Environmental influences on human innovation and behavioural diversity in southern Africa 92-80 thousand years ago.

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.

A worked bone assemblage from 120,000-90,000 year old deposits at Contrebandiers Cave, Atlantic Coast, Morocco.

The emergence of Homo sapiens in Pleistocene Africa is associated with a profound reconfiguration of technology. Symbolic expression and personal ornamentation, new tool forms, and regional technological traditions are widely recognized as the earliest indicators of complex culture and cognition in humans. Here we describe a bone tool tradition from Contrebandiers Cave on the Atlantic coast of Morocco, dated between 120,000-90,000 years ago. The bone tools were produced for different activities, including likely leather and fur working, and were found in association with carnivore remains that were possibly skinned for fur. A cetacean tooth tip bears what is likely a combination of anthropogenic and non-anthropogenic modification and shows the use of a marine mammal tooth by early humans. The evidence from Contrebandiers Cave demonstrates that the pan-African emergence of complex culture included the use of multiple and diverse materials for specialized tool manufacture.

Reconstructing Late Pleistocene paleoclimate at the scale of human behavior: an example from the Neandertal occupation of La Ferrassie (France).

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.

Non-destructive ZooMS identification reveals strategic bone tool raw material selection by Neandertals.

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.

Subsistence strategies throughout the African Middle Pleistocene: Faunal evidence for behavioral change and continuity across the Earlier to Middle Stone Age transition.

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.

Time wears on: Assessing how bone wears using 3D surface texture analysis.

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.

Clarifying distinct models of modern human origins in Africa.

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.

The age of the hominin fossils from Jebel Irhoud, Morocco, and the origins of the Middle Stone Age.

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.

A New Chronology for Rhafas, Northeast Morocco, Spanning the North African Middle Stone Age through to the Neolithic.

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.

Neandertals made the first specialized bone tools in Europe.

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.

Archaeological shellfish size and later human evolution in Africa.

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.

On the industrial attributions of the Aterian and Mousterian of the Maghreb.

North Africa is quickly emerging as one of the more important regions yielding information on the origins of modern Homo sapiens. Associated with significant fossil hominin remains are two stone tool industries, the Aterian and Mousterian, which have been differentiated, respectively, primarily on the basis of the presence and absence of tanged, or stemmed, stone tools. Largely because of historical reasons, these two industries have been attributed to the western Eurasian Middle Paleolithic rather than the African Middle Stone Age. In this paper, drawing on our recent excavation of Contrebandiers Cave and other published data, we show that, aside from the presence or absence of tanged pieces, there are no other distinctions between these two industries in terms of either lithic attributes or chronology. Together, these results demonstrate that these two 'industries' are instead variants of the same entity. Moreover, several additional characteristics of these assemblages, such as distinctive stone implements and the manufacture and use of bone tools and possible shell ornaments, suggest a closer affinity to other Late Pleistocene African Middle Stone Age industries rather than to the Middle Paleolithic of western Eurasia.

Neandertal mobility and large-game hunting: the exploitation of reindeer during the Quina Mousterian at Chez-Pinaud Jonzac (Charente-Maritime, France).

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 relationships 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.

Strontium isotope evidence for migration in late Pleistocene Rangifer: implications for Neanderthal hunting strategies at the Middle Palaeolithic site of Jonzac, France.

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.

In silico QTL mapping of basal liver iron levels in inbred mouse strains.

Both iron deficiency and iron excess are detrimental in many organisms, and previous studies in both mice and humans suggest that genetic variation may influence iron status in mammals. However, these genetic factors are not well defined. To address this issue, we measured basal liver iron levels in 18 inbred strains of mice of both sexes on a defined iron diet and found ∼4-fold variation in liver iron in males (lowest 153 µg/g, highest 661 µg/g) and ∼3-fold variation in females (lowest 222 µg/g, highest 658 µg/g). We carried out a genome-wide association mapping to identify haplotypes underlying differences in liver iron and three other related traits (copper and zinc liver levels, and plasma diferric transferrin levels) in a subset of 14 inbred strains for which genotype information was available. We identified two putative quantitative trait loci (QTL) that contain genes with a known role in iron metabolism: Eif2ak1 and Igf2r. We also identified four putative QTL that reside in previously identified iron-related QTL and 22 novel putative QTL. The most promising putative QTL include a 0.22 Mb region on Chromosome 7 and a 0.32 Mb region on Chromosome 11 that both contain only one candidate gene, Adam12 and Gria1, respectively. Identified putative QTL are good candidates for further refinement and subsequent functional studies.

From the Cover: A Howiesons Poort tradition of engraving ostrich eggshell containers dated to 60,000 years ago at Diepkloof Rock Shelter, South Africa.

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 approximately 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.

The mammalian fauna associated with an archaic hominin skullcap and later Acheulean artifacts at Elandsfontein, Western Cape Province, South Africa.

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.