Middle Stone Age Ochre Processing and Behavioural Complexity in the Horn of Africa: Evidence from Porc-Epic Cave,Dire Dawa,Ethiopia

Ochre is a common feature at Middle Stone Age (MSA) sites and has often been interpreted as a proxy for the origin of modern behaviour. However, few ochre processing tools, ochre containers, and ochre-stained artefacts from MSA contexts have been studied in detail within a theoretical framework aimed at inferring the technical steps involved in the acquisition, production and use of these artefacts. Here we analyse 21 ochre processing tools, i.e. upper and lower grindstones, and two ochre-stained artefacts from the MSA layers of Porc-Epic Cave, Dire Dawa, Ethiopia, dated to ca. 40 cal kyr BP. These tools, and a large proportion of the 4213 ochre fragments found at the site, were concentrated in an area devoted to ochre processing. Lower grindstones are made of a variety of raw materials, some of which are not locally available. Traces of use indicate that different techniques were employed to process ochre. Optical microscopy, XRD, μ-Raman spectroscopy, and SEM-EDS analyses of residues preserved on worn areas of artefacts show that different types of ferruginous rocks were processed in order to produce ochre powder of different coarseness and shades. A round stone bearing no traces of having been used to process ochre is half covered with residues as if it had been dipped in a liquid ochered medium to paint the object or to use it as a stamp to apply pigment to a soft material. We argue that the ochre reduction sequences identified at Porc-Epic Cave reflect a high degree of behavioural complexity, and represent ochre use, which was probably devoted to a variety of functions.     

Nassarius kraussianus shell beads from Blombos Cave: evidence for symbolic behaviour in the Middle Stone Age

Since 1991, excavations at Blombos Cave have yielded a well-preserved sample of faunal and cultural material in Middle Stone Age (MSA) levels. The uppermost MSA phase, M1, is dated to c. 75 ka by optically stimulated luminescence (OSL) and thermoluminescence, and the middle M2 phase to a provisional c. 78 ka. Artefacts unusual in a MSA context from these phases include bifacial points, bone tools, engraved ochre and engraved bone. In this paper, we describe forty-one marine tick shell beads recovered from these MSA phases and tick shell beads from Later Stone Age (LSA) levels at Blombos Cave and the Die Kelders site. Thirty-nine shell beads come from the upper M1 phase and two from M2. Morphometric, taphonomic and microscopic analysis of modern assemblages of living and dead tick shell demonstrate that the presence of perforated Nassarius kraussianus shells in the Blombos MSA levels cannot be due to natural processes or accidental transport by humans. The types of perforation seen on the MSA shells are absent on modern accumulations of dead shells and not attributable to post-depositional damage. Their location, size, and microscopic features are similar to those obtained experimentally by piercing the shell wall, through the aperture, with a sharp bone point. Use-wear, recorded on the perforation edge, the outer lip, and the parietal wall of the aperture indicates the shells having being strung and worn. MSA shell beads differ significantly in size, perforation type, wear pattern and shade compared to LSA beads and this eliminates the possibility of mixing across respective levels. Thirty-one beads were found in four groups of five to twelve beads, each group being recovered in a single square or in two adjacent sub-squares during a single excavation day. Within a group, shells display a similar shade, use-wear pattern and perforation size suggesting their provenance from the same beadwork item, lost or disposed during a single event. The likely symbolic significance of these finds suggests levels of cognitively modern behaviour not previously associated with MSA people.     

Engraved ochres from the Middle Stone Age levels at Blombos Cave, South Africa

Powerful categories of evidence for symbolically mediated behaviour, variously described as ‘modern’ or ‘cognitively modern’ human behaviour, are geometric or iconographic representations. After 40,000 years ago such evidence is well documented in much of the Old World and is widely considered as typifying ‘modern human culture,’ but earlier evidence is rare. In Africa, this includes two deliberately engraved ochre pieces from c. 75,000 year old levels at Blombos Cave, Western Cape, South Africa and the greater than 55,000 year old incised ostrich egg shell from the Diepkloof shelter, located in the same province. Here we report on thirteen additional pieces of incised ochre recovered from c. 75,000-100,000 year old levels at Blombos Cave. These finds, taken together with other engraved objects reported from other southern African sites, suggest that symbolic intent and tradition were present in this region at an earlier date than previously thought.     

Putting ochre to the test: replication studies of adhesives that may have been used for hafting tools in the Middle Stone Age

Substantial frequencies of Middle Stone Age (MSA) lithics from Rose Cottage and Sibudu Caves in South Africa have red ochre on their proximal and medial portions. Residue studies suggest that the tools were hafted and that the ochre may be part of the adhesive used for hafting the tools. Replication studies show that ochre is indeed a useful loading agent for adhesive; however, there are other potential loading agents. It is also possible to use unloaded plant resin, but this agent is brittle and difficult to work with. It appears that people living in the MSA had wide knowledge of ingredients suitable for hafting tools, and that they chose different adhesive recipes because of the required properties of the adhesive. Brittle, unloaded adhesive allows a projectile head to disengage its haft and implant itself in an animal; robust adhesive keeps a spearhead safely in its shaft.     

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.     

Human genome diversity

Human genome diversity studies analyse genetic variation among individuals and between populations in order to understand the origins and evolution of anatomically modern humans (Homo sapiens sapiens). The availability of thousands of DNA polymorphisms (genetic markers) brings analytic power to these studies. Human genome diversity studies have clearly shown that the large part of genetic variability is due to differences among individuals within populations rather than to differences between populations, effectively discrediting a genetic basis of the concept of ‘race’. Evidence from paleontology, archaeology and genetic diversity studies is quite consistent with an African origin of modern humans more than 100 000 years ago. The evidence favors migrations out of African as the source of the original peopling of Asia, Australia, Europe and Oceania. An international program for the scientific analysis of human genome diversity and of human evolution has been developed. The Human Genome Diversity Project (HGDP) aims to collect and preserve biologic samples from hundreds of populations throughout the world, make DNA from these samples available to scientists and distribute to the scientific community the results of DNA typing with hundreds of genetic markers.     

Middle and late Pleistocene faunas of Pinnacle Point and their paleoecological implications

The Western Cape region of South Africa is home to a unique type of mediterranean vegetation called fynbos, as well as some of the earliest sites of modern human occupation in southern Africa. Reconstructing the paleohabitats during occupations of these early anatomically modern Homo sapiens is important for understanding the availability of resources to the humans during the development of behaviors that are often considered advanced. These reconstructions are critical to understanding the nature of the changes in the environment and resources over time. Here we analyze the craniodental fossils of the larger mammals recovered from two Pleistocene assemblages in the Pinnacle Point complex, Mossel Bay, Western Cape Region, South Africa. We reconstruct the paleohabitats as revealed by multivariate analyses of the mammalian community structures. Pinnacle Point 30 is a carnivore assemblage and Pinnacle Point 13B includes early evidence of a suite of modern human behavior; together they present an opportunity to identify environmental change over time at a localized geographic scale. Further, this is the first such study to include dated Western Cape localities from Marine Isotope Stage 6, a time of environmental pressure that may have marginalized human populations. Results indicate that environmental change in the Western Cape was more complex than generalized C(4) grassland expansions replacing fynbos habitats during glacial lowered sea levels, and thus, resources available to early modern humans in the region may not have been entirely predictable.     

Shells and ochre in Middle Paleolithic Qafzeh Cave,Israel: indications for modern behavior

Qafzeh Cave, the burial grounds of several anatomically modern humans, producers of Mousterian industry, yielded archaeological evidence reflecting their modern behavior. Dated to 92 ka BP, the lower layers at the site contained a series of hearths, several human graves, flint artifacts, animal bones, a collection of sea shells, lumps of red ochre, and an incised cortical flake. The marine shells were recovered from layers earlier than most of the graves except for one burial. The shells were collected and brought from the Mediterranean Sea shore some 35 km away, and are complete Glycymeris bivalves, naturally perforated. Several valves bear traces of having been strung, and a few had ochre stains on them.     

Brain Size Growth and Life History in Human Evolution

Increases in endocranial volume (a measure of brain size) play a major role in human evolution. Despite the importance of brain size increase, the developmental bases of human brain size evolution remain poorly characterized. Comparative analyses of endocranial volume size growth illustrate that distinctions between humans and other primates are consequences of differences in rates of brain size growth, with little evidence for differences in growth duration. Evaluation of available juvenile fossils shows that earliest hominins do not differ perceptibly from chimpanzees (Pan). However, rapid and human-like early brain growth apparently characterized Homo erectus at about 1?Ma before present. Neandertals show patterns of brain growth consistent with modern humans during infancy, but reach larger sizes than modern humans as a result of differences in later growth. Growth analyses reveal commonalities in patterns of early brain size growth during the last million years human evolution, despite major increases in adult size. This result implies consistency across hominins in terms of maternal metabolic costs of infancy. Continued size growth past infancy in Neandertals and modern humans, when compared to earlier hominins, may have cognitive implications. Differences between Neandertals and modern humans are implied, but difficult to define with certainty.     

Trans-Generational Influence of Human Disturbances in Japanese Quail: Egg Quality Influences Male Social and Sexual Behaviour

In many bird species prenatal exposure to yolk androgens of maternal origin has been found to influence offspring behavioural phenotype. In contrast to altricial birds, far less is known about maternal effects in precocial birds. In a previous experiment we found that female quail ( Coturnix japonica ) that were not previously habituated to humans (NH) produced eggs with less androgens (testosterone, androstenedione) and more progesterone when exposed to human disturbances than females habituated to humans (H). Here, we analysed social motivation and sexual behaviour of the male offspring of NH and H females. Classical behavioural test procedures were applied including separation, runway, partner choice and female encounter tests. As chicks, offspring of the NH females spent more time far from conspecifics than offspring of the H females. As adults, the same NH males showed less crowing and courtship behaviour (ritual preening) in female encounter tests than H males. Thus, maternal environment and egg quality may be key factors in the emergence of individual variability of appetitive behaviour, such as social proximity and courtship behaviour. Human disturbance of the mother seems to have triggered trans-generational effects resulting in consistently reduced social and sexual motivation in offspring until adulthood.     

Human urinary and blood toxicokinetics of beryllium after accidental exposure

PurposeBeryllium is known to have adverse health effects and is classified as carcinogenic to humans. However, data on systemic beryllium exposure in humans are rare and especially human toxicokinetics are largely uncharted. As such, the first reported multi-annual course of blood and urine concentrations after a high exposure scenario provides important new insights.MethodsFor a medical follow-up biomonitoring samples were collected for 56 months from a male subject after an accidental and multi-faceted high exposure. Sampling started on day 2 post-exposure for urine and day 147 for blood. The samples were analyzed by inductively coupled mass spectrometry (ICP-MS) and plotted longitudinally as a function of time. Terminal half-lives were calculated assuming a first-order elimination process.Main findingsBoth matrices showed highly increased initial concentrations (about 100-fold), despite the 147-day delay in blood sampling, and a marked decline over time. In urine, a two-phase excretion process was suspected based on the longitudinal data. Calculations gave terminal half-lives of 117.5 days and 666.5 days for phases 1 and 2, respectively. Blood kinetics called for a terminal half-life of 103.5 days. Elimination kinetics in blood and urine were comparable, simultaneously gathered samples showed an excellent correlation (R² = 0.985).Principal conclusionsThe long-term follow-up after a high initial exposure to beryllium provides the first detailed insights into the elimination course of systemically available beryllium in humans. Conform kinetics of beryllium in urine and blood and the strong correlation between both parameters indicate high data validity and support the good representation of the current systemically available beryllium by urine and blood concentration in humans. The relatively long terminal half-lives in both matrices suggest a possible accumulation in humans in case of repeated exposures.     

What Biomonitoring Can and Cannot Tell Us about Causality in Human Health and Ecological Risk Assessments

Biomonitoring can provide exposure and effects information on various stressors (chemical or biological) that can be useful for human health and ecological risk assessments. It has been applied over the years where harmful changes in human health or the environment were observed and which may have warranted more detailed investigation. Sometimes biomonitoring programs may have been useful in determining the significance and/or cause of these harmful observations. These data can help to infer, but not confirm, causality as exemplified in classical studies conducted in humans and wildlife. However, in most cases we note that additional work was needed to provide the information necessary to support or refute causality. Today modern technology provides the ability to measure a wide variety of parameters in environmental media, plants, animals, and humans. Finding a chemical in an environmental medium or biological tissue may be helpful in understanding potential exposure (and perhaps to begin estimating hazard) to humans and ecological receptors, but mere presence does not necessarily help to establish effects or assign causality. In this article we evaluate the strengths and weaknesses, in a risk assessment context, of the use of biomonitoring data to support a determination of causality.     

Leadership in Moving Human Groups

How is movement of individuals coordinated as a group? This is a fundamental question of social behaviour, encompassing phenomena such as bird flocking, fish schooling, and the innumerable activities in human groups that require people to synchronise their actions. We have developed an experimental paradigm, the HoneyComb computer-based multi-client game, to empirically investigate human movement coordination and leadership. Using economic games as a model, we set monetary incentives to motivate players on a virtual playfield to reach goals via players'' movements. We asked whether (I) humans coordinate their movements when information is limited to an individual group member''s observation of adjacent group member motion, (II) whether an informed group minority can lead an uninformed group majority to the minority''s goal, and if so, (III) how this minority exerts its influence. We showed that in a human group – on the basis of movement alone – a minority can successfully lead a majority. Minorities lead successfully when (a) their members choose similar initial steps towards their goal field and (b) they are among the first in the whole group to make a move. Using our approach, we empirically demonstrate that the rules of swarming behaviour apply to humans. Even complex human behaviour, such as leadership and directed group movement, follow simple rules that are based on visual perception of local movement.     

Introgression of Neandertal- and Denisovan-like Haplotypes Contributes to Adaptive Variation in Human Toll-like Receptors

Pathogens and the diseases they cause have been among the most important selective forces experienced by humans during their evolutionary history. Although adaptive alleles generally arise by mutation, introgression can also be a valuable source of beneficial alleles. Archaic humans, who lived in Europe and Western Asia for more than 200,000 years, were probably well adapted to this environment and its local pathogens. It is therefore conceivable that modern humans entering Europe and Western Asia who admixed with them obtained a substantial immune advantage from the introgression of archaic alleles. Here we document a cluster of three Toll-like receptors (TLR6-TLR1-TLR10) in modern humans that carries three distinct archaic haplotypes, indicating repeated introgression from archaic humans. Two of these haplotypes are most similar to the Neandertal genome, and the third haplotype is most similar to the Denisovan genome. The Toll-like receptors are key components of innate immunity and provide an important first line of immune defense against bacteria, fungi, and parasites. The unusually high allele frequencies and unexpected levels of population differentiation indicate that there has been local positive selection on multiple haplotypes at this locus. We show that the introgressed alleles have clear functional effects in modern humans; archaic-like alleles underlie differences in the expression of the TLR genes and are associated with reduced microbial resistance and increased allergic disease in large cohorts. This provides strong evidence for recurrent adaptive introgression at the TLR6-TLR1-TLR10 locus, resulting in differences in disease phenotypes in modern humans.     

Glacial cycling, large mammal community composition, and trophic adaptations in the Western Cape, South Africa

Some of the earliest evidence for modern human behavior has been recovered from the Western Cape Province, South Africa. Archaeological and paleontological sites in the Western Cape are typically described as “glacial” or “interglacial” in aspect based on the numbers of grazers found in the faunal assemblage, as glacial periods are often thought to have been characterized by spreading C₄ grasslands that replaced endemic C₃ shrubland vegetation found in the Western Cape today. Here, we test the hypothesis that glacial and interglacial time periods were associated with a predictable change in large mammal trophic adaptations by analyzing the proportions of grazing larger mammals from 118 levels of 15 Western Cape fossil assemblages sampling marine isotope stage (MIS) 6 to the present time to determine whether there is a change in composition in these communities that might reflect a shift in ecology and habitat. Our results indicate that trophic proportions did not significantly change over time in the Western Cape as a whole, and thus the hypothesis for habitat changes affecting the subsistence ecology of modern humans during the development of modern behavior is not supported. However, our results show that the southwestern subregion of the Western Cape was characterized by the presence of more grazing species through time than the western subregion. Thus, if ecological and population isolation during glacial periods were integral to catalyzing the development of modern behaviors in the Western Cape region of South Africa, then a complex model including the development of possible mosaic habitats is needed.     

Characterizing the Late Pleistocene MSA Lithic Technology of Sibudu,KwaZulu-Natal,South Africa

Studies of the African Middle Stone Age (MSA) have become central for defining the cultural adaptations that accompanied the evolution of modern humans. While much of recent research in South Africa has focused on the Still Bay and Howiesons Poort (HP), periods following these technocomplexes were often neglected. Here we examine lithic assemblages from Sibudu that post-date the HP to further the understanding of MSA cultural variability during the Late Pleistocene. Sibudu preserves an exceptionally thick, rich, and high-resolution archaeological sequence that dates to ∼58 ka, which has recently been proposed as type assemblage for the “Sibudan”. This study presents a detailed analysis of the six uppermost lithic assemblages from these deposits (BM-BSP) that we excavated from 2011–2013. We define the key elements of the lithic technology and compare our findings to other assemblages post-dating the HP. The six lithic assemblages provide a distinct and robust cultural signal, closely resembling each other in various technological, techno-functional, techno-economic, and typological characteristics. These results refute assertions that modern humans living after the HP possessed an unstructured and unsophisticated MSA lithic technology. While we observed several parallels with other contemporaneous MSA sites, particularly in the eastern part of southern Africa, the lithic assemblages at Sibudu demonstrate a distinct and so far unique combination of techno-typological traits. Our findings support the use of the Sibudan to help structuring this part of the southern African MSA and emphasize the need for further research to identify the spatial and temporal extent of this proposed cultural unit.     

Extending the chronology of deposits at Blombos Cave, South Africa, back to 140 ka using optical dating of single and multiple grains of quartz

Optically stimulated luminescence (OSL) measurements are reported for both single aliquots (of two different sizes) and single grains of quartz from deposits within Blombos Cave. Ages have been obtained for six sediments from the Middle Stone Age (MSA) occupation levels and for two sterile sands, one underlying the archaeological sediment and one overlying the Later Stone Age occupation levels. The ages for the archaeological sediments were obtained from single-grain measurements that enabled unrepresentative grains to be rejected. The MSA occupation levels have ages that, within error limits, are in stratigraphic order and fall between the OSL age for the oldest dune sand (143.2+/-5.5 ka) and a previously published OSL age for the sterile sand ( approximately 70 ka) that separates the Middle and Later Stone Age deposits. The earliest MSA archaeological phase, M3, from where fragments of ochre were found as well as human teeth, is dated to 98.9+/-4.5 ka, coinciding with the sea-level high of oxygen isotope substage 5c. The cave then appears to be unoccupied until oxygen isotope substage 5a on the basis of four OSL ages for archaeological phase M2, ranging from 84.6+/-5.8 to 76.8+/-3.1 ka; these levels contained large hearths and bone tools. An age of 72.7+/-3.1 ka was obtained for the final MSA archaeological phase, M1, from which deliberately engraved ochre and shell beads were recovered along with bifacial stone points. We conclude that the periods of occupation were determined by changes in sea level, with abundant sources of seafood available in times of high sea level and with the cave being closed by the accumulation of large dunes during periods of low sea level, such as during oxygen isotope stages 4 and 6.     

40Ar/(39)Ar dating of the Kapthurin Formation, Baringo, Kenya.

The(40)Ar/(39)Ar radiometric dating technique has been applied to tuffs and lavas of the Kapthurin Formation in the Tugen Hills, Kenya Rift Valley. Two variants of the(40)Ar/(39)Ar technique, single-crystal total fusion (SCTF) and laser incremental heating (LIH) have been employed to date five marker horizons within the formation: near the base, the Kasurein Basalt at 0.61+/-0.04 Ma; the Pumice Tuff at 0.543+/-0.004 Ma; the Upper Kasurein Basalt at 0.552+/-0.015 Ma; the Grey Tuff at 0.509+/-0.009 Ma; and within the upper part of the formation, the Bedded Tuff at 0.284+/-0.012 Ma. The new, precise radiometric age determination for the Pumice Tuff also provides an age for the widespread Lake Baringo Trachyte, since the Pumice Tuff is the early pyroclastic phase of this voluminous trachyte eruption.These results establish the age of fossil hominids KNM-BK 63-67 and KNM-BK 8518 at approximately 0.510-0.512 Ma, a significant finding given that few Middle Pleistocene hominids are radiometrically dated. The Kapthurin hominids are thus the near contemporaries of those from Bodo, Ethiopia and Tanzania. A flake and core industry from lacustrine sediments in the lower part of the formation is constrained by new dates of 0.55-0.52 Ma, a period during which the Acheulian industry, characterized by handaxes, is known throughout East Africa. Points, typical of the Middle Stone Age (MSA), are found in Kapthurin Formation sediments now shown to date to between 0.509+/-0.009 Ma and 0.284+/-0.012 Ma. This date exceeds previous estimates for the age of the MSA elsewhere in East Africa by 49 ka, and establishes the age of Acheulian to MSA transition for the region. Evidence of the use of the Levallois technique for the manufacture of both small flakes and biface preforms, the systematic production of blades, and the use and processing of red ochre also occurs in this interval. The presence of blades and red ochre at this depth is important as blades signify a high degree of technical competence and red ochre suggests symbolic behavior.     

The Xujiayao 14 Mandibular Ramus and Pleistocene Homo Mandibular Variation

The earlier Late Pleistocene mandibular ramus from Xujiayao (northern China) preserves traits that vary distributionally among western Old World Pleistocene Homo samples and between Early/Middle Pleistocene archaic humans and Late Pleistocene modern humans in eastern Eurasia. Xujiayao 14 presents a lateral mandibular notch crest, an open mandibular foramen, a wide ramus, an asymmetrical mandibular notch, an enlarged superior medial pterygoid tubercle, (probably) a retromolar space, and gonial eversion, as well as an unusual depression in the planum triangulare. The first two traits appear ancestral for Later Pleistocene and recent Homo and are dominant among modern humans. The second two traits largely separate Xujiayao 14 and archaic Homo from modern humans. The next two traits are found in the highest frequency among the Neandertals, although gonial eversion contrasts with Late Pleistocene Neandertals. Xujiayao 14, in the context of Pleistocene and recent Homo samples and the other Xujiayao human remains, therefore provides a morphological mosaic, highlighting regional variation through the Pleistocene.