those of the modern humans arriving into Europe, including the use of specialized bone tools, body ornaments, and small blades. It is highly debated whether these modern behaviors developed before or as a result of contact with modern humans. Here we report the identification of a type of specialized bone tool, lissoir , previously only associated with modern humans. The microwear preserved on one of these lissoir is consistent with the use of lissoir in modern times to obtain supple, lustrous, and more impermeable hides. These tools are from a Neandertal context proceeding the replacement period and are the oldest specialized bone tools in Europe. As such, they are either a demonstration of independent invention by Neandertals or an indication that modern humans started influencing European Neandertals much earlier than previously believed. Because these finds clearly predate the oldest known age for the use of similar objects in Europe by anatomically modern humans, they could also be evidence for cultural diffusion from Neandertals to modern humans.

International audience; Neandertals were effective hunters of large ungulates throughout their geographic and temporal ranges. Equipped with this knowledge, researchers in paleoanthropology continue to seek insight on the rela- tionships between hunting and subsistence strategies with other components of the Neandertals' niche, such as mobility, site use, and lithic technology. The Quina Mousterian deposits from the rockshelter site of Chez Pinaud Jonzac (Charente-Maritime, France; hereafter Jonzac) offer an excellent opportunity to pursue these issues. This paper focuses on the extensive and well-preserved skeletal remains of reindeer (Rangifer tarandus) recovered from recent excavations of the site, representing at least 18 individuals that were hunted by Neandertals during the fall through winter. Our zooarchaeological results indicate that all ages of reindeer were hunted but adult individuals predominate. No bias is evident in the comparable frequencies of males and females. These prey were butchered on-site, with abundant evidence of meat filleting and marrow exploitation. In the excavated sample, the absence of hearths and the almost complete lack of burned bones or stones suggest that Neandertals were not using fire to assist with processing the reindeer carcasses. The zooarchaeological results presented here indicate that reindeer were hunted during a restricted window of time when they were seasonally abundant in the local area near Jonzac. Taken together with the lithic industry based on bifacial elements, the evidence is consistent with a pattern of site use by highly mobile hunter-gatherers making frequent, short-term visits. Ongoing research at Jonzac and other Quina Mousterian localities will contribute to a better understanding of Neandertal behavior during cold climate phases.

Zooarchaeological evidence has featured prominently in the debate of the mode and tempo of modern human origins during the Late Pleistocene of Africa. However, most of the assemblages included in the discussion are from southern Africa, and we have limited knowledge about human paleoecology from other parts of the continent. This chapter aims to review the zooarchaeological record for Late Pleistocene human subsistence in northwestern Africa. This region provides some of the earliest and most complete fossil evidence for anatomically modern humans, and it provides a unique aspect of human technological development in the Aterian stone tool industry. Understanding the subsistence of these early people is important for understanding human behavioral evolution. Until recently, most faunal analyses from the region have been focused on paleontology, biostratigraphy, and environmental reconstructions. Zooarchaeological studies have been limited by sample size, excavation methods, and the lack of good comparative data. What is known of the Northwest African record is placed in the context of our knowledge of Middle Stone Age subsistence from the rest of Africa, and future research directions are offered. The current projects in Morocco, as discussed in this volume, will provide valuable insights into North African subsistence during the Late Pleistocene.

While there has long been an interest in Holocene coastal adaptations globally (see other papers in this volume and Erlandson 2001), only more recently has attention focused on marine resource exploitation during the more ancient times of the Middle Paleolithic (MP) and Middle Stone Age (MSA) of Europe and Africa (∼250,000–50,000 years ago). The ancestors of all living humans originated in Africa sometime at the beginning, during, or at the end of the MSA, and therefore, investigations into the full variation of human adaptations during this time provide critical data on how changing diets and environments are linked to modern human origins. McBrearty and Brooks (2000) include “shellfishing” starting at 140,000 years ago (140 kya) and “fishing” at 110 kya in their list of behavioral innovations of the MSA (their Fig. 13:530) and as part of a general increase in diet breadth (their Table 3:492) that is indicative of fully modern human behavior. The earliest evidence of shellfishing has since been revised to at least 164 kya (Marean et al. 2007). In addition, Parkington (2003; see also Crawford et al. 1999) has proposed that the nutrients provided by shellfish were integral to building effective modern human brains during the MSA and, therefore, to modern human origins. Others see the exploitation of marine resources as typical of coastal living, and that real indicators of changing behavior, evidenced by further increased diet breadth and increased human population densities, come only after the end of the MSA (Klein et al. 2004). Because studies into coastal adaptations allow us to investigate the relationship between subsistence, technology, human population size and density, and the environment, these studies are uniquely positioned to inform on modern human origins, and therefore, it is important that future studies focus on marine resource exploitation during these ancient times.

Ongoing debates about the emergence of modern human behavior, however defined, regularly incorporate observations from the later part of the southern African Middle Stone Age and emphasize the early appearance of artifacts thought to reflect symbolic practice. Here we report a large sample of 270 fragments of intentionally marked ostrich eggshell from the Howiesons Poort at Diepkloof Rock Shelter, Western Cape, South Africa. Dating from ≈60,000 years ago, these pieces attest to an engraving tradition that is the earliest reliable evidence of what is a widespread modern practice. These abstract linear depictions were made on functional items (eggshell containers), which were curated and involved in daily hunter-gatherer life. The standardized production of repetitive patterns, including a hatched band motif, suggests a system of symbolic representation in which collective identities and individual expressions are clearly communicated, suggesting social, cultural, and cognitive underpinnings that overlap with those of modern people.

Stringer et al. (1) report 149 mollusk shells, 5 seal bones, 3 dolphin bones, and 3 fish bones from Mousterian layers in 2 caves on Gibraltar. From this they argue that the Neanderthals had “focused coastal visits… repeated during particular times, possibly seasonal” (p. 14323) and that in this respect they surpassed …

International audience; We report here on the isotopic analysis (carbon and nitrogen) of collagen extracted from a Neanderthal tooth and animal bone from the late Mousterian site of Jonzac (Charente-Maritime, France). This study was undertaken to test whether the isotopic evidence indicates that animal protein was the main source of dietary protein for this relatively late Neanderthal, as suggested by previous studies. This was of particular interest here because this is the first isotopic study of a relatively late Neanderthal associated with Mousterian of Acheulian Tradition (MTA, dating to approximately 55,000 to 40,000 BP) technology. We found that the Jonzac Neanderthal had isotopic values consistent with a diet in which the main protein sources were large herbivores, particularly bovids and horses. We also found evidence of different dietary niches between the Neanderthal and a hyena at the site, with the hyena consuming mainly reindeer.

The Elandsfontein site, Western Cape Province, South Africa, is well known for an archaic hominin skullcap associated with later Acheulean artifacts. The site has also provided nearly 13,000 mammalian bones that can be identified to skeletal part and taxon. The assemblage derives from 49 species, 15 of which have no historic descendants. Comparisons to radiometrically dated faunas in eastern Africa indicate an age between 1 million and 600 thousand years ago. Unique features of the fauna, including the late occurrence of a dirk-toothed cat and a sivathere, may reflect its geographic origin in a region that was notable historically for its distinctive climate and high degree of biotic endemism. Together, taxonomic composition, geomorphic setting, and pollen extracted from coprolites indicate the proximity of a large marsh or pond, maintained by a higher water table. The small average size of the black-backed jackals implies relatively mild temperatures. The sum of the evidence places bone accumulation during one of the mid-Pleistocene interglacials that were longer and cooler than later ones, including the Holocene. The geomorphic context of the fauna presents no evidence for catastrophe, and most deaths probably resulted from attritional factors that disproportionately killed the young and old. However, only the dental-age profile of long-horned buffalo supports this directly. Field collection methods biased skeletal-part representation, but originally, it probably resembled the pattern in the younger, marsh-edge Acheulean occurrence at Duinefontein 2, 45 km to the south. Excavation there exposed multiple vertebral spreads, which probably mark carcasses from which hominins or large carnivores removed the meatier elements. Bone damage at both sites suggests that, despite abundant artifacts, hominins were much less important than carnivores in the bone accumulation. Together with limited observations from other sites, Elandsfontein and Duinefontein provisionally suggest that Acheulean-age hominins obtained few large mammals, whether by hunting or scavenging. 2006 Elsevier Ltd. All rights reserved.