Reconstructing Prehistoric African Population Structure.

We assembled genome-wide data from 16 prehistoric Africans. We show that the anciently divergent lineage that comprises the primary ancestry of the southern African San had a wider distribution in the past, contributing approximately two-thirds of the ancestry of Malawi hunter-gatherers ∼8,100-2,500 years ago and approximately one-third of the ancestry of Tanzanian hunter-gatherers ∼1,400 years ago. We document how the spread of farmers from western Africa involved complete replacement of local hunter-gatherers in some regions, and we track the spread of herders by showing that the population of a ∼3,100-year-old pastoralist from Tanzania contributed ancestry to people from northeastern to southern Africa, including a ∼1,200-year-old southern African pastoralist. The deepest diversifications of African lineages were complex, involving either repeated gene flow among geographically disparate groups or a lineage more deeply diverging than that of the San contributing more to some western African populations than to others. We finally leverage ancient genomes to document episodes of natural selection in southern African populations. PAPERCLIP.

Multiple hominin dispersals into Southwest Asia over the past 400,000 years.

Pleistocene hominin dispersals out of, and back into, Africa necessarily involved traversing the diverse and often challenging environments of Southwest Asia1-4. Archaeological and palaeontological records from the Levantine woodland zone document major biological and cultural shifts, such as alternating occupations by Homo sapiens and Neanderthals. However, Late Quaternary cultural, biological and environmental records from the vast arid zone that constitutes most of Southwest Asia remain scarce, limiting regional-scale insights into changes in hominin demography and behaviour1,2,5. Here we report a series of dated palaeolake sequences, associated with stone tool assemblages and vertebrate fossils, from the Khall Amayshan 4 and Jubbah basins in the Nefud Desert. These findings, including the oldest dated hominin occupations in Arabia, reveal at least five hominin expansions into the Arabian interior, coinciding with brief 'green' windows of reduced aridity approximately 400, 300, 200, 130-75 and 55 thousand years ago. Each occupation phase is characterized by a distinct form of material culture, indicating colonization by diverse hominin groups, and a lack of long-term Southwest Asian population continuity. Within a general pattern of African and Eurasian hominin groups being separated by Pleistocene Saharo-Arabian aridity, our findings reveal the tempo and character of climatically modulated windows for dispersal and admixture.

Earliest known human burial in Africa.

The origin and evolution of hominin mortuary practices are topics of intense interest and debate1-3. Human burials dated to the Middle Stone Age (MSA) are exceedingly rare in Africa and unknown in East Africa1-6. Here we describe the partial skeleton of a roughly 2.5- to 3.0-year-old child dating to 78.3 ± 4.1 thousand years ago, which was recovered in the MSA layers of Panga ya Saidi (PYS), a cave site in the tropical upland coast of Kenya7,8. Recent excavations have revealed a pit feature containing a child in a flexed position. Geochemical, granulometric and micromorphological analyses of the burial pit content and encasing archaeological layers indicate that the pit was deliberately excavated. Taphonomical evidence, such as the strict articulation or good anatomical association of the skeletal elements and histological evidence of putrefaction, support the in-place decomposition of the fresh body. The presence of little or no displacement of the unstable joints during decomposition points to an interment in a filled space (grave earth), making the PYS finding the oldest known human burial in Africa. The morphological assessment of the partial skeleton is consistent with its assignment to Homo sapiens, although the preservation of some primitive features in the dentition supports increasing evidence for non-gradual assembly of modern traits during the emergence of our species. The PYS burial sheds light on how MSA populations interacted with the dead.

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

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

Trajectories of cultural innovation from the Middle to Later Stone Age in Eastern Africa: Personal ornaments, bone artifacts, and ocher from Panga ya Saidi, Kenya.

African Middle Stone Age (MSA) populations used pigments, manufactured and wore personal ornaments, made abstract engravings, and produced fully shaped bone tools. However, ongoing research across Africa reveals variability in the emergence of cultural innovations in the MSA and their subsequent development through the Later Stone Age (LSA). When present, it appears that cultural innovations manifest regional variability, suggestive of distinct cultural traditions. In eastern Africa, several Late Pleistocene sites have produced evidence for novel activities, but the chronologies of key behavioral innovations remain unclear. The 3 m deep, well-dated, Panga ya Saidi sequence in eastern Kenya, encompassing 19 layers covering a time span of 78 kyr beginning in late Marine Isotope Stage 5, is the only known African site recording the interplay between cultural and ecological diversity in a coastal forested environment. Excavations have yielded worked and incised bones, ostrich eggshell beads (OES), beads made from seashells, worked and engraved ocher pieces, fragments of coral, and a belemnite fossil. Here, we provide, for the first time, a detailed analysis of this material. This includes a taphonomic, archeozoological, technological, and functional study of bone artifacts; a technological and morphometric analysis of personal ornaments; and a technological and geochemical analysis of ocher pieces. The interpretation of the results stemming from the analysis of OES beads is guided by an ethnoarcheological perspective and field observations. We demonstrate that key cultural innovations on the eastern African coast are evident by 67 ka and exhibit remarkable diversity through the LSA and Iron Age. We suggest the cultural trajectories evident at Panga ya Saidi were shaped by both regional traditions and cultural/demic diffusion.

Ancient crops provide first archaeological signature of the westward Austronesian expansion.

The Austronesian settlement of the remote island of Madagascar remains one of the great puzzles of Indo-Pacific prehistory. Although linguistic, ethnographic, and genetic evidence points clearly to a colonization of Madagascar by Austronesian language-speaking people from Island Southeast Asia, decades of archaeological research have failed to locate evidence for a Southeast Asian signature in the island's early material record. Here, we present new archaeobotanical data that show that Southeast Asian settlers brought Asian crops with them when they settled in Africa. These crops provide the first, to our knowledge, reliable archaeological window into the Southeast Asian colonization of Madagascar. They additionally suggest that initial Southeast Asian settlement in Africa was not limited to Madagascar, but also extended to the Comoros. Archaeobotanical data may support a model of indirect Austronesian colonization of Madagascar from the Comoros and/or elsewhere in eastern Africa.

Childhood and the evolution of higher-effort teaching.

Kline presents an excellent synthesis of teaching theory and research, with cogent arguments regarding its prevalence. In this, she claims that "active teaching" is human specific, and presents tangible reasons why. But in doing so, she overlooks a critical aspect of the human condition that may have arisen only recently in our evolutionary history: Childhood as a life stage.

Abrupt onset of intensive human occupation 44,000 years ago on the threshold of Sahul.

Archaeological evidence attests multiple early dispersals of Homo sapiens out of Africa, but genetic evidence points to the primacy of a single dispersal 70-40 ka. Laili in Timor-Leste is on the southern dispersal route between Eurasia and Australasia and has the earliest record of human occupation in the eastern Wallacean archipelago. New evidence from the site shows that, unusually in the region, sediment accumulated in the shelter without human occupation, in the window 59-54 ka. This was followed by an abrupt onset of intensive human habitation beginning ~44 ka. The initial occupation is distinctive from overlying layers in the aquatic focus of faunal exploitation, while it has similarities in material culture to other early Homo sapiens sites in Wallacea. We suggest that the intensive early occupation at Laili represents a colonisation phase, which may have overwhelmed previous human dispersals in this part of the world.

Plant, pigment, and bone processing in the Neolithic of northern Arabia-New evidence from Use-wear analysis of grinding tools at Jebel Oraf.

Archaeological sites with surface hearths are a ubiquitous feature across the arid zones of the Arabian interior. At Jebel Oraf, in the Jubbah basin of the Nefud Desert of northern Arabia, numerous grinding stone fragments were found in association with hearths, though the original purpose of these stones was unclear owing to the poor preservation of faunal and botanic remains. Here we describe results from use-wear analysis on five grinding tools at Jebel Oraf, demonstrating that such artefacts were used during the Neolithic for plant processing, bone processing, and pigment production. Grinding stones were often broken up after initial use and fragments were subsequently re-used for alternative purposes, before finally being placed on hearths or discarded. More specifically, plants were ground or prepared and possibly cooked in the hearths, and bones were processed as well. The analyses also highlight the importance of pigment processing at Neolithic sites and provide a link to painted rock art. The frequent use of pigment in the archaeological record suggests that pigment was widely used, and that Neolithic painted art may have been more common than the surviving images suggest.

Stone toolmaking difficulty and the evolution of hominin technological skills.

Stone tools are a manifestation of the complex cognitive and dexterous skills of our hominin ancestors. As such, much research has been devoted to understanding the skill requirements of individual lithic technologies. Yet, comparing skill across different technologies, and thus across the vast timespan of the Palaeolithic, is an elusive goal. We seek to quantify a series of commensurable metrics of knapping skill across four different lithic technologies (discoids, handaxes, Levallois, and prismatic blades). To compare the requisite dexterity, coordination, and care involved in each technology, we analysed video footage and lithic material from a series of replicative knapping experiments to quantify deliberation (strike time), precision (platform area), intricacy (flake size relative to core size), and success (relative blank length). According to these four metrics, discoidal knapping appears to be easiest among the sample. Levallois knapping involved an intricate reduction sequence, but did not require as much motor control as handaxes and especially prismatic blades. Compared with the other Palaeolithic technologies, we conclude that prismatic blade knapping is set apart by being a skill intensive means of producing numerous standardised elongate end-products.

Talking Dead. New burials from Tron Bon Lei (Alor Island, Indonesia) inform on the evolution of mortuary practices from the terminal Pleistocene to the Holocene in Southeast Asia.

Burial elaborations are a human behaviour that, in recent contexts can inform on social diversification, belief systems, and the introduction of new practices resulting from migration or cultural transmission. The study of mortuary practices in Mainland and Island Southeast Asia has revealed complex and diverse treatments of the deceased. This paper contributes to this topic with the description of three new burials excavated in Tron Bon Lei (Alor Island, Indonesia) dated to 7.5, 10, and 12 kya cal BP. In addition to the bioskeletal profiles and palaeohealth observations, we propose the adoption of archaeothanatological methods to characterise burial types in the region. Through the analysis of skeletal element representation, body position, articulation, and grave associations, we provide an example of a holistic approach to mortuary treatments in the Lesser Sunda Islands. Our results provide significant new data for understanding the evolution and diversification of burial practices in Southeast Asia, contributing to a growing body of literature describing prehistoric socio-cultural behaviour in this region.

67,000 years of coastal engagement at Panga ya Saidi, eastern Africa.

The antiquity and nature of coastal resource procurement is central to understanding human evolution and adaptations to complex environments. It has become increasingly apparent in global archaeological studies that the timing, characteristics, and trajectories of coastal resource use are highly variable. Within Africa, discussions of these issues have largely been based on the archaeological record from the south and northeast of the continent, with little evidence from eastern coastal areas leaving significant spatial and temporal gaps in our knowledge. Here, we present data from Panga ya Saidi, a limestone cave complex located 15 km from the modern Kenyan coast, which represents the first long-term sequence of coastal engagement from eastern Africa. Rather than attempting to distinguish between coastal resource use and coastal adaptations, we focus on coastal engagement as a means of characterising human relationships with marine environments and resources from this inland location. We use aquatic mollusc data spanning the past 67,000 years to document shifts in the acquisition, transportation, and discard of these materials, to better understand long-term trends in coastal engagement. Our results show pulses of coastal engagement beginning with low-intensity symbolism, and culminating in the consistent low-level transport of marine and freshwater food resources, emphasising a diverse relationship through time. Panga ya Saidi has the oldest stratified evidence of marine engagement in eastern Africa, and is the only site in Africa which documents coastal resources from the Late Pleistocene through the Holocene, highlighting the potential archaeological importance of peri-coastal sites to debates about marine resource relationships.

Collagen fingerprinting traces the introduction of caprines to island Eastern Africa.

The human colonization of eastern Africa's near- and offshore islands was accompanied by the translocation of several domestic, wild and commensal fauna, many of which had long-term impacts on local environments. To better understand the timing and nature of the introduction of domesticated caprines (sheep and goat) to these islands, this study applied collagen peptide fingerprinting (Zooarchaeology by Mass Spectrometry or ZooMS) to archaeological remains from eight Iron Age sites, dating between ca 300 and 1000 CE, in the Zanzibar, Mafia and Comoros archipelagos. Where previous zooarchaeological analyses had identified caprine remains at four of these sites, this study identified goat at seven sites and sheep at three, demonstrating that caprines were more widespread than previously known. The ZooMS results support an introduction of goats to island eastern Africa from at least the seventh century CE, while sheep in our sample arrived one-two centuries later. Goats may have been preferred because, as browsers, they were better adapted to the islands' environments. The results allow for a more accurate understanding of early caprine husbandry in the study region and provide a critical archaeological baseline for examining the potential long-term impacts of translocated fauna on island ecologies.

Homo neanderthalensis and the evolutionary origins of ritual in Homo sapiens.

There is a large, if disparate, body of archaeological literature discussing specific instantiations of symbolic material culture and the possibility of ritual practices in Neanderthal populations. Despite this attention, however, no single synthesis exists that draws upon cognitive, psychological and cultural evolutionary theories of ritual. Here, we review the evidence for ritual-practice among now-extinct Homo neanderthalensis, as well as the necessary cognitive pre-conditions for such behaviour, in order to explore the evolution of ritual in Homo sapiens. We suggest that the currently available archaeological evidence indicates that Neanderthals may have used 'ritualization' to increase the successful transmission of technical knowledge across generations-providing an explanation for the technological stability of the Middle Palaeolithic and attesting to a survival strategy differing from near-contemporary H. sapiens. This article is part of the theme issue 'Ritual renaissance: new insights into the most human of behaviours'.

Early ground axe technology in Wallacea: The first excavations on Obi Island.

The first excavations on Obi Island, north-east Wallacea, reveal three phases of occupation beginning in the terminal Pleistocene. Ground shell artefacts appear at the end of the terminal Pleistocene, the earliest examples in Wallacea. In the subsequent early Holocene occupation phase, ground stone axe flakes appear, which are again the earliest examples in Wallacea. Ground axes were likely instrumental to subsistence in Obi's dense tropical forest. From ~8000 BP there was a hiatus lasting several millennia, perhaps because increased precipitation and forest density made the sites inhospitable. The site was reoccupied in the Metal Age, with this third phase including quadrangular ground stone artefacts, as well as pottery and pigs; reflecting Austronesian influences. Greater connectivity at this time is also indicated by an Oliva shell bead tradition that occurs in southern Wallacea and an exotic obsidian artefact. The emergence of ground axes on Obi is an independent example of a broader pattern of intensification at the Pleistocene-Holocene transition in Wallacea and New Guinea, evincing human innovation in response to rapid environmental change.

Monumental landscapes of the Holocene humid period in Northern Arabia: The mustatil phenomenon.

Between 10 and six thousand years ago the Arabian Peninsula saw the most recent of the 'Green Arabia' periods, when increased rainfall transformed this generally arid region. The transition to the Neolithic in Arabia occurred during this period of climatic amelioration. Various forms of stone structures are abundant in northern Arabia, and it has been speculated that some of these dated to the Neolithic, but there has been little research on their character and chronology. Here we report a study of 104 'mustatil' stone structures from the southern margins of the Nefud Desert in northern Arabia. We provide the first chronometric age estimate for this type of structure - a radiocarbon date of ca. 5000 BC - and describe their landscape positions, architecture and associated material culture and faunal remains. The structure we have dated is the oldest large-scale stone structure known from the Arabian Peninsula. The mustatil phenomenon represents a remarkable development of monumental architecture, as hundreds of these structures were built in northwest Arabia. This 'monumental landscape' represents one of the earliest large-scale forms of monumental stone structure construction anywhere in the world. Further research is needed to understand the function of these structures, but we hypothesise that they were related to rituals in the context of the adoption of pastoralism and resulting territoriality in the challenging environments of northern Arabia.

Isotopic evidence for initial coastal colonization and subsequent diversification in the human occupation of Wallacea.

The resource-poor, isolated islands of Wallacea have been considered a major adaptive obstacle for hominins expanding into Australasia. Archaeological evidence has hinted that coastal adaptations in Homo sapiens enabled rapid island dispersal and settlement; however, there has been no means to directly test this proposition. Here, we apply stable carbon and oxygen isotope analysis to human and faunal tooth enamel from six Late Pleistocene to Holocene archaeological sites across Wallacea. The results demonstrate that the earliest human forager found in the region c. 42,000 years ago made significant use of coastal resources prior to subsequent niche diversification shown for later individuals. We argue that our data provides clear insights into the huge adaptive flexibility of our species, including its ability to specialize in the use of varied environments, particularly in comparison to other hominin species known from Island Southeast Asia.

Human occupation of northern India spans the Toba super-eruption ~74,000 years ago.

India is located at a critical geographic crossroads for understanding the dispersal of Homo sapiens out of Africa and into Asia and Oceania. Here we report evidence for long-term human occupation, spanning the last ~80 thousand years, at the site of Dhaba in the Middle Son River Valley of Central India. An unchanging stone tool industry is found at Dhaba spanning the Toba eruption of ~74 ka (i.e., the Youngest Toba Tuff, YTT) bracketed between ages of 79.6 ± 3.2 and 65.2 ± 3.1 ka, with the introduction of microlithic technology ~48 ka. The lithic industry from Dhaba strongly resembles stone tool assemblages from the African Middle Stone Age (MSA) and Arabia, and the earliest artefacts from Australia, suggesting that it is likely the product of Homo sapiens as they dispersed eastward out of Africa.

Correction: Acheulean technology and landscape use at Dawadmi, central Arabia.

[This corrects the article DOI: 10.1371/journal.pone.0200497.].