Defining paleoclimatic routes and opportunities for hominin dispersals across Iran.

Fossil and archaeological evidence indicates that hominin dispersals into Southwest Asia occurred throughout the Pleistocene, including the expansion of Homo sapiens populations out of Africa. While there is evidence for hominin occupations in the Pleistocene in Iran, as evidenced by the presence of Lower to Upper Paleolithic archaeological sites, the extent to which humid periods facilitated population expansions into western Asia has remained unclear. To test the role of humid periods on hominin dispersals here we assess Paleolithic site distributions and paleoenvironmental records across Iran. We developed the first spatially comprehensive, high-resolution paleohydrological model for Iran in order to assess water availability and its influence on hominin dispersals. We highlight environmentally mediated routes which likely played a key role in Late Pleistocene hominin dispersals, including the expansion of H. sapiens and Neanderthals eastwards into Asia. Our combined analyses indicate that, during MIS 5, there were opportunities for hominins to traverse a northern route through the Alborz and Kopet Dagh Mountains and the Dasht-I Kavir desert owing to the presence of activated fresh water sources. We recognize a new southern route along the Zagros Mountains and extending eastwards towards Pakistan and Afghanistan. We find evidence for a potential northern route during MIS 3, which would have permitted hominin movements and species interactions in Southwest Asia. Between humid periods, these interconnections would have waned, isolating populations in the Zagros and Alborz Mountains, where hominins may have continued to have had access to water.

Paleolithic occupation of arid Central Asia in the Middle Pleistocene.

Central Asia is positioned at a crossroads linking several zones important to hominin dispersal during the Middle Pleistocene. However, the scarcity of stratified and dated archaeological material and paleoclimate records makes it difficult to understand dispersal and occupation dynamics during this time period, especially in arid zones. Here we compile and analyze paleoclimatic and archaeological data from Pleistocene Central Asia, including examination of a new layer-counted speleothem-based multiproxy record of hydrological changes in southern Uzbekistan at the end of MIS 11. Our findings indicate that Lower Palaeolithic sites in the steppe, semi-arid, and desert zones of Central Asia may have served as key areas for the dispersal of hominins into Eurasia during the Middle Pleistocene. In agreement with previous studies, we find that bifaces occur across these zones at higher latitudes and in lower altitudes relative to the other Paleolithic assemblages. We argue that arid Central Asia would have been intermittently habitable during the Middle Pleistocene when long warm interglacial phases coincided with periods when the Caspian Sea was experiencing consistently high water levels, resulting in greater moisture availability and more temperate conditions in otherwise arid regions. During periodic intervals in the Middle Pleistocene, the local environment of arid Central Asia was likely a favorable habitat for paleolithic hominins and was frequented by Lower Paleolithic toolmakers producing bifaces.

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.

The expansion of Acheulean hominins into the Nefud Desert of Arabia.

The Arabian Peninsula is a critical geographic landmass situated between Africa and the rest of Eurasia. Climatic shifts across the Pleistocene periodically produced wetter conditions in Arabia, dramatically altering the spatial distribution of hominins both within and between continents. This is particularly true of Acheulean hominins, who appear to have been more tethered to water sources than Middle Palaeolithic hominins. However, until recently, chrono-cultural knowledge of the Acheulean of Arabia has been limited to one dated site, which indicated a hominin presence in Marine Isotope Stages (MIS) 7-6. Here, we report the first dated Acheulean site from the Nefud Desert of northern Saudi Arabia, together with palaeoecological evidence for an associated deep, probably fresh-water, lake. The site of An Nasim features varied and often finely flaked façonnage handaxes. Luminescence ages together with geomorphological and palaeoecological evidence indicates that the associated artefacts date to MIS 9. At present, An Nasim represents the oldest yet documented Acheulean sites in Arabia, and adds to a growing picture of regionally diverse stone tool assemblages used by Middle Pleistocene hominins, and likely indicative of repeated population re-entry into the peninsula in wet 'Green Arabia' phases.

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.

Human footprints provide snapshot of last interglacial ecology in the Arabian interior.

The nature of human dispersals out of Africa has remained elusive because of the poor resolution of paleoecological data in direct association with remains of the earliest non-African people. Here, we report hominin and non-hominin mammalian tracks from an ancient lake deposit in the Arabian Peninsula, dated within the last interglacial. The findings, it is argued, likely represent the oldest securely dated evidence for Homo sapiens in Arabia. The paleoecological evidence indicates a well-watered semi-arid grassland setting during human movements into the Nefud Desert of Saudi Arabia. We conclude that visitation to the lake was transient, likely serving as a place to drink and to forage, and that late Pleistocene human and mammalian migrations and landscape use patterns in Arabia were inexorably linked.

Human responses to climate and ecosystem change in ancient Arabia.

Recent interdisciplinary archaeological and paleoenvironmental research in the Arabian peninsula is transforming our understanding of ancient human societies in their ecological contexts. Hypotheses about the cultural and demographic impacts of a series of droughts have primarily been developed from the environmental and archaeological records of southeastern Arabia. Here we examine these human-environment interactions by integrating ongoing research from northern Arabia. While droughts and extreme environmental variability in the Holocene had significant impacts on human societies, responses varied across space and time and included mobility at various scales, as well as diverse social, economic and cultural adaptations, such as the management of water resources, the introduction of pastoral lifeways, and the construction of diverse types of stone structures. The long-term story of human societies in Arabia is one of resilience in the face of climate change, yet future challenges include rising temperatures and flash flooding. The history of human responses to climatic and ecosystem changes in Arabia can provide important lessons for a planet facing catastrophic global warming and environmental change.

The expansion of later Acheulean hominins into the Arabian Peninsula.

The Acheulean is the longest lasting cultural-technological tradition in human evolutionary history. However, considerable gaps remain in understanding the chronology and geographical distribution of Acheulean hominins. We present the first chronometrically dated Acheulean site from the Arabian Peninsula, a vast and poorly known region that forms more than half of Southwest Asia. Results show that Acheulean hominin occupation expanded along hydrological networks into the heart of Arabia from Marine Isotope Stage (MIS) 7 until at least ~190 ka ̶ the youngest documented Acheulean in Southwest Asia. The site of Saffaqah features Acheulean technology, characterized by large flakes, handaxes and cleavers, similar to Acheulean assemblages in Africa. These findings reveal a climatically-mediated later Acheulean expansion into a poorly known region, amplifying the documented diversity of Middle Pleistocene hominin behaviour across the Old World and elaborating the terminal archaic landscape encountered by our species as they dispersed out of Africa.

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

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

Acheulean technology and landscape use at Dawadmi, central Arabia.

Despite occupying a central geographic position, investigations of hominin populations in the Arabian Peninsula during the Lower Palaeolithic period are rare. The colonization of Eurasia below 55 degrees latitude indicates the success of the genus Homo in the Early and Middle Pleistocene, but the extent to which these hominins were capable of innovative and novel behavioural adaptations to engage with mid-latitude environments is unclear. Here we describe new field investigations at the Saffaqah locality (206-76) near Dawadmi, in central Arabia that aim to establish how hominins adapted to this region. The site is located in the interior of Arabia over 500 km from both the Red Sea and the Gulf, and at the headwaters of two major extinct river systems that were likely used by Acheulean hominins to cross the Peninsula. Saffaqah is one of the largest Acheulean sites in Arabia with nearly a million artefacts estimated to occur on the surface, and it is also the first to yield stratified deposits containing abundant artefacts. It is situated in the unusual setting of a dense and well-preserved landscape of Acheulean localities, with sites and isolated artefacts occurring regularly for tens of kilometres in every direction. We describe both previous and recent excavations at Saffaqah and its large lithic assemblage. We analyse thousands of artefacts from excavated and surface contexts, including giant andesite cores and flakes, smaller cores and retouched artefacts, as well as handaxes and cleavers. Technological assessment of stratified lithics and those from systematic survey, enable the reconstruction of stone tool life histories. The Acheulean hominins at Dawadmi were strong and skilful, with their adaptation evidently successful for some time. However, these biface-makers were also technologically conservative, and used least-effort strategies of resource procurement and tool transport. Ultimately, central Arabia was depopulated, likely in the face of environmental deterioration in the form of increasing aridity.

Homo sapiens in Arabia by 85,000 years ago.

Understanding the timing and character of the expansion of Homo sapiens out of Africa is critical for inferring the colonization and admixture processes that underpin global population history. It has been argued that dispersal out of Africa had an early phase, particularly ~130-90 thousand years ago (ka), that reached only the East Mediterranean Levant, and a later phase, ~60-50 ka, that extended across the diverse environments of Eurasia to Sahul. However, recent findings from East Asia and Sahul challenge this model. Here we show that H. sapiens was in the Arabian Peninsula before 85 ka. We describe the Al Wusta-1 (AW-1) intermediate phalanx from the site of Al Wusta in the Nefud desert, Saudi Arabia. AW-1 is the oldest directly dated fossil of our species outside Africa and the Levant. The palaeoenvironmental context of Al Wusta demonstrates that H. sapiens using Middle Palaeolithic stone tools dispersed into Arabia during a phase of increased precipitation driven by orbital forcing, in association with a primarily African fauna. A Bayesian model incorporating independent chronometric age estimates indicates a chronology for Al Wusta of ~95-86 ka, which we correlate with a humid episode in the later part of Marine Isotope Stage 5 known from various regional records. Al Wusta shows that early dispersals were more spatially and temporally extensive than previously thought. Early H. sapiens dispersals out of Africa were not limited to winter rainfall-fed Levantine Mediterranean woodlands immediately adjacent to Africa, but extended deep into the semi-arid grasslands of Arabia, facilitated by periods of enhanced monsoonal rainfall.

Rethinking the dispersal of Homo sapiens out of Africa.

Current fossil, genetic, and archeological data indicate that Homo sapiens originated in Africa in the late Middle Pleistocene. By the end of the Late Pleistocene, our species was distributed across every continent except Antarctica, setting the foundations for the subsequent demographic and cultural changes of the Holocene. The intervening processes remain intensely debated and a key theme in hominin evolutionary studies. We review archeological, fossil, environmental, and genetic data to evaluate the current state of knowledge on the dispersal of Homo sapiens out of Africa. The emerging picture of the dispersal process suggests dynamic behavioral variability, complex interactions between populations, and an intricate genetic and cultural legacy. This evolutionary and historical complexity challenges simple narratives and suggests that hybrid models and the testing of explicit hypotheses are required to understand the expansion of Homo sapiens into Eurasia.