Human origins in Southern African palaeo-wetlands? Strong claims from weak evidence

Attempts to identify a ‘homeland’ for our species from genetic data are widespread in the academic literature. However, even when putting aside the question of whether a ‘homeland’ is a useful concept, there are a number of inferential pitfalls in attempting to identify the geographic origin of a species from contemporary patterns of genetic variation. These include making strong claims from weakly informative data, treating genetic lineages as representative of populations, assuming a high degree of regional population continuity over hundreds of thousands of years, and using circumstantial observations as corroborating evidence without considering alternative hypotheses on an equal footing, or formally evaluating any hypothesis. In this commentary we review the recent publication that claims to pinpoint the origins of ‘modern humans’ to a very specific region in Africa (Chan et al., 2019), demonstrate how it fell into these inferential pitfalls, and discuss how this can be avoided

Rock art imagery as a proxy for Holocene environmental change: A view from Shuwaymis, NW Saudi Arabia

The animal species depicted in the rock art of Shuwaymis, Saudi Arabia, provide a record of Holocene climatic changes, as seen by the engravers. Of 1903 animal engravings, 1514 contained sufficient detail to allow identification with confidence. In addition, the stratigraphy of the engravings and the depiction of domesticates provide a broad chronological framework that allows a division into images created during the Holocene humid phase and animals represented after the onset of desert conditions. Despite the large sample size, only 16 animal species could be identified, which represents an extraordinarily narrow species spectrum. Comparison with the scarce faunal record of the Arabian Peninsula shows that all larger animals that are thought to have been present in the area were also depicted in the rock art. The contemporaneous presence of at least four large carnivores during the Holocene humid phase suggests that prey animals were abundant, and that the landscape consisted of a mosaic of habitats, potentially with thicker vegetation along the water courses of the wadis and more open vegetation in the landscape around them. Community Earth System Models (COSMOS) climate simulations show that Shuwaymis was at the northern edge of the African Summer Monsoon rainfall regime. It is therefore possible that Shuwaymis was ecologically connected with southwestern Arabia, and that an arid barrier remained in place to the north, restricting the dispersal of Levantine species into Arabia

Middle Pleistocene vertebrate fossils from the Nefud Desert, Saudi Arabia: Implications for biogeography and palaeoecology

The current paucity of Pleistocene vertebrate records from the Arabian Peninsula - a landmass of over 3 million km2 - is a significant gap in our knowledge of the Quaternary. Such data are critical lines of contextual evidence for considering animal and hominin dispersals between Africa and Eurasia generally, and hominin palaeoecology in the Pleistocene landscapes of the Arabian interior specifically. Here, we describe an important contribution to the record and report stratigraphically-constrained fossils of mammals, birds and reptiles from recent excavations at Ti’s al Ghadah in the southwestern Nefud Desert. Combined U-series and ESR analyses of Oryx sp. teeth indicate that the assemblage is Middle Pleistocene in age and dates to ca. 500 ka. The identified fauna is a biogeographical admixture that consists of likely endemics and taxa of African and Eurasian affinity and includes extinct and extant (or related Pleistocene forms of) mammals (Palaeoloxodon cf. recki, Panthera cf. gombaszogenis, Equus hemionus, cf. Crocuta crocuta, Vulpes sp., Canis anthus, Oryx sp.), the first Pleistocene records of birds from the Arabian Peninsula (Struthio sp., Neophron percnopterus, Milvus cf. migrans, Tachybaptus sp. Anas sp., Pterocles orientalis, Motacilla cf. alba) and reptiles (Varanidae/Uromastyx sp.). We infer that the assemblage reflects mortality in populations of herbivorous animals and their predators and scavengers that were attracted to freshwater and plant resources in the inter-dune basin. At present, there is no evidence to suggest hominin agency in the accumulation of the bone assemblages. The inferred ecological characteristics of the taxa recovered indicate the presence, at least periodically, of substantial water-bodies and open grassland habitats

Middle Pleistocene vertebrate fossils from the Nefud Desert, Saudi Arabia: Implications for biogeography and palaeoecology

The current paucity of Pleistocene vertebrate records from the Arabian Peninsula – a landmass of over 3 million km2 – is a significant gap in our knowledge of the Quaternary. Such data are critical lines of contextual evidence for considering animal and hominin dispersals between Africa and Eurasia generally, and hominin palaeoecology in the Pleistocene landscapes of the Arabian interior specifically. Here, we describe an important contribution to the record and report stratigraphically-constrained fossils of mammals, birds and reptiles from recent excavations at Ti’s al Ghadah in the southwestern Nefud Desert. Combined U-series and ESR analyses of Oryx sp. teeth indicate that the assemblage is Middle Pleistocene in age and dates to ca. 500 ka. The identified fauna is a biogeographical admixture that consists of likely endemics and taxa of African and Eurasian affinity and includes extinct and extant (or related Pleistocene forms of) mammals (Palaeoloxodon cf. recki, Panthera cf. gombaszogenis, Equus hemionus, cf. Crocuta crocuta, Vulpes sp., Canis anthus, Oryx sp.), the first Pleistocene records of birds from the Arabian Peninsula (Struthio sp., Neophron percnopterus, Milvus cf. migrans, Tachybaptus sp. Anas sp., Pterocles orientalis, Motacilla cf. alba) and reptiles (Varanidae/Uromastyx sp.). We infer that the assemblage reflects mortality in populations of herbivorous animals and their predators and scavengers that were attracted to freshwater and plant resources in the inter-dune basin. At present, there is no evidence to suggest hominin agency in the accumulation of the bone assemblages. The inferred ecological characteristics of the taxa recovered indicate the presence, at least periodically, of substantial water-bodies and open grassland habitats

Coring, profiling, and trenching: Archaeological field strategies for investigating the Pleistocene-Holocene-Anthropocene continuum

Archaeologists have long emphasized the importance of large-scale excavations and multi-year or even decades-long projects at a single site or site complex. Here, we highlight archaeological field strategies, termed coring, profiling, and trenching (CPT), that rely on relatively small-scale excavations or the collection of new samples from intact deposits in previously excavated trenches (aka test units or pits). Examples from multiple sites in Africa, Asia, and North America demonstrate that CPT is highly effective for obtaining high-resolution archaeobiological and geoarchaeological samples (e.g., faunal and botanical remains, sediments) and artefacts from areas that have seen limited or no archaeological research, little systematic application of archaeological science methods, or research only on a relatively narrow time period or geographic scale. Designed to complement large-scale excavations at single sites, CPT is ideal for multi-scalar research that works in tandem with remote sensing techniques, providing samples for detailed laboratory analyses and offering a bridge between surface surveys and large-scale excavation. Given the threats facing archaeological sites around the world from climate change and human development, as well as financial, training and infrastructure constraints, and concerns from many Indigenous communities about large excavations, we argue that CPT is an important method for addressing 21st century human-environmental research questions

Mapping human dispersals into the Horn of Africa from Arabian Ice Age refugia using mitogenomes

Rare mitochondrial lineages with relict distributions can sometimes be disproportionately informative about deep events in human prehistory. We have studied one such lineage, haplogroup R0a, which uniquely is most frequent in Arabia and the Horn of Africa, but is distributed much more widely, from Europe to India. We conclude that: (1) the lineage ancestral to R0a is more ancient than previously thought, with a relict distribution across the Mediterranean/Southwest Asia; (2) R0a has a much deeper presence in Arabia than previously thought, highlighting the role of at least one Pleistocene glacial refugium, perhaps on the Red Sea plains; (3) the main episode of dispersal into Eastern Africa, at least concerning maternal lineages, was at the end of the Late Glacial, due to major expansions from one or more refugia in Arabia; (4) there was likely a minor Late Glacial/early postglacial dispersal from Arabia through the Levant and into Europe, possibly alongside other lineages from a Levantine refugium; and (5) the presence of R0a in Southwest Arabia in the Holocene at the nexus of a trading network that developed after ~3 ka between Africa and the Indian Ocean led to some gene flow even further afield, into Iran, Pakistan and India

Homo sapiens in Arabia by 85,000 years ago

Understanding the timing and character of Homo sapiens expansion out of Africa is critical for inferring the colonisation 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 only reached 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

Fossil herbivore stable isotopes reveal middle Pleistocene hominin palaeoenvironment in ‘Green Arabia’

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 c. 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 lithics and probable cutmarks. The results demonstrate productive grasslands in the interior of the Arabian Peninsula c. 300-500 ka, as well as aridity levels similar to those found in open savanna 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 dispersal into the interior of the Arabian Peninsula required no major novel adaptation

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

Genomic analyses inform on migration events during the peopling of Eurasia.

High-coverage whole-genome sequence studies have so far focused on a limited number of geographically restricted populations, or been targeted at specific diseases, such as cancer. Nevertheless, the availability of high-resolution genomic data has led to the development of new methodologies for inferring population history and refuelled the debate on the mutation rate in humans. Here we present the Estonian Biocentre Human Genome Diversity Panel (EGDP), a dataset of 483 high-coverage human genomes from 148 populations worldwide, including 379 new genomes from 125 populations, which we group into diversity and selection sets. We analyse this dataset to refine estimates of continent-wide patterns of heterozygosity, long- and short-distance gene flow, archaic admixture, and changes in effective population size through time as well as for signals of positive or balancing selection. We find a genetic signature in present-day Papuans that suggests that at least 2% of their genome originates from an early and largely extinct expansion of anatomically modern humans (AMHs) out of Africa. Together with evidence from the western Asian fossil record, and admixture between AMHs and Neanderthals predating the main Eurasian expansion, our results contribute to the mounting evidence for the presence of AMHs out of Africa earlier than 75,000 years ago.Support was provided by: Estonian Research Infrastructure Roadmap grant no 3.2.0304.11-0312; Australian Research Council Discovery grants (DP110102635 and DP140101405) (D.M.L., M.W. and E.W.); Danish National Research Foundation; the Lundbeck Foundation and KU2016 (E.W.); ERC Starting Investigator grant (FP7 - 261213) (T.K.); Estonian Research Council grant PUT766 (G.C. and M.K.); EU European Regional Development Fund through the Centre of Excellence in Genomics to Estonian Biocentre (R.V.; M.Me. and A.Me.), and Centre of Excellence for Genomics and Translational Medicine Project No. 2014-2020.4.01.15-0012 to EGC of UT (A.Me.) and EBC (M.Me.); Estonian Institutional Research grant IUT24-1 (L.S., M.J., A.K., B.Y., K.T., C.B.M., Le.S., H.Sa., S.L., D.M.B., E.M., R.V., G.H., M.K., G.C., T.K. and M.Me.) and IUT20-60 (A.Me.); French Ministry of Foreign and European Affairs and French ANR grant number ANR-14-CE31-0013-01 (F.-X.R.); Gates Cambridge Trust Funding (E.J.); ICG SB RAS (No. VI.58.1.1) (D.V.L.); Leverhulme Programme grant no. RP2011-R-045 (A.B.M., P.G. and M.G.T.); Ministry of Education and Science of Russia; Project 6.656.2014/K (S.A.F.); NEFREX grant funded by the European Union (People Marie Curie Actions; International Research Staff Exchange Scheme; call FP7-PEOPLE-2012-IRSES-number 318979) (M.Me., G.H. and M.K.); NIH grants 5DP1ES022577 05, 1R01DK104339-01, and 1R01GM113657-01 (S.Tis.); Russian Foundation for Basic Research (grant N 14-06-00180a) (M.G.); Russian Foundation for Basic Research; grant 16-04-00890 (O.B. and E.B); Russian Science Foundation grant 14-14-00827 (O.B.); The Russian Foundation for Basic Research (14-04-00725-a), The Russian Humanitarian Scientific Foundation (13-11-02014) and the Program of the Basic Research of the RAS Presidium “Biological diversity” (E.K.K.); Wellcome Trust and Royal Society grant WT104125AIA & the Bristol Advanced Computing Research Centre (http://www.bris.ac.uk/acrc/) (D.J.L.); Wellcome Trust grant 098051 (Q.A.; C.T.-S. and Y.X.); Wellcome Trust Senior Research Fellowship grant 100719/Z/12/Z (M.G.T.); Young Explorers Grant from the National Geographic Society (8900-11) (C.A.E.); ERC Consolidator Grant 647787 ‘LocalAdaptatio’ (A.Ma.); Program of the RAS Presidium “Basic research for the development of the Russian Arctic” (B.M.); Russian Foundation for Basic Research grant 16-06-00303 (E.B.); a Rutherford Fellowship (RDF-10-MAU-001) from the Royal Society of New Zealand (M.P.C.)