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Recent advances in interdisciplinary archaeological research in Arabia have focused on the evolution and historical development of regional human populations as well as the diverse patterns of cultural change, migration, and adaptations to environmental fluctuations. Obtaining a comprehensive understanding of cultural developments such as the emergence and lifeways of Neolithic groups has been hindered by the limited preservation of stratified archaeological assemblages and organic remains, a common challenge in arid environments. Underground settings like caves and lava tubes, which are prevalent in Arabia but which have seen limited scientific exploration, offer promising opportunities for addressing these issues. Here, we report on an archaeological excavation and a related survey at and around Umm Jirsan lava tube in the Harrat Khaybar, north-western Saudi Arabia. Our results reveal repeated phases of human occupation of the site ranging from at least the Neolithic through to the Chalcolithic/Bronze Age. Pastoralist use of the lava tube and surrounding landscape is attested in rock art and faunal records, suggesting that Umm Jirsan was situated along a pastoral route linking key oases. Isotopic data indicates that herbivores primarily grazed on wild grasses and shrubs rather than being provided with fodder, while humans had a diet consistently high in protein but with increasing consumption of C3 plants through-time, perhaps related to the emergence of oasis agriculture. While underground and naturally sheltered localities are globally prominent in archaeology and Quaternary science, our work represents the first such combined records for Saudi Arabia and highlight the potential for interdisciplinary studies in caves and lava tubes., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2024 Stewart et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

The behavioral origins of Homo sapiens can be traced back to the first material culture produced by our species in Africa, the Middle Stone Age (MSA). Beyond this broad consensus, the origins, patterns, and causes of behavioral complexity in modern humans remain debated. Here, we consider whether recent findings continue to support popular scenarios of: (1) a modern human 'package,' (2) a gradual and 'pan-African' emergence of behavioral complexity, and (3) a direct connection to changes in the human brain. Our geographically structured review shows that decades of scientific research have continuously failed to find a discrete threshold for a complete 'modernity package' and that the concept is theoretically obsolete. Instead of a continent-wide, gradual accumulation of complex material culture, the record exhibits a predominantly asynchronous presence and duration of many innovations across different regions of Africa. The emerging pattern of behavioral complexity from the MSA conforms to an intricate mosaic characterized by spatially discrete, temporally variable, and historically contingent trajectories. This archaeological record bears no direct relation to a simplistic shift in the human brain but rather reflects similar cognitive capacities that are variably manifested. The interaction of multiple causal factors constitutes the most parsimonious explanation driving the variable expression of complex behaviors, with demographic processes such as population structure, size, and connectivity playing a key role. While much emphasis has been given to innovation and variability in the MSA record, long periods of stasis and a lack of cumulative developments argue further against a strictly gradualistic nature in the record. Instead, we are confronted with humanity's deep, variegated roots in Africa, and a dynamic metapopulation that took many millennia to reach the critical mass capable of producing the ratchet effect commonly used to define contemporary human culture. Finally, we note a weakening link between 'modern' human biology and behavior from around 300 ka ago., Competing Interests: Declaration of competing interest The authors declare no conflict of interest., (Copyright © 2023 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

The Ihò Eléérú (or Iho Eleru) rock shelter, located in Southwest Nigeria, is the only site from which Pleistocene-age hominin fossils have been recovered in western Africa. Excavations at Iho Eleru revealed regular human occupations ranging from the Later Stone Age (LSA) to the present day. Here, we present chronometric, archaeobotanical, and paleoenvironmental findings, which include the taxonomic, taphonomic, and isotopic analyses of what is the only Pleistocene faunal assemblage documented in western Africa. Our results indicate that the local landscape surrounding Iho Eleru, although situated within a regional open-canopy biome, was forested throughout the past human occupation of the site. At a regional scale, a shift from forest- to savanna-dominated ecotonal environment occurred during a mid-Holocene warm event 6,000 years ago, with a subsequent modern reforestation of the landscape. Locally, no environmental shift was observable, placing Iho Eleru in a persistent forested "island" during the period of occupation., Competing Interests: The authors declare no competing interest., (© 2023 The Author(s).)

Since Darwin, studies of human evolution have tended to give primacy to open 'savannah' environments as the ecological cradle of our lineage, with dense tropical forests cast as hostile, unfavourable frontiers. These perceptions continue to shape both the geographical context of fieldwork as well as dominant narratives concerning hominin evolution. This paradigm persists despite new, ground-breaking research highlighting the role of tropical forests in the human story. For example, novel research in Africa's rainforests has uncovered archaeological sites dating back into the Pleistocene; genetic studies have revealed very deep human roots in Central and West Africa and in the tropics of Asia and the Pacific; an unprecedented number of coexistent hominin species have now been documented, including Homo erectus , the 'Hobbit' ( Homo floresiensis ), Homo luzonensis , Denisovans, and Homo sapiens . Some of the earliest members of our own species to reach South Asia, Southeast Asia, Oceania and the tropical Americas have shown an unexpected rapidity in their adaptation to even some of the more 'extreme' tropical settings. This includes the early human manipulation of species and even habitats. This volume builds on these currently disparate threads and, for the first time, draws together a group of interdisciplinary, agenda-setting papers that firmly places a broader spectrum of tropical environments at the heart of the deep human past. This article is part of the theme issue 'Tropical forests in the deep human past'.

The most profound shift in the African hydroclimate of the last 1 million years occurred around 300 thousand years (ka) ago. This change in African hydroclimate is manifest as an east-west change in moisture balance that cannot be fully explained through linkages to high latitude climate systems. The east-west shift is, instead, probably driven by a shift in the tropical Walker Circulation related to sea surface temperature change driven by orbital forcing. Comparing records of past vegetation change, and hominin evolution and development, with this breakpoint in the climate system is challenging owing to the paucity of study sites available and uncertainties regarding the dating of records. Notwithstanding these uncertainties we find that, broadly speaking, both vegetation and hominins change around 300 ka. The vegetative backdrop suggests that relative abundance of vegetative resources shifted from western to eastern Africa, although resources would have persisted across the continent. The climatic and vegetation changes probably provided challenges for hominins and are broadly coincident with the appearance of Homo sapiens ( ca 315 ka) and the emergence of Middle Stone Age technology. The concomitant changes in climate, vegetation and hominin evolution suggest that these factors are closely intertwined. This article is part of the theme issue 'Tropical forests in the deep human past'.

Homo sapiens have adapted to an incredible diversity of habitats around the globe. This capacity to adapt to different landscapes is clearly expressed within Africa, with Late Pleistocene Homo sapiens populations occupying savannahs, woodlands, coastlines and mountainous terrain. As the only area of the world where Homo sapiens have clearly persisted through multiple glacial-interglacial cycles, Africa is the only continent where classic refugia models can be formulated and tested to examine and describe changing patterns of past distributions and human phylogeographies. The potential role of refugia has frequently been acknowledged in the Late Pleistocene palaeoanthropological literature, yet explicit identification of potential refugia has been limited by the patchy nature of palaeoenvironmental and archaeological records, and the low temporal resolution of climate or ecological models. Here, we apply potential climatic thresholds on human habitation, rooted in ethnographic studies, in combination with high-resolution model datasets for precipitation and biome distributions to identify persistent refugia spanning the Late Pleistocene (130-10 ka). We present two alternate models suggesting that between 27% and 66% of Africa may have provided refugia to Late Pleistocene human populations, and examine variability in precipitation, biome and ecotone distributions within these refugial zones. This article is part of the theme issue 'Tropical forests in the deep human past'.

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., Competing Interests: The authors declare no competing interests., (© 2021 The Authors.)

Pleistocene hominin dispersals out of, and back into, Africa necessarily involved traversing the diverse and often challenging environments of Southwest Asia 1-4 . 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 behaviour 1,2,5 . 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., (© 2021. The Author(s).)

In this study, we synthesize terrestrial and marine proxy records, spanning the past 620 ky, to decipher pan-African climate variability and its drivers and potential linkages to hominin evolution. We find a tight correlation between moisture availability across Africa to El Niño Southern Ocean oscillation (ENSO) variability, a manifestation of the Walker Circulation, that was most likely driven by changes in Earth's eccentricity. Our results demonstrate that low-latitude insolation was a prominent driver of pan-African climate change during the Middle to Late Pleistocene. We argue that these low-latitude climate processes governed the dispersion and evolution of vegetation as well as mammals in eastern and western Africa by increasing resource-rich and stable ecotonal settings thought to have been important to early modern humans., Competing Interests: The authors declare no competing interest.

Marine Isotope Stage (MIS) 5, ~ 130 to 71 thousand years ago, was a key period for the geographic expansion of Homo sapiens, including engagement with new landscapes within Africa and dispersal into Asia. Occupation of the Levant by Homo sapiens in MIS 5 is well established, while recent research has documented complementary evidence in Arabia. Here, we undertake the first detailed comparison of Levallois core technology from eastern Africa, Arabia, and the Levant during MIS 5, including multiple sites associated with Homo sapiens fossils. We employ quantitative comparisons of individual artefacts that provides a detailed appraisal of Levallois reduction activity in MIS 5, thereby enabling assessment of intra- and inter-assemblage variability for the first time. Our results demonstrate a pattern of geographically structured variability embedded within a shared focus on centripetal Levallois reduction schemes and overlapping core morphologies. We reveal directional changes in core shaping and flake production from eastern Africa to Arabia and the Levant that are independent of differences in geographic or environmental parameters. These results are consistent with a common cultural inheritance between these regions, potentially stemming from a shared late Middle Pleistocene source in eastern Africa.

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.

Neanderthals occurred widely across north Eurasian landscapes, but between ~ 70 and 50 thousand years ago (ka) they expanded southwards into the Levant, which had previously been inhabited by Homo sapiens. Palaeoanthropological research in the first half of the twentieth century demonstrated alternate occupations of the Levant by Neanderthal and Homo sapiens populations, yet key early findings have largely been overlooked in later studies. Here, we present the results of new examinations of both the fossil and archaeological collections from Shukbah Cave, located in the Palestinian West Bank, presenting new quantitative analyses of a hominin lower first molar and associated stone tool assemblage. The hominin tooth shows clear Neanderthal affinities, making it the southernmost known fossil specimen of this population/species. The associated Middle Palaeolithic stone tool assemblage is dominated by Levallois reduction methods, including the presence of Nubian Levallois points and cores. This is the first direct association between Neanderthals and Nubian Levallois technology, demonstrating that this stone tool technology should not be considered an exclusive marker of Homo sapiens.

New finds in the palaeoanthropological and genomic records have changed our view of the origins of modern human ancestry. Here we review our current understanding of how the ancestry of modern humans around the globe can be traced into the deep past, and which ancestors it passes through during our journey back in time. We identify three key phases that are surrounded by major questions, and which will be at the frontiers of future research. The most recent phase comprises the worldwide expansion of modern humans between 40 and 60 thousand years ago (ka) and their last known contacts with archaic groups such as Neanderthals and Denisovans. The second phase is associated with a broadly construed African origin of modern human diversity between 60 and 300 ka. The oldest phase comprises the complex separation of modern human ancestors from archaic human groups from 0.3 to 1 million years ago. We argue that no specific point in time can currently be identified at which modern human ancestry was confined to a limited birthplace, and that patterns of the first appearance of anatomical or behavioural traits that are used to define Homo sapiens are consistent with a range of evolutionary histories.

The African Middle Stone Age (MSA, typically considered to span ca. 300-30 thousand years ago [ka]), represents our species' first and longest lasting cultural phase. Although the MSA to Later Stone Age (LSA) transition is known to have had a degree of spatial and temporal variability, recent studies have implied that in some regions, the MSA persisted well beyond 30 ka. Here we report two new sites in Senegal that date the end of the MSA to around 11 ka, the youngest yet documented MSA in Africa. This shows that this cultural phase persisted into the Holocene. These results highlight significant spatial and temporal cultural variability in the African Late Pleistocene, consistent with genomic and palaeoanthropological hypotheses that significant, long-standing inter-group cultural differences shaped the later stages of human evolution in Africa.

Despite its largely hyper-arid and inhospitable climate today, the Arabian Peninsula is emerging as an important area for investigating Pleistocene hominin dispersals. Recently, a member of our own species was found in northern Arabia dating to ca. 90 ka, while stone tools and fossil finds have hinted at an earlier, middle Pleistocene, hominin presence. However, there remain few direct insights into Pleistocene environments, and associated hominin adaptations, that accompanied the movement of populations into this region. Here, we apply stable carbon and oxygen isotope analysis to fossil mammal tooth enamel (n = 21) from the middle Pleistocene locality of Ti's al Ghadah in Saudi Arabia associated with newly discovered stone tools and probable cutmarks. The results demonstrate productive grasslands in the interior of the Arabian Peninsula ca. 300-500 ka, as well as aridity levels similar to those found in open savannah settings in eastern Africa today. The association between this palaeoenvironmental information and the earliest traces for hominin activity in this part of the world lead us to argue that middle Pleistocene hominin dispersals into the interior of the Arabian Peninsula required no major novel adaptation.

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.

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

We challenge the view that our species, Homo sapiens, evolved within a single population and/or region of Africa. The chronology and physical diversity of Pleistocene human fossils suggest that morphologically varied populations pertaining to the H. sapiens clade lived throughout Africa. Similarly, the African archaeological record demonstrates the polycentric origin and persistence of regionally distinct Pleistocene material culture in a variety of paleoecological settings. Genetic studies also indicate that present-day population structure within Africa extends to deep times, paralleling a paleoenvironmental record of shifting and fractured habitable zones. We argue that these fields support an emerging view of a highly structured African prehistory that should be considered in human evolutionary inferences, prompting new interpretations, questions, and interdisciplinary research directions., (Copyright © 2018 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

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., Competing Interests: The authors have declared that no competing interests exist.

The North African Middle Stone Age (NAMSA, ∼300-24 thousand years ago, or ka) features what may be the oldest fossils of our species as well as extremely early examples of technological regionalization and 'symbolic' material culture (d'Errico, Vanhaeren, Barton, Bouzouggar, Mienis, Richter, Hublin, McPherron, Louzouet, & Klein, ; Scerri, ; Richter, Grün, Joannes-Boyau, Steele, Amani, Rué, Fernandes, Raynal, Geraads, Ben-Ncer Hublin, McPherron, ). The geographic situation of North Africa and an increased understanding of the wet-dry climatic pulses of the Sahara Desert also show that North Africa played a strategic role in continental-scale evolutionary processes by modulating human dispersal and demographic structure (Drake, Blench, Armitage, Bristow, & White, ; Blome, Cohen, Tryon, Brooks, & Russell, ). However, current understanding of the NAMSA remains patchy and subject to a bewildering array of industrial nomenclatures that mask underlying variability. These issues are compounded by a geographic research bias skewed toward non-desert regions. As a result, it has been difficult to test long-established narratives of behavioral and evolutionary change in North Africa and to resolve debates on their wider significance. In order to evaluate existing data and identify future research directions, this paper provides a critical overview of the component elements of the NAMSA and shows that the timing of many key behaviors has close parallels with others in sub-Saharan Africa and Southwest Asia., (© 2017 Wiley Periodicals, Inc.)