The genetic legacy of the expansion of Bantu-speaking peoples in Africa.

The expansion of people speaking Bantu languages is the most dramatic demographic event in Late Holocene Africa and fundamentally reshaped the linguistic, cultural and biological landscape of the continent1-7. With a comprehensive genomic dataset, including newly generated data of modern-day and ancient DNA from previously unsampled regions in Africa, we contribute insights into this expansion that started 6,000-4,000 years ago in western Africa. We genotyped 1,763 participants, including 1,526 Bantu speakers from 147 populations across 14 African countries, and generated whole-genome sequences from 12 Late Iron Age individuals8. We show that genetic diversity amongst Bantu-speaking populations declines with distance from western Africa, with current-day Zambia and the Democratic Republic of Congo as possible crossroads of interaction. Using spatially explicit methods9 and correlating genetic, linguistic and geographical data, we provide cross-disciplinary support for a serial-founder migration model. We further show that Bantu speakers received significant gene flow from local groups in regions they expanded into. Our genetic dataset provides an exhaustive modern-day African comparative dataset for ancient DNA studies10 and will be important to a wide range of disciplines from science and humanities, as well as to the medical sector studying human genetic variation and health in African and African-descendant populations.

New investigations at Kalambo Falls, Zambia: Luminescence chronology, site formation, and archaeological significance.

Fluvial deposits can provide excellent archives of early hominin activity but may be complex to interpret, especially without extensive geochronology. The Stone Age site of Kalambo Falls, northern Zambia, has yielded a rich artefact record from dominantly fluvial deposits, but its significance has been restricted by uncertainties over site formation processes and a limited chronology. Our new investigations in the centre of the Kalambo Basin have used luminescence to provide a chronology and have provided key insights into the geomorphological and sedimentological processes involved in site formation. Excavations reveal a complex assemblage of channel and floodplain deposits. Single grain quartz optically stimulated luminescence (OSL) measurements provide the most accurate age estimates for the youngest sediments, but in older deposits the OSL signal from some grains is saturated. A different luminescence signal from quartz, thermally transferred OSL (TT-OSL), can date these older deposits. OSL and TT-OSL results are combined to provide a chronology for the site. Ages indicate four phases of punctuated deposition by the dominantly laterally migrating and vertically aggrading Kalambo River (∼500-300 ka, ∼300-50 ka, ∼50-30 ka, ∼1.5-0.49 ka), followed by deep incision and renewed lateral migration at a lower topographic level. A conceptual model for site formation provides the basis for improved interpretation of the generation, preservation, and visibility of the Kalambo archaeological record. This model highlights the important role of intrinsic meander dynamics in site formation and does not necessarily require complex interpretations that invoke periodic blocking of the Kalambo River, as has previously been suggested. The oldest luminescence ages place the Mode 2/3 transition between ∼500 and 300 ka, consistent with other African and Asian sites where a similar transition can be found. The study approach adopted here can potentially be applied to other fluvial Stone Age sites throughout Africa and beyond.

BaTwa populations from Zambia retain ancestry of past hunter-gatherer groups.

Sub-equatorial Africa is today inhabited predominantly by Bantu-speaking groups of Western African descent who brought agriculture to the Luangwa valley in eastern Zambia ~2000 years ago. Before their arrival the area was inhabited by hunter-gatherers, who in many cases were subsequently replaced, displaced or assimilated. In Zambia, we know little about the genetic affinities of these hunter-gatherers. We examine ancestry of two isolated communities in Zambia, known as BaTwa and possible descendants of recent hunter-gatherers. We genotype over two million genome-wide SNPs from two BaTwa populations (total of 80 individuals) and from three comparative farming populations to: (i) determine if the BaTwa carry genetic links to past hunter-gatherer-groups, and (ii) characterise the genetic affinities of past Zambian hunter-gatherer-groups. The BaTwa populations do harbour a hunter-gatherer-like genetic ancestry and Western African ancestry. The hunter-gatherer component is a unique local signature, intermediate between current-day Khoe-San ancestry from southern Africa and central African rainforest hunter-gatherer ancestry.

The dating and interpretation of a Mode 1 site in the Luangwa Valley, Zambia.

Flake based assemblages (Mode 1) comprise the earliest stone technologies known, with well-dated Oldowan sites occurring in eastern Africa between ~2.6-1.7 Ma, and in less securely dated contexts in central, southern and northern Africa. Our understanding of the spread and local development of this technology outside East Africa remains hampered by the lack of reliable numerical dating techniques applicable to non-volcanic deposits. This study applied the still relatively new technique of cosmogenic nuclide burial dating ((10)Be/(26)Al) to calculate burial ages for fluvial gravels containing Mode 1 artefacts in the Luangwa Valley, Zambia. The Manzi River, a tributary of the Luangwa River, has exposed a 4.7 m deep section of fluvial sands with discontinuous but stratified gravel layers bearing Mode 1, possibly Oldowan, artefacts in the basal layers. An unconformity divides the Manzi section, separating Mode 1 deposits from overlying gravels containing Mode 3 (Middle Stone Age) artefacts. No diagnostic Mode 2 (Acheulean) artefacts were found. Cosmogenic nuclide burial dating was attempted for the basal gravels as well as exposure ages for the upper Mode 3 gravels, but was unsuccessful. The complex depositional history of the site prevented the calculation of reliable age models. A relative chronology for the full Manzi sequence was constructed, however, from the magnetostratigraphy of the deposit (N>R>N sequence). Isothermal thermoluminescence (ITL) dating of the upper Mode 3 layers also provided consistent results (~78 ka). A coarse but chronologically coherent sequence now exists for the Manzi section with the unconformity separating probable mid- or early Pleistocene deposits below from late Pleistocene deposits above. The results suggest Mode 1 technology in the Luangwa Valley may post-date the Oldowan in eastern and southern Africa. The dating programme has contributed to a clearer understanding of the geomorphological processes that have shaped the valley and structured its archaeological record.

Genetic perspectives on forager-farmer interaction in the Luangwa valley of Zambia.

The transformation from a foraging way of life to a reliance on domesticated plants and animals often led to the expansion of agropastoralist populations at the expense of hunter-gatherers (HGs). In Africa, one of these expansions involved the Niger-Congo Bantu-speaking populations that started to spread southwards from Cameroon/Nigeria approximately 4,000 years ago, bringing agricultural technologies. Genetic studies have shown different degrees of gene flow (sometimes involving sex-biased migrations) between Bantu agriculturalists and HGs. Although these studies have covered many parts of sub-Saharan Africa, the central part (e.g. Zambia) was not yet studied, and the interactions between immigrating food-producers and local HGs are still unclear. Archeological evidence from the Luangwa Valley of Zambia suggests a long period of coexistence ( approximately 1,700 years) of early food-producers and HGs. To investigate if this apparent coexistence was accompanied by genetic admixture, we analyzed the mtDNA control region, Y chromosomal unique event polymorphisms, and 12 associated Y- short tandem repeats in two food-producing groups (Bisa and Kunda) that live today in the Luangwa Valley, and compared these data with available published data on African HGs. Our results suggest that both the Bisa and Kunda experienced at most low levels of admixture with HGs, and these levels do not differ between the maternal and paternal lineages. Coalescent simulations indicate that the genetic data best fit a demographic scenario with a long divergence (62,500 years) and little or no gene flow between the ancestors of the Bisa/Kunda and existing HGs. This scenario contrasts with the archaeological evidence for a long period of coexistence between the two different communities in the Luangwa Valley, and suggests a process of sociocultural boundary maintenance may have characterized their interaction.

A Dynamical Analysis of the Suitability of Prehistoric Spheroids from the Cave of Hearths as Thrown Projectiles.

Spheroids are ball-shaped stone objects found in African archaeological sites dating from 1.8 million years ago (Early Stone Age) to at least 70,000 years ago (Middle Stone Age). Spheroids are either fabricated or naturally shaped stones selected and transported to places of use making them one of the longest-used technologies on record. Most hypotheses about their use suggest they were percussive tools for shaping or grinding other materials. However, their size and spherical shape make them potentially useful as projectile weapons, a property that, uniquely, humans have been specialised to exploit for millions of years. Here we show (using simulations of projectile motions resulting from human throwing) that 81% of a sample of spheroids from the late Acheulean (Bed 3) at the Cave of Hearths, South Africa afford being thrown so as to inflict worthwhile damage to a medium-sized animal over distances up to 25 m. Most of the objects have weights that produce optimal levels of damage from throwing, rather than simply being as heavy as possible (as would suit other functions). Our results show that these objects were eminently suitable for throwing, and demonstrate how empirical research on behavioural tasks can inform and constrain our theories about prehistoric artefacts.

Clarifying some fundamental errors in herries' "a chronological perspective on the acheulian and its transition to the middle stone age in southern Africa: the question of the fauresmith" (2011).

Herries provides a timely review of the archaeological and dating evidence of the transition from the Acheulean to the Middle Stone Age (MSA) in southern Africa, however, in relation to the site of Twin Rivers, Zambia he makes several fundamental errors of interpretation that demand correction. The stratigraphic sequence of the site is admittedly complex, but it deserves a more careful analysis than that offered by Herries. This detailed response by the most recent excavator of the site addresses Herries critique by placing the site in its historical context and then dealing with the central issue of the association of dated speleothem with the surviving archaeological deposits. Herries is shown to have mistakenly combined the dates from two separate cave passages and to have misunderstood the published sections, plans, and taphonomic assessment of each excavation area. His reinterpretation of the site as being significantly younger than published is based on a conflation of unrelated data.

Backed tools in Middle Pleistocene central Africa and their evolutionary significance.

The fashioning of stone inserts for composite tools by blunting flakes and blades is a technique usually associated with Late Pleistocene modern humans. Recent reports from two sites in south central Africa (Twin Rivers and Kalambo Falls) suggest that this backed tool technology originated in the later Middle Pleistocene with early or "archaic" Homo sapiens. This paper investigates these claims critically from the perspective of the potential mixing of Middle and Later Stone Age deposits at the two sites and the possible creation of misleading assemblages. The review shows that backed tools form a statistically minor, but technologically significant feature of the early Middle Stone Age of south central Africa. They first appear in the Lupemban industry at approximately 300 ka and remain an element of the Middle Stone Age technological repertoire of the region. Comparisons are made with early backed tool assemblages of east Africa and with the much younger Howiesons Poort industry of southern Africa. The paper concludes that Lupemban tools lack the standardization of the Howiesons Poort backed pieces, but form part of a regionally distinctive and diverse assemblage of heavy and light duty tools. Some modern-like behaviours appear to have emerged by the later Middle Pleistocene in south central Africa.