First evidence for human occupation of a lava tube in Arabia: The archaeology of Umm Jirsan Cave and its surroundings, northern Saudi Arabia.

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.

Longstanding behavioural stability in West Africa extends to the Middle Pleistocene at Bargny, coastal Senegal.

Middle Stone Age (MSA) technologies first appear in the archaeological records of northern, eastern and southern Africa during the Middle Pleistocene epoch. The absence of MSA sites from West Africa limits evaluation of shared behaviours across the continent during the late Middle Pleistocene and the diversity of subsequent regionalized trajectories. Here we present evidence for the late Middle Pleistocene MSA occupation of the West African littoral at Bargny, Senegal, dating to 150 thousand years ago. Palaeoecological evidence suggests that Bargny was a hydrological refugium during the MSA occupation, supporting estuarine conditions during Middle Pleistocene arid phases. The stone tool technology at Bargny presents characteristics widely shared across Africa in the late Middle Pleistocene but which remain uniquely stable in West Africa to the onset of the Holocene. We explore how the persistent habitability of West African environments, including mangroves, contributes to distinctly West African trajectories of behavioural stability.

Human interactions with tropical environments over the last 14,000 years at Iho Eleru, Nigeria.

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.

Homo sapiens lithic technology and microlithization in the South Asian rainforest at Kitulgala Beli-lena (c. 45 - 8,000 years ago).

Recent archaeological investigations in Sri Lanka have reported evidence for the exploitation and settlement of tropical rainforests by Homo sapiens since c. 48,000 BP. Information on technological approaches used by human populations in rainforest habitats is restricted to two cave sites, Batadomba-lena and Fa-Hien Lena. Here, we provide detailed study of the lithic assemblages of Kitulgala Beli-lena, a recently excavated rockshelter preserving a sedimentary sequence from the Late Pleistocene to the Holocene. Our analysis indicates in situ lithic production and the recurrent use of the bipolar method for the production of microliths. Stone tool analyses demonstrate long-term technological stability from c. 45,000 to 8,000 years BP, a pattern documented in other rainforest locations. Foraging behaviour is characterised by the use of lithic bipolar by-products together with osseous projectile points for the consistent targeting of semi-arboreal/arboreal species, allowing for the widespread and recurrent settlement of the wet zone of Sri Lanka.

A spatiotemporally explicit paleoenvironmental framework for the Middle Stone Age of eastern Africa.

Eastern Africa has played a prominent role in debates about human evolution and dispersal due to the presence of rich archaeological, palaeoanthropological and palaeoenvironmental records. However, substantial disconnects occur between the spatial and temporal resolutions of these data that complicate their integration. Here, we apply high-resolution climatic simulations of two key parameters, mean annual temperature and precipitation, and a biome model, to produce a highly refined characterisation of the environments inhabited during the eastern African Middle Stone Age. Occupations are typically found in sub-humid climates and landscapes dominated by or including tropical xerophytic shrubland. Marked expansions from these core landscapes include movement into hotter, low-altitude landscapes in Marine Isotope Stage 5 and cooler, high-altitude landscapes in Marine Isotope Stage 3, with the recurrent inhabitation of ecotones between open and forested habitats. Through our use of high-resolution climate models, we demonstrate a significant independent relationship between past precipitation and patterns of Middle Stone Age stone tool production modes overlooked by previous studies. Engagement with these models not only enables spatiotemporally explicit examination of climatic variability across Middle Stone Age occupations in eastern Africa but enables clearer characterisation of the habitats early human populations were adapted to, and how they changed through time.

Evaluating refugia in recent human evolution in Africa.

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'.

Constraining the chronology and ecology of Late Acheulean and Middle Palaeolithic occupations at the margins of the monsoon.

South Asia hosts the world's youngest Acheulean sites, with dated records typically restricted to sub-humid landscapes. The Thar Desert marks a major adaptive boundary between monsoonal Asia to the east and the Saharo-Arabian desert belt to the west, making it a key threshold to examine patterns of hominin ecological adaptation and its impacts on patterns of behaviour, demography and dispersal. Here, we investigate Palaeolithic occupations at the western margin of the South Asian monsoon at Singi Talav, undertaking new chronometric, sedimentological and palaeoecological studies of Acheulean and Middle Palaeolithic occupation horizons. We constrain occupations of the site between 248 and 65 thousand years ago. This presents the first direct palaeoecological evidence for landscapes occupied by South Asian Acheulean-producing populations, most notably in the main occupation horizon dating to 177 thousand years ago. Our results illustrate the potential role of the Thar Desert as an ecological, and demographic, frontier to Palaeolithic populations.

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.

Directional changes in Levallois core technologies between Eastern Africa, Arabia, and the Levant during MIS 5.

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.

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.

Nubian Levallois technology associated with southernmost Neanderthals.

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.

Continuity of the Middle Stone Age into the Holocene.

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.

Neural networks differentiate between Middle and Later Stone Age lithic assemblages in eastern Africa.

The Middle to Later Stone Age transition marks a major change in how Late Pleistocene African populations produced and used stone tool kits, but is manifest in various ways, places and times across the continent. Alongside changing patterns of raw material use and decreasing artefact sizes, changes in artefact types are commonly employed to differentiate Middle Stone Age (MSA) and Later Stone Age (LSA) assemblages. The current paper employs a quantitative analytical framework based upon the use of neural networks to examine changing constellations of technologies between MSA and LSA assemblages from eastern Africa. Network ensembles were trained to differentiate LSA assemblages from Marine Isotope Stage 3&4 MSA and Marine Isotope Stage 5 MSA assemblages based upon the presence or absence of 16 technologies. Simulations were used to extract significant indicator and contra-indicator technologies for each assemblage class. The trained network ensembles classified over 94% of assemblages correctly, and identified 7 key technologies that significantly distinguish between assemblage classes. These results clarify both temporal changes within the MSA and differences between MSA and LSA assemblages in eastern Africa.

Microliths in the South Asian rainforest ~45-4 ka: New insights from Fa-Hien Lena Cave, Sri Lanka.

Microliths-small, retouched, often-backed stone tools-are often interpreted to be the product of composite tools, including projectile weapons, and efficient hunting strategies by modern humans. In Europe and Africa these lithic toolkits are linked to hunting of medium- and large-sized game found in grassland or woodland settings, or as adaptations to risky environments during periods of climatic change. Here, we report on a recently excavated lithic assemblage from the Late Pleistocene cave site of Fa-Hien Lena in the tropical evergreen rainforest of Sri Lanka. Our analyses demonstrate that Fa-Hien Lena represents the earliest microlith assemblage in South Asia (c. 48,000-45,000 cal. years BP) in firm association with evidence for the procurement of small to medium size arboreal prey and rainforest plants. Moreover, our data highlight that the lithic technology of Fa-Hien Lena changed little over the long span of human occupation (c. 48,000-45,000 cal. years BP to c. 4,000 cal. years BP) indicating a successful, stable technological adaptation to the tropics. We argue that microlith assemblages were an important part of the environmental plasticity that enabled Homo sapiens to colonise and specialise in a diversity of ecological settings during its expansion within and beyond Africa. The proliferation of diverse microlithic technologies across Eurasia c. 48-45 ka was part of a flexible human 'toolkit' that assisted our species' spread into all of the world's environments, and the development of specialised technological and cultural approaches to novel ecological situations.

Specialized rainforest hunting by Homo sapiens ~45,000 years ago.

Defining the distinctive capacities of Homo sapiens relative to other hominins is a major focus for human evolutionary studies. It has been argued that the procurement of small, difficult-to-catch, agile prey is a hallmark of complex behavior unique to our species; however, most research in this regard has been limited to the last 20,000 years in Europe and the Levant. Here, we present detailed faunal assemblage and taphonomic data from Fa-Hien Lena Cave in Sri Lanka that demonstrates specialized, sophisticated hunting of semi-arboreal and arboreal monkey and squirrel populations from ca. 45,000 years ago, in a tropical rainforest environment. Facilitated by complex osseous and microlithic technologies, we argue these data highlight that the early capture of small, elusive mammals was part of the plastic behavior of Homo sapiens that allowed it to rapidly colonize a series of extreme environments that were apparently untouched by its hominin relatives.

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.