Khoi-San peoples of southern Africa: The Khoekhoen people have an “Indigenous nomadic pastoralist culture,” and the San people have an “Indigenous hunter-gatherer culture,” which is one of the oldest surviving cultures of the region

Damien Marie AtHope’s Art

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Khoi-san and Hadza peoples of southern Africa: The Khoekhoen people have an “Indigenous nomadic pastoralist culture,” and the San people have an “Indigenous hunter-gatherer culture,” which is one of the oldest surviving cultures of the region. The Hadza people, who have an “indigenous hunter-gatherer culture,” were a pre-Bantu expansion culture not closely related to Khoisan speakers.

Khoisan

“Khoisan /ˈkɔɪsɑːn/ KOY-sahn, or Khoe-Sān (pronounced [kxʰoesaːn]), is a catch-all term for the indigenous peoples of Southern Africa who traditionally speak non-Bantu languages, combining the Khoekhoen (formerly “Hottentots“) and the Sān peoples (also called “Bushmen”). Khoisan populations traditionally speak click languages and are considered to be the historical communities throughout Southern Africa, remaining predominant until European colonisation in areas climatically unfavorable to Bantu (sorghum-based) agriculture, such as the Cape region, through to Namibia, where Khoekhoe populations of Nama and Damara people are prevalent groups, and Botswana. Considerable mingling with Bantu-speaking groups is evidenced by prevalence of click phonemes in many especially Xhosa Southern African Bantu languages.

“Many Khoesān peoples are the descendants of a very early dispersal of anatomically modern humans to Southern Africa before 150,000 years ago. (However, see below for recent work supporting a multi-regional hypothesis that suggests the Khoisan may be a source population for anatomically modern humans.) Their languages show a vague typological similarity, largely confined to the prevalence of click consonants. They are not verifiably derived from a common proto-language, but are today split into at least three separate and unrelated language families (Khoe-Kwadi, Tuu and Kxʼa). It has been suggested that the Khoekhoe may represent Late Stone Age arrivals to Southern Africa, possibly displaced by Bantu expansion reaching the area roughly between 1,500 and 2,000 years ago.” ref  

“Sān are popularly thought of as foragers in the Kalahari Desert and regions of Botswana, Namibia, Angola, Zambia, Zimbabwe, Lesotho and Northern South Africa. The word sān is from the Khoekhoe language and refers to foragers (“those who pick things up from the ground”) who do not own livestock. As such, it was used in reference to all hunter-gatherer populations who came into contact with Khoekhoe-speaking communities, and was largely referring to the lifestyle, distinct from a pastoralist or agriculturalist one, and not to any particular ethnicity. While there are attendant cosmologies and languages associated with this way of life, the term is an economic designator rather than a cultural or ethnic one.” ref

“It is suggested that the ancestors of the modern Khoisan expanded to southern Africa (from East or Central Africa) before 150,000 years ago, possibly as early as before 260,000 years ago, so that by the beginning of the MIS 5 “megadrought” 130,000 years ago, there were two ancestral population clusters in Africa, bearers of mt-DNA haplogroup L0 in southern Africa ancestral to the Khoi-San, and bearers of haplogroup L1-6 in central/eastern Africa ancestral to everyone else. This group gave rise to the San population of hunter gatherers. A much later wave of migration, around or before the beginning of the Common Era, gave rise to the Khoe people, who were pastoralists. This group carried DNA from Eurasian as well as some Neanderthal groups.” ref

“Due to their early expansion and separation, the populations ancestral to the Khoisan have been estimated as having represented the “largest human population” during the majority of the anatomically modern human timeline, from their early separation before 150 kya until the recent peopling of Eurasia some 70,000 years ago. They were much more widespread than today, their modern distribution being due to their decimation in the course of the Bantu expansion. They were dispersed throughout much of southern and southeastern Africa. There was also a significant back-migration of bearers of L0 towards eastern Africa between 120,000 and 75,000 years ago.” ref  

“Rito et al. (2013) speculate that pressure from such back-migration may even have contributed to the dispersal of East African populations out of Africa at about 70,000 years ago. Recent work has suggested that the multi-regional hypothesis may be supported by current human population genetic data. A 2023 study published in the journal Nature suggests that current genetic data may be best understood as reflecting internal admixtures of multiple population sources across Africa, including ancestral populations of the Khoisan.” ref

“The San populations ancestral to the Khoisan were spread throughout much of southern and eastern Africa throughout the Late Stone Age after about 75,000 years ago. A further expansion dated to about 20,000 years ago has been proposed based on the distribution of the L0d haplogroup. Rosti et al. suggest a connection of this recent expansion with the spread of click consonants to eastern African languages (Hadza language).” ref  

“The Late Stone Age Sangoan industry occupied southern Africa in areas where annual rainfall is less than a metre (1000 mm; 39.4 in). The contemporary San and Khoi peoples resemble those represented by the ancient Sangoan skeletal remains. Against the traditional interpretation that finds a common origin for the Khoi and San, other evidence has suggested that the ancestors of the Khoi peoples are relatively recent pre-Bantu agricultural immigrants to southern Africa who abandoned agriculture as the climate dried and either joined the San as hunter-gatherers or retained pastoralism.” ref

“With the hypothesized arrival of pastoralists & bantoid agro-pastoralists in southern Africa starting around 2,300 years ago, linguistic development is later seen in the click consonants and loan words from ancient Khoe-san languages into the evolution of blended agro-pastoralist & hunter-gatherer communities that would eventually evolve into the now extant, amalgamated modern native linguistic communities found in South Africa, Botswana & Namibia (e.g. in South African Xhosa, Sotho, Tswana, Zulu people.) The Khoikhoi enter the historical record with their first contact with Portuguese explorers, about 1,000 years after their displacement by the Bantu.” ref 

“In 2019, scientists from the University of the Free State discovered 8,000-year-old carvings made by the Khoisan people. The carvings depicted a hippopotamus, horse, and antelope in the ‘Rain Snake’ Dyke of the Vredefort impact structure, which may have spiritual significance regarding the rain-making mythology of the Khoisan. In the 1990s, genomic studies of the world’s peoples found that the Y chromosome of San men share certain patterns of polymorphisms that are distinct from those of all other populations. Because the Y chromosome is highly conserved between generations, this type of DNA test is used to determine when different subgroups separated from one another, and hence their last common ancestry. The authors of these studies suggested that the San may have been one of the first populations to differentiate from the most recent common paternal ancestor of all extant humans.” ref

“Various Y-chromosome studies since confirmed that the Khoisan carry some of the most divergent (oldest) Y-chromosome haplogroups. These haplogroups are specific sub-groups of haplogroups A and B, the two earliest branches on the human Y-chromosome tree. Similar to findings from Y-chromosome studies, mitochondrial DNA studies also showed evidence that the Khoisan people carry high frequencies of the earliest haplogroup branches in the human mitochondrial DNA tree. The most divergent (oldest) mitochondrial haplogroup, L0d, has been identified at its highest frequencies in the southern African Khoi and San groups. The distinctiveness of the Khoisan in both matrilineal and patrilineal groupings is a further indicator that they represent a population historically distinct from other Africans.” ref

“Some genomic studies have further revealed that Khoisan groups have been influenced by 9 to 30% genetic admixture in the last few thousand years from an East African population who carried a Eurasian admixture component. Furthermore, they place an East African origin for the paternal haplogroup E1b1b found in these Southern African populations, as well as the introduction of pastoralism into the region. The paper also noted that the Bantu expansion had a notable genetic impact in a number of Khoisan groups. On the basis of PCA projections, the East African ancestry identified in the genomes of Khoe-Kwadi speakers and other southern Africans is related to an individual from the Tanzanian Luxmanda.” ref

Khoekhoe

“Khoekhoe (or Khoikhoi in former orthography) are the traditionally nomadic pastoralist indigenous population of South Africa. They are often grouped with the hunter-gatherer San (literally “Foragers”) peoples. The designation “Khoekhoe” is actually a kare or praise address, not an ethnic endonym, but it has been used in the literature as an ethnic term for Khoe-speaking peoples of Southern Africa, particularly pastoralist groups, such as the Griqua, Gona, Nama, Khoemana and Damara nations. The Khoekhoe were once known as Hottentots, a term now considered offensive.” ref

“While the presence of Khoekhoe in Southern Africa predates the Bantu expansion, according to a scientific theory based mainly on linguistic evidence, it is not clear when, possibly in the Late Stone Age, the Khoekhoe began inhabiting the areas where the first contact with Europeans occurred. At that time, in the 17th century, the Khoekhoe maintained large herds of Nguni cattle in the Cape region. Their Khoekhoe language is related to certain dialects spoken by foraging San peoples of the Kalahari, such as the Khwe and Tshwa, forming the Khoe language family. Khoekhoe subdivisions today are the Nama people of Namibia, Botswana and South Africa (with numerous clans), the Damara of Namibia, the Orana clans of South Africa (such as Nama or Ngqosini), the Khoemana or Griqua nation of South Africa, and the Gqunukhwebe or Gona clans which fall under the Xhosa-speaking polities.” ref

“The broad ethnic designation of “Khoekhoen”, meaning the peoples originally part of a pastoral culture and language group to be found across Southern Africa, is thought to refer to a population originating in the northern area of modern Botswana. This culture steadily spread southward, eventually reaching the Cape approximately 2,000 years ago. “Khoekhoe” groups include ǀAwakhoen to the west, and ǀKx’abakhoena of South and mid-South Africa, and the Eastern Cape. Both of these terms mean “Red People”, and are equivalent to the IsiXhosa term “amaqaba”. Husbandry of sheep, goats and cattle grazing in fertile valleys across the region provided a stable, balanced diet, and allowed these lifestyles to spread, with larger groups forming in a region previously occupied by the subsistence foragers.” ref

“Ntu-speaking agriculturalist culture is thought to have entered the region in the 3rd century AD, pushing pastoralists into the Western areas. The example of the close relation between the ǃUriǁ’aes (High clan), a cattle-keeping population, and the !Uriǁ’aeǀ’ona (High clan children), a more-or-less sedentary forager population (also known as “Strandlopers”), both occupying the area of ǁHuiǃgaeb, shows that the strict distinction between these two lifestyles is unwarranted, as well as the ethnic categories that are derived. Foraging peoples who ideologically value non-accumulation as a social value system would be distinct, however, but the distinctions among “Khoekhoe pastoralists”, “San hunter-gatherers” and “Bantu agriculturalists” do not hold up to scrutiny, and appear to be historical reductionism.” ref

The religious mythology of the Khoe-speaking cultures gives special significance to the Moon, which may have been viewed as the physical manifestation of a supreme being associated with heaven. Thiǁoab (Tsui’goab) is also believed to be the creator and the guardian of health, while ǁGaunab is primarily an evil being, who causes sickness or death. Many Khoe-speakers have converted to Christianity and Nama Muslims make up a large percentage of Namibia’s Muslims.” ref 

Nomadic pastoralism

“Nomadic pastoralism is a form of pastoralism in which livestock are herded in order to seek for fresh pastures on which to graze. True nomads follow an irregular pattern of movement, in contrast with transhumance, where seasonal pastures are fixed. However, this distinction is often not observed and the term ‘nomad’ used for both—and in historical cases the regularity of movements is often unknown in any case. The herded livestock include cattle, water buffalo, yaks, llamas, sheep, goats, reindeer, horses, donkeys or camels, or mixtures of species. Nomadic pastoralism is commonly practised in regions with little arable land, typically in the developing world, especially in the steppe lands north of the agricultural zone of Eurasia. Pastoralists often trade with sedentary agrarians, exchanging meat for grains, however have been known to raid.” ref

“Nomadic pastoralism was a result of the Neolithic Revolution and the rise of agriculture. During that revolution, humans began domesticating animals and plants for food and started forming cities. Nomadism generally has existed in symbiosis with such settled cultures trading animal products (meat, hides, wool, cheese and other animal products) for manufactured items not produced by the nomadic herders. Henri Fleisch tentatively suggested the Shepherd Neolithic industry of Lebanon may date to the Epipaleolithic and that it may have been used by one of the first cultures of nomadic shepherds in the Beqaa valley. Andrew Sherratt demonstrates that “early farming populations used livestock mainly for meat, and that other applications were explored as agriculturalists adapted to new conditions, especially in the semi‐arid zone.” ref

“In the past it was asserted that pastoral nomads left no presence archaeologically or were impoverished, but this has now been challenged, and was clearly not so for many ancient Eurasian nomads, who have left very rich kurgan burial sites. Pastoral nomadic sites are identified based on their location outside the zone of agriculture, the absence of grains or grain-processing equipment, limited and characteristic architecture, a predominance of sheep and goat bones, and by ethnographic analogy to modern pastoral nomadic peoples Juris Zarins has proposed that pastoral nomadism began as a cultural lifestyle in the wake of the 6200 BCE or around 8,200 years ago climatic crisis when Harifian pottery making hunter-gatherers in the Sinai fused with Pre-Pottery Neolithic B agriculturalists to produce the Munhata culture, a nomadic lifestyle based on animal domestication, developing into the Yarmoukian and thence into a circum-Arabian nomadic pastoral complex, and spreading Proto-Semitic languages.” ref

“In Bronze Age Central Asia, nomadic populations are associated with the earliest transmissions of millet and wheat grains through the region that eventually became central for the Silk Road. Early Indo-European migrations from the Pontic–Caspian steppe spread Yamnaya Steppe pastoralist ancestry and Indo-European languages across large parts of Eurasia. By the medieval period in Central Asia, nomadic communities exhibited isotopically diverse diets, suggesting a multitude of subsistence strategies. David Christian made the following observations about pastoralism. The agriculturist lives from domesticated plants and the pastoralist lives from domesticated animals.” ref

“Since animals are higher on the food chain, pastoralism supports a thinner population than agriculture. Pastoralism predominates where low rainfall makes farming impractical. Full pastoralism required the Secondary products revolution when animals began to be used for wool, milk, riding and traction as well as meat. Where grass is poor herds must be moved, which leads to nomadism. Some peoples are fully nomadic while others live in sheltered winter camps and lead their herds into the steppe in summer. Some nomads travel long distances, usually north in summer and south in winter. Near mountains, herds are led uphill in summer and downhill in winter (transhumance). Pastoralists often trade with or raid their agrarian neighbors.” ref

“David Christian distinguished ‘Inner Eurasia’, which was pastoral with a few hunter-gatherers in the far north, from ‘Outer Eurasia’, a crescent of agrarian civilizations from Europe through India to China. High civilization is based on agriculture where tax-paying peasants support landed aristocrats, kings, cities, literacy and scholars. Pastoral societies are less developed and as a result, according to Christian, more egalitarian. One tribe would often dominate its neighbors, but these ’empires’ usually broke up after a hundred years or so. The heartland of pastoralism is the Eurasian steppe. In the center of Eurasia pastoralism extended south to Iran and surrounded agrarian oasis cities. When pastoral and agrarian societies went to war, horse-borne mobility counterbalanced greater numbers.” ref

“Attempts by agrarian civilizations to conquer the steppe usually failed until the last few centuries. Pastoralists frequently raided and sometimes collected regular tribute from their farming neighbors. Especially in north China and Iran, they would sometimes conquer agricultural societies, but these dynasties were usually short-lived and broke up when the nomads became ‘civilized’ and lost their warlike virtues. Nomadic pastoralism was historically widespread throughout less fertile regions of Earth. It is found in areas of low rainfall such as the Arabian Peninsula inhabited by Bedouins, as well as Northeast Africa inhabited, among other ethnic groups, by Somalis (where camel, cattle, sheep and goat nomadic pastoralism is especially common).” ref

“Nomadic transhumance is also common in areas of harsh climate, such as Northern Europe and Russia inhabited by the indigenous Sami people, Nenets people and Chukchis. There are an estimated 30–40 million nomads in the world. Pastoral nomads and semi-nomadic pastoralists form a significant but declining minority in such countries as Saudi Arabia (probably less than 3%), Iran (4%), and Afghanistan (at most 10%). They comprise less than 2% of the population in the countries of North Africa except Libya and Mauritania.” ref

“The Eurasian steppe has been largely populated by pastoralist nomads since the late prehistoric times, with a succession of peoples known by the names given to them by surrounding literate sedentary societies, including the Bronze Age Proto-Indo-Europeans, and later Proto-Indo-Iranians, Scythians, Sarmatians, Cimmerians, Massagetae, Alans, Pechenegs, Cumans, Kipchaks, Karluks, Saka, Yuezhi, Wusun, Jie, Xiongnu, Xianbei, Khitan, Pannonian Avars, Huns, Mongols, Dzungars and various Turkics.” ref

“The Mongols in what is now Mongolia, Russia and China, and the Tatars or Turkic people of Eastern Europe and Central Asia were nomadic people who practiced nomadic transhumance on harsh Asian steppes. Some remnants of these populations are nomadic to this day. In Mongolia, about 40% of the population continues to live a traditional nomadic lifestyle. In China, it is estimated that a little over five million herders are dispersed over the pastoral counties, and more than 11 million over the semi-pastoral counties. This brings the total of the (semi)nomadic herder population to over 16 million, in general living in remote, scattered and resource-poor communities.” ref

“In Chad, nomadic pastoralists include the Zaghawa, Kreda, and Mimi. Farther north in Egypt and western Libya, the Bedouins also practice pastoralism. Sometimes nomadic pastoralists move their herds across international borders in search of new grazing terrain or for trade. This cross-border activity can occasionally lead to tensions with national governments as this activity is often informal and beyond their control and regulation. In East Africa, for example, over 95% of cross-border trade is through unofficial channels and the unofficial trade of live cattle, camels, sheep and goats from Ethiopia sold to Somalia, Kenya and Djibouti generates an estimated total value of between US$250 and US$300 million annually (100 times more than the official figure). This trade helps lower food prices, increase food security, relieve border tensions and promote regional integration. However, there are also risks as the unregulated and undocumented nature of this trade runs risks, such as allowing disease to spread more easily across national borders. Furthermore, governments are unhappy with lost tax revenue and foreign exchange revenues.” ref

Pastoralism

“Pastoralism is a form of animal husbandry where domesticated animals (known as “livestock“) are released onto large vegetated outdoor lands (pastures) for grazing, historically by nomadic people who moved around with their herds. The animal species involved include cattle, camels, goats, yaks, llamas, reindeer, horses, and sheep. Pastoralism occurs in many variations throughout the world, generally where environmental characteristics such as aridity, poor soils, cold or hot temperatures, and lack of water make crop-growing difficult or impossible. Operating in more extreme environments with more marginal lands means that pastoral communities are very vulnerable to the effects of global warming.” ref

“Pastoralism remains a way of life in many geographic areas, including Africa, the Tibetan plateau, the Eurasian steppes, the Andes, Patagonia, the Pampas, Australia and many other places. As of 2019, between 200 million and 500 million people globally practiced pastoralism, and 75% of all countries had pastoral communities. Pastoral communities have different levels of mobility. Sedentary pastoralism has become more common as the hardening of political borders, land tenures, expansion of crop farming, and construction of fences and dedicated agricultural buildings all reduce the ability to move livestock around freely, leading to the rise of pastoral farming on established grazing-zones (sometimes called “ranches“).” ref

“Sedentary pastoralists may also raise crops and livestock together in the form of mixed farming, for the purpose of diversifying productivity, obtaining manure for organic farming, and improving pasture conditions for their livestock. Mobile pastoralism includes moving herds locally across short distances in search of fresh forage and water (something that can occur daily or even within a few hours); as well as transhumance, where herders routinely move animals between different seasonal pastures across regions; and nomadism, where nomadic pastoralists and their families move with the animals in search for any available grazing-grounds—without much long-term planning. Grazing in woodlands and forests may be referred to as silvopastoralism. Those who practice pastoralism are called “pastoralists.” ref

“Pastoralist herds interact with their environment, and mediate human relations with the environment as a way of turning uncultivated plants (like wild grass) into food. In many places, grazing herds on savannas and in woodlands can help maintain the biodiversity of such landscapes and prevent them from evolving into dense shrublands or forests. Grazing and browsing at the appropriate levels often can increase biodiversity in Mediterranean climate regions. Pastoralists shape ecosystems in different ways: some communities use fire to make ecosystems more suitable for grazing and browsing animals. One theory suggests that pastoralism developed from mixed farming. Bates and Lees proposed that the incorporation of irrigation into farming resulted in specialization.” ref

“Advantages of mixed farming include reducing risk of failure, spreading labor, and re-utilizing resources. The importance of these advantages and disadvantages to different farmers or farming societies differs according to the sociocultural preferences of the farmers and the biophysical conditions as determined by rainfall, radiation, soil type, and disease. The increased productivity of irrigation agriculture led to an increase in population and an added impact on resources. Bordering areas of land remained in use for animal breeding. This meant that large distances had to be covered by herds to collect sufficient forage. Specialization occurred as a result of the increasing importance of both intensive agriculture and pastoralism. Both agriculture and pastoralism developed alongside each other, with continuous interactions.” ref

“A different theory suggests that pastoralism evolved from the hunting and gathering. Hunters of wild goats and sheep were knowledgeable about herd mobility and the needs of the animals. Such hunters were mobile and followed the herds on their seasonal rounds. Undomesticated herds were chosen to become more controllable for the proto-pastoralist nomadic hunter and gatherer groups by taming and domesticating them. Hunter-gatherers’ strategies in the past have been very diverse and contingent upon the local environmental conditions, like those of mixed farmers. Foraging strategies have included hunting or trapping big game and smaller animals, fishing, collecting shellfish or insects, and gathering wild-plant foods such as fruits, seeds, and nuts.” ref 

“These diverse strategies for survival amongst the migratory herds could also provide an evolutionary route towards nomadic pastoralism. Pastoralism occurs in uncultivated areas. Wild animals eat the forage from the marginal lands and humans survive from milk, blood, and often meat of the herds and often trade by-products like wool and milk for money and food. Pastoralists do not exist at basic subsistence. Some pastoralists supplement herding with hunting and gathering, fishing and/or small-scale farming or pastoral farming.” ref

“Pastoralists often compile wealth and participate in international trade. Pastoralists have trade relations with agriculturalists, horticulturalists, and other groups. Pastoralists are not extensively dependent on milk, blood, and meat of their herd. McCabe noted that when common property institutions are created, in long-lived communities, resource sustainability is much higher, which is evident in the East African grasslands of pastoralist populations. However, the property rights structure is only one of the many different parameters that affect the sustainability of resources, and common or private property per se, does not necessarily lead to sustainability.” ref

“Mobility allows pastoralists to adapt to the environment, which opens up the possibility for both fertile and infertile regions to support human existence. Important components of pastoralism include low population density, mobility, vitality, and intricate information systems. The system is transformed to fit the environment rather than adjusting the environment to support the “food production system.” Mobile pastoralists can often cover a radius of a hundred to five hundred kilometers. The violent herder–farmer conflicts in Nigeria, Mali, Sudan, Ethiopia and other countries in the Sahel and Horn of Africa regions have been exacerbated by climate change, land degradation, and population growth.” ref

“Pastoralists and their livestock have impacted the environment. Lands long used for pastoralism have transformed under the forces of grazing livestock and anthropogenic fire. Fire was a method of revitalizing pastureland and preventing forest regrowth. The collective environmental weights of fire and livestock browsing have transformed landscapes in many parts of the world. Fire has permitted pastoralists to tend the land for their livestock. Political boundaries are based on environmental boundaries. The Maquis shrublands of the Mediterranean region are dominated by pyrophytic plants that thrive under conditions of anthropogenic fire and livestock grazing.” ref

“Nomadic pastoralists have a global food-producing strategy depending on the management of herd animals for meat, skin, wool, milk, blood, manure, and transport. Nomadic pastoralism is practiced in different climates and environments with daily movement and seasonal migration. Pastoralists are among the most flexible populations. Pastoralist societies have had field armed men protect their livestock and their people and then to return into a disorganized pattern of foraging. The products of the herd animals are the most important resources, although the use of other resources, including domesticated and wild plants, hunted animals, and goods accessible in a market economy are not excluded. The boundaries between states impact the viability of subsistence and trade relations with cultivators.” ref

“Some pastoralists are constantly moving, which may put them at odds with sedentary people of towns and cities. The resulting conflicts can result in war for disputed lands. These disputes are recorded in ancient times in the Middle East, as well as for East Asia. Other pastoralists are able to remain in the same location which results in longer-standing housing. Different mobility patterns can be observed: Somali pastoralists keep their animals in one of the harshest environments but they have evolved of the centuries. Somalis have well developed pastoral culture where complete system of life and governance has been refined. Somali poetry depicts humans interactions, pastoral animals, beasts on the prowl, and other natural things such the rain, celestial events and historic events of significance.” ref 

“The basis of pastoral organization almost everywhere in the world is the clan, a set of patrilineally related households traced (in theory) to an apical ancestor. The great majority of pastoral societies are patrilineal and male-dominated.” ref

“The basis of pastoral organization almost everywhere in the world is the clan, a set of patrilineally related households traced (in theory) to an apical ancestor. Such groupings can be very small, and the ancestry stretch back for only a short time span, or so great that the ancestral figure is semi-mythical, in which case the working kin group is a lineage. The preservation of these genealogies is very important – especially to the aristocratic strata of nomad society, because it makes their position legitimate.” ref

“Well-known exceptions to the rule are the Tuareg, who had matrilineal descent groups in some areas, and subarctic peoples such as the Saami, the Chukchi and the Koryak, who had neither unilineal descent groups nor elaborate genealogies. One of the most distinctive features of pastoralism in East Africa and the Horn of Africa is the system of age-sets. Among the Boran of southern Ethiopia, for example, men born within a seven-year cohort fall into a named age-set, which has rights and privileges within society as well as acting as a powerful force for cohesion and a calendrical system.” ref

“A key aspect of pastoral systems is the strong relationship between wealth in livestock and labor. Herds that grow beyond a certain size cannot be managed with household labor alone, and outside herders must be sought. In the twentieth century, this is generally through hired labor, but formerly it was often through slavery or vassal castes. The great herds of cattle owned today by Fuel herders in the Niger were managed by slave labor in the nineteenth century, and many pastoral societies in Africa and the Near East developed elaborate caste systems based on slaves and non-slaves. In the case of the Tuareg, for example, society was divided into:

• Imajer.en: nobles;
• Iklan: (former) slaves;
• Izeggar.en: agricultural laborers;
• Ineden: blacksmiths.” ref

“Marriages between these groups were formerly forbidden, and even today remain uncommon. When slaves were freed in the colonial era, they stayed with their original camps for some time, but have gradually broken away and now form independent households, often remote from their original site so that traditional authority cannot be brought to bear. Similar systems were found throughout much of the Bedouin areas and in the Horn of Africa.” ref

ROLE OF WOMEN IN PASTORAL SOCIETY

“The role of women among pastoralists has been much discussed, in part because pastoral societies are more male-dominated than most other subsistence systems. Despite the well-known exception of the Saharan Tuareg, the great majority of pastoral societies are patrilineal and male-dominated. The reasons for this are much debated, but the root cause appears to be related to the importance of not dispersing viable herds. In an exogamous system, if women can own significant herds of their own, they will take these away on marriage to a new camp and potentially deplete the herd of an individual household. Many pastoral societies practice pre-inheritance, the father dispersing the herd among his sons prior to his death, since the principle of patrilocality means that the animals will anyway remain in the same physical herd.” ref

“In pastoral societies, particularly those affected by Islamic inheritance rules, some animals go to daughters on the death of the household head, but these are then “managed” by the women’s brothers. In most pastoral societies gender roles are strongly marked, and patterns seem to be extremely similar across the world. Women are typically responsible for milking and dairy processing; they may or may not sell the milk, and they usually have control over the proceeds in order to feed the family. Men are responsible for herding and selling meat animals. In systems in which herds are split, women usually stay at fixed homesteads while men go away with the animals.” ref

“Pastoral societies typically tend towards monogamy because of the importance of the division of labor. In other words, for a pastoral household to be viable, there must be a wife to carry out key tasks. If there are too many polygynous households, the system will become unviable. There are exceptions to this rule, the Maasai being one well-known example. The Maasai system of age-sets, in which young men are assigned to a social category, makes it possible for older men to have several wives because moran warriors are not allowed to marry. Only after a young man has graduated from being a moran is he able to marry.” ref

PASTORAL IDENTITIES

“Throughout much of Eurasia, pastoralism is interwoven with the culture of itinerants; groups who move around supplying services to fixed communities. The most well-known of these are the gypsies, who are spread from Wales to India under a variety of names and associated with a variety of occupations. Rao (1982, 1987) calls these groups “peripatetics” and describes some of their activities, notably those concerned with crafts. As with gypsies and horse-coping, some peripatetics play an important role in livestock trade, although they generally do not produce food. Such groups are particularly numerous in the area between Afghanistan and India. They fall into casted, endogamous groups and are often stereotyped as ethnically distinct, as are pastoralists, whom national governments often put into the same category as these other. In Afghanistan, both pastoral nomads and peripatetics live in tents; those of livestock producers are made out of black goat-hair, while peripatetic tents are white.” ref

LAND TENURE AND THE COMMON PROPERTY RESOURCES DEBATE

“Pastoral systems have been at the heart of many debates on the nature of common property resources. While settled farmers usually develop relatively explicit systems of tenure, many pastoral peoples have fluid systems that are hard to pin down. This is in keeping with their opportunistic grazing strategies. When pasture is extremely patchy and likely to appear at different sites each year, investing heavily in ownership of a specific piece of land is hardly worthwhile. The negative side of this is that farmers can come and cultivate the land that herders regularly use for grazing their stock, without having to ask for permission. Because they are generally operating in remote areas without access to schools, pastoralists rarely have the literacy necessary to register land claims and so are outcompeted by both farmers and urban-based ranchers. In Jordan, the Badia rangelands were the preserve of sheep-herders because agriculture was considered to be impossible.” ref

“However, a combination of more boreholes and new irrigation techniques is pushing farms ever further into traditional grazing land, and the government is unwilling to halt this process because of its own political constituency. Tenure is thus divided by both ecology and the potential for agriculture. In much of the snowy steppe, agriculture is not practical, so pastoralists compete with one another for prime sites. The same is true in the subarctic regions, where reindeer herders do not interact with farmers. In much of Central Asia, the command economies overrode traditional access rights and created mapped and demarcated territories for collectivized units. These are in the process of being dismantled, and more traditional access rights are being reasserted. However, legal frameworks for this new situation are only now being developed.” ref

“The tenure of pastoralists in all parts of the world is not deemed sufficiently strong to prevent it from being overridden by the State in its search for minerals. Land can be appropriated for building and transport infrastructure, generally without compensation. There is no doubt that, if pastoralism is to survive, effective tenure must be developed in many parts of the world. This is proving difficult, because few governments have the political will to protect pastoralists against the vested interests of urban groups. The usual indicator of tenure in the ranching areas is the fence, a high-investment strategy that is only effective in countries where specific legal frameworks are in place.” ref 

Double Marginalized Livelihoods: Invisible Gender Inequality in Pastoral Societies

“Abstract: Achieving gender equality is the Third Millennium Development Goal, and the major challenge to poverty reduction is the inability of governments to address this at grass root levels. This study is therefore aimed at assessing gender inequality as it pertains to socio-economic factors in (agro-) pastoral societies. It tries to explain how “invisible” forces perpetuate gender inequality, based on data collected from male and female household heads and community representatives. The findings indicate that in comparison with men, women lack access to control rights over livestock, land, and income, which are critical to securing a sustainable livelihood. However, this inequality remains invisible to women who appear to readily submit to local customs, and to the community at large due to a lack of public awareness and gender based interventions. In addition, violence against women is perpetuated through traditional beliefs and sustained by tourists to the area. As a result, (agro-) pastoral woman face double marginalization, for being pastoralist, and for being a woman.” ref

Nomads

“Nomads are communities without fixed habitation who regularly move to and from areas. Such groups include hunter-gatherers, pastoral nomads (owning livestock), tinkers and trader nomads. Nomadic hunting and gathering—following seasonally available wild plants and game—is by far the oldest human subsistence method. Pastoralists raise herds of domesticated livestock, driving or accompanying them in patterns that normally avoid depleting pastures beyond their ability to recover. Nomadism is also a lifestyle adapted to infertile regions such as steppe, tundra, or ice and sand, where mobility is the most efficient strategy for exploiting scarce resources. For example, many groups living in the tundra are reindeer herders and are semi-nomadic, following forage for their animals. Sometimes also described as “nomadic” are various itinerant populations who move among densely populated areas to offer specialized services (crafts or trades) to their residents—external consultants, for example. These groups are known as “peripatetic nomads.” ref

“Nomadic pastoralism seems to have developed first as a part of the secondary-products revolution proposed by Andrew Sherratt, in which early pre-pottery Neolithic cultures that had used animals as live meat (“on the hoof”) also began using animals for their secondary products, for example: milk and its associated dairy products, wool and other animal hair, hides (and consequently leather), manure (for fuel and fertilizer), and traction.” ref

“The first nomadic pastoral society developed in the period from 8,500 to 6,500 BCE or around 10,500 to 8,500 years ago in the area of the southern Levant. There, during a period of increasing aridity, Pre-Pottery Neolithic B (PPNB) cultures in the Sinai were replaced by a nomadic, pastoral pottery-using culture, which seems to have been a cultural fusion between them and a newly-arrived Mesolithic people from Egypt (the Harifian culture), adopting their nomadic hunting lifestyle to the raising of stock.” ref

“Harifian is a specialized regional cultural development of the Epipalaeolithic of the Negev Desert. It corresponds to the latest stages of the Natufian culture. Like the Natufian, Harifian is characterized by semi-subterranean houses. These are often more elaborate than those found at Natufian sites. For the first time arrowheads are found among the stone tool kit. The Harifian dates to between approximately 10,800/10,500 and 10,000/10,200 years ago. It is restricted to the Sinai and Negev, and is probably broadly contemporary with the Late Natufian or Pre-Pottery Neolithic A (PPNA). Microlithic points are a characteristic feature of the industry, with the Harif point being both new and particularly diagnostic – Bar-Yosef (1998) suggests that it is an indication of improved hunting techniques. Lunates, isosceles and other triangular forms were backed with retouch, and some Helwan lunates are found. This industry contrasts with the Desert Natufian which did not have the roughly triangular points in its assemblage. There are two main groups within the Harifian. One group consists of ephemeral base camps in the north of Sinai and western Negev, where stone points comprise up to 88% of all microliths, accompanied by only a few lunates and triangles. The other group consists of base camps and smaller campsites in the Negev and features a greater number of lunates and triangles than points. These sites probably represent functional rather than chronological differences. The presence of Khiam points in some sites indicates that there was communication with other areas in the Levant at this time.” ref

“This lifestyle quickly developed into what Jaris Yurins has called the circum-Arabian nomadic pastoral techno-complex and is possibly associated with the appearance of Semitic languages in the region of the Ancient Near East. The rapid spread of such nomadic pastoralism was typical of such later developments as of the Yamnaya culture of the horse and cattle nomads of the Eurasian steppe (c. 3300–2600 BCE), and of the Mongol spread in the later Middle Ages. Yamnaya steppe pastoralists from the Pontic–Caspian steppe, who were among the first to master horseback riding, played a key role in Indo-European migrations and in the spread of Indo-European languages across Eurasia.” ref

“Trekboers in southern Africa adopted nomadism from the 17th century. Some elements of gaucho culture in colonial South America also re-invented nomadic lifestyles. Hunter-gatherers (also known as foragers) move from campsite to campsite, following game and wild fruits and vegetables. Hunting and gathering describes early peoples’ subsistence living style. Following the development of agriculture, most hunter-gatherers were eventually either displaced or converted to farming or pastoralist groups. Only a few contemporary societies, such as the Pygmies, the Hadza people, and some uncontacted tribes in the Amazon rainforest, are classified as hunter-gatherers; some of these societies supplement, sometimes extensively, their foraging activity with farming or animal husbandry.” ref

“Pastoral nomads are nomads moving between pastures. Nomadic pastoralism is thought to have developed in three stages that accompanied population growth and an increase in the complexity of social organization. Karim Sadr has proposed the following stages:

• Pastoralism: This is a mixed economy with a symbiosis within the family.
• Agropastoralism: This is when symbiosis is between segments or clans within an ethnic group.
• True Nomadism: This is when symbiosis is at the regional level, generally between specialised nomadic and agricultural populations.” ref

“The pastoralists are sedentary to a certain area, as they move between the permanent spring, summer, autumn and winter (or dry and wet season) pastures for their livestock. The nomads moved depending on the availability of resources.” ref 

Nguni cattle

“The Nguni is a cattle breed indigenous to Southern Africa. A hybrid of different Indian and later European cattle breeds, they were introduced by pastoralist tribes ancestral to modern Nguni people to Southern Africa during their migration from the North of the continent. Nguni cattle are dairy and beef cattle. Nguni cattle are known for their fertility and resistance to diseases, being the favourite and most beloved breed amongst the local Bantu-speaking people of southern Africa (South Africa, Eswatini, Namibia, Zimbabwe, Botswana, and Angola). They are a principal form of Sanga cattle (collective name for indigenous cattle of some regions in Africa), which originated as hybrids of Zebu (originating in South Aisa, first domesticated between 7,000 and 6,000 years ago, and In some regions, zebu have significant religious meaning) and humpless cattle in East Africa. DNA analyses have confirmed that they are a combination of Bos indicus and Bos taurus, that is a combination of different Zebu and European cattle breeds.” ref 

Zebu cattle

“Zebu cattle were found to derive from the Indian form of aurochs and have first been domesticated between 7,000 and 6,000 years ago at Mehrgarh, present-day Pakistan, by people linked to or coming from Mesopotamia. Its wild ancestor, the Indian aurochs, became extinct during the Indus Valley civilisation likely due to habitat loss, caused by expanding pastoralism and interbreeding with domestic zebu. Its latest remains ever found were dated to 3,800 years ago, making it the first of the three aurochs subspecies to die out. Archaeological evidence including depictions on pottery and rocks suggests that humped cattle likely imported from the Near East was present in Egypt around 4,000 years ago. Its first appearance in the Subsahara is dated to after 700 CE and it was introduced to the Horn of Africa around 1000 CE. Phylogenetic analysis revealed that all the zebu Y chromosome haplotype groups are found in three different lineages: Y3A, the most predominant and cosmopolitan lineage; Y3B, only observed in West Africa; and Y3C, predominant in south and northeast India.” ref 

Nguni languages

“The Nguni languages are a group of Bantu languages spoken in southern Africa (mainly South Africa, Zimbabwe and Eswatini) by the Nguni people. Nguni languages include Xhosa, Hlubi, Zulu, Ndebele, and Swati. The appellation “Nguni” derives from the Nguni cattle type. Ngoni (see below) is an older, or a shifted, variant. It is sometimes argued that the use of Nguni as a generic label suggests a historical monolithic unity of the people in question, where in fact the situation may have been more complex. The linguistic use of the label (referring to a subgrouping of Bantu) is relatively stable.” ref

“Within a subset of Southern Bantu, the label “Nguni” is used both genetically (in the linguistic sense) and typologically (quite apart from any historical significance). The Nguni languages are closely related, and in many instances different languages are mutually intelligible; in this way, Nguni languages might better be construed as a dialect continuum than as a cluster of separate languages. On more than one occasion, proposals have been put forward to create a unified Nguni language.” ref

“In scholarly literature on southern African languages, the linguistic classificatory category “Nguni” is traditionally considered to subsume two subgroups: “Zunda Nguni” and “Tekela Nguni”. This division is based principally on the salient phonological distinction between corresponding coronal consonants: Zunda /z/ and Tekela /t/ (thus the native form of the name Swati and the better-known Zulu form Swazi), but there is a host of additional linguistic variables that enables a relatively straightforward division into these two substreams of Nguni.” ref 

Nguni people

“The Nguni people are a linguistic cultural group of Bantu cattle herders who migrated from central Africa into Southern Africa, made up of ethnic groups formed from hunter-gatherer pygmy and proto-agrarians, with offshoots in neighboring colonially-created countries in Southern Africa. Swazi (or Swati) people live in both South Africa and Eswatini, while Ndebele people live in both South Africa and Zimbabwe.” ref

“The Xhosa were formed out of pastoralist blended from pygmies and proto-Bantu agro-pastoralists and established federations (AbaThembu, AmaMpondo, AmaXhosa, and AmaMpondomise) in the 5th century AD. The Xhosa trace their name back to the Khoikhoi, as the Khoi named them Xhosa, signifying “violent angry” people. The recent homeland of the Xhosa people is marked by lands in the Eastern Cape from the Gamtoos River up to Umzimkhulu near Natal, which confined and restricted their pastoral ancestors from the rest of the Cape by an expanding and setting of the VOC Cape Colony frontier. This closed frontier was set in the late 1700s. The Xhosa often called the “Red Blanket People,” are speakers of Bantu languages living in south-east South Africa and in the last two centuries throughout the southern and central-southern parts of the country.” ref

“Most of what is believed about ancient Nguni history comes from oral history and legends. Traditionally, their partial ancestors are said to have migrated to Africa’s Great Lakes region from the north. According to linguistic evidence and historians (including John H. Robertson, Rebecca Bradley, T. Russell, Fabio Silva, and James Steele), some of the ancestors of the Nguni people migrated from west of the geographic centre of Africa towards modern-day South Africa 7000 years ago (5000 BCE or around 7,000 years ago). Nguni ancestors had migrated within South Africa to KwaZulu-Natal by the 1st century CE and were also present in the Transvaal region at the same time. These partially nomadic ancestors of the modern Nguni people brought with them sheep, cattle, goats, and horticultural crops, many of which had never been used in South Africa at that time.” ref

“Other provinces in present-day South Africa, such as the Cape, saw the emergence of Nguni speakers around the same time. Some groups split off and settled along the way, while others kept going. Thus, the following settlement pattern formed: the southern Ndebele in the north, the Swazi in the northeast, the Xhosa in the south, and the Zulu towards the east. Because these peoples had a common origin, their languages and cultures show marked similarities. Partial ancestors of the Nguni eventually met and merged with San hunters, which accounts for the use of click consonants in the languages of the Nguni.” ref

“Within the Nguni nations, the clan, based on male ancestry, formed the highest social unit. Each clan was led by a chieftain. Influential men tried to achieve independence by creating their own clan. The power of a chieftain often depended on how well he could hold his clan together. From about 1800, the rise of the Zulu clan of the Nguni, and the consequent Mfecane that accompanied the expansion of the Zulus under Shaka helped to drive a process of alliance and consolidation among many of the smaller clans.” ref

“For example, the kingdom of Eswatini was formed in the early nineteenth century by different Nguni groups allying with the Dlamini clan against the threat of external attack. Today, the kingdom encompasses many different clans that speak a Nguni language called Swati and are loyal to the king of Eswatini, who is also the head of the Dlamini clan. Ngunis may be Christians (Catholics or Protestants), practitioners of African traditional religions or members of forms of Christianity modified with traditional African values.They also follow a mix of these two religions, usually not separately.” ref

Khoe-San Genomes Reveal Unique Variation and Confirm the Deepest Population Divergence in Homo sapiens

“Abstract: The southern African indigenous Khoe-San populations harbor the most divergent lineages of all living peoples. Exploring their genomes is key to understanding deep human history. We sequenced 25 full genomes from five Khoe-San populations, revealing many novel variants, that 25% of variants are unique to the Khoe-San, and that the Khoe-San group harbors the greatest level of diversity across the globe. In line with previous studies, we found several gene regions with extreme values in genome-wide scans for selection, potentially caused by natural selection in the lineage leading to Homo sapiens and more recent in time. These gene regions included immunity-, sperm-, brain-, diet-, and muscle-related genes. When accounting for recent admixture, all Khoe-San groups display genetic diversity approaching the levels in other African groups and a reduction in effective population size starting around 100,000 years ago. Hence, all human groups show a reduction in effective population size commencing around the time of the Out-of-Africa migrations, which coincides with changes in the paleoclimate records, changes that potentially impacted all humans at the time.” ref

“Genetics has played an increasingly important role in revealing human evolutionary history, by demonstrating that Homo sapiens emerged from, with some groups outside Africa admixing with archaic humans. Our deepest roots include indigenous groups of current-day southern Africa, with modern-day Khoe-San representing one branch in the earliest population divergence in Homo sapiens, and all other Africans and non-Africans representing the other branch. Southern African hunter-gatherers (San) and herders (Khoekhoe) are collectively referred to as Khoe-San. Khoe-San people speak Khoisan languages, a group of languages that rely heavily on “click” sounds. Three out of the five major Khoisan language families are spoken in southern Africa, namely, Kx’a (formerly called Northern Khoisan), Tuu (formerly Southern Khoisan), and Khoe-Kwadi (formerly Central Khoisan). These three language families show no linguistic relatedness to each other. A few complete genomes from Khoe-San individuals have been investigated with poor representation among the different groups. As the Khoe-San represents one of two branches of the deepest population divergence within Homo sapiens, it is crucial to reveal their evolutionary history and their genetic diversity in order to understand the early evolutionary history of our species.” ref

“Researchers sequenced and analyzed 25 complete high-coverage genomes from five different Khoe-San groups, representing the three main Khoisan linguistic phyla, across an extensive geographic area. These genomes were placed into a global context by jointly investigating 11 previously published genomes from the HGDP panel, sequenced on the same platform and subjected to similar single nucleotide polymorphism (SNP) calling procedures, and another 67 genomes sequenced on the Complete Genomics platform. Using these data sets, we characterized genome variation across the world and inferred past population history, where Khoe-San groups showed greater genetic diversity than any other group, but still revealed a reduction in effective population size coinciding with the Out-of-Africa migrations and bottleneck. We further discovered a number of selection targets in the Khoe-San and other groups, and within our common ancestors of >300,000 years ago. These results shed new light on Pleistocene human demographic history and evolution.” ref

“Researchers found a distinct signal of eastern African/non-African affinity and shared private variants among the Khoe-San, particularly for the Nama. This outcome is consistent with recent migration of mixed (eastern African-Eurasian) herding groups to southern Africa, and potentially long-term gene flow between eastern African hunter-gatherers (e.g., Hadza) and Khoe-San. This pattern can also be seen in mtDNA and Y chromosome data, with haplogroup sharing detected between the Ju|’hoansi and Hadza.” ref

“Researchers estimated population divergence between the Khoe-San and various other groups using different and complementary approaches. We applied a mutation rate of 1.25 × 10⁻⁸ per base pair per generation and a generation time of 30 years to convert estimates to years ago in the past. Consistent with previous studies, the deepest divergences included the Khoe-San populations; a result probably not caused by “archaic admixture” into the Khoe-San.” ref

“Modern-day Khoe-San have, however, >10% of their genetic material tracing to a recent admixture with external groups. By sequencing the genome of the Stone Age boy from Ballito Bay (BBA), South Africa, the deepest population divergence in Homo sapiens was estimated to 350,000–260,000 years ago. Consistent with the recent admixture into all modern-day Khoe-San groups, which reduces population divergence time estimates, we found the mean divergence time of all Khoe-San populations from all other groups to be within the 200,000–300,000 years ago range.” ref

“These dates correlate well with previous estimates that also fall within the 200,000–300,000 years ago range when applying the mutation rate used here. The Ju|’hoansi (with the lowest level of recent admixture) had a point estimate of ∼270,000 years ago (∼9,000 generations), SD 20,000 years ago (GphoCS method; TT method: ∼260,000 years ago, SD 12,000 years ago), whereas the Nama (with the greatest level of recent admixture) had a point estimate of ∼210,000 years ago, SD 30,000 years ago (TT method: ∼210,000 years ago, SD 30,000 years ago). The Mbuti then diverged around ∼220 ka, SD 10 ka (TT method: 215,000 years ago, SD 9,000 years ago), with the other population divergences occurring subsequently. We inferred a mean divergence time of ∼160,000 years ago, SD 20,000 years ago (TT method: ∼190,000 years ago, SD 20,000 years ago) among the different San groups, consistent with previous estimates.” ref

“If researchers jointly analyze two individuals (four haploid genomes) instead of one, it should provide more resolution on the timing of the bottleneck, because the mean time to first coalescence for four haploid genomes is 85,000 years ago (assuming an average ancestral N_e of 17,000 and a generation time of 30 years). For this analysis, we found that all Khoe-San groups showed a reduction to 1/3 of the previous N_e between 100,000 and 20,000 years ago. The same pattern was also observed with samples of five individuals (ten haploid genomes), though it could not be detected with samples of one single modern-day Khoe-San individual.” ref

“Researchers simulated data under a bottleneck model and ran MSMC on samples of one, two, four, and five individuals under a range of varying conditions of bottleneck strength, duration, and age. From this investigation, we observed a qualitatively similar pattern of reduced power to infer population-size changes around 80 ka when basing the inference on single genomes. Thus, all human groups appeared to have suffered reduced N_e, of varying degrees, between ∼100,000 and ∼20,000 years ago; declining to between 50% and 10% of an N_e of ∼30,000 at ∼300,000 years ago a.” ref

“The genetic diversity among the Khoe-San is the greatest among all human groups across the world, which, in part, is explained by relatively recent (pre-colonial) admixture. When the admixed DNA portion was excluded, the genetic diversity of the Khoe-San approached levels seen in other African populations. All human groups, including the Khoe-San, showed a reduction in N_e (between 1/3 and 1/10) between ∼100,000 and 20,000 years ago. The early phase of the reduction coincides with the Out-of-Africa bottleneck for non-Africans.” ref

“Sub-Saharan African populations would not have been impacted by this migration bottleneck, but they all (including the Khoe-San) show a reduction in N_e. This observation suggests that an additional factor—beyond the migration out of Africa—impacted all humans at this time, perhaps the change in climate. For example, work on the Lake Malawi core indicates severe drought and low-lake stage occurring between ∼109,000 and 92,000 years ago when the area is also shifting from leaf- to grass-dominated vegetation, which roughly aligns with a change from warm toward colder temperatures for Africa. These events may have caused a reduction in the number of humans; potentially also driving them out of arid African regions, such as the Sahara, and into western Asia.” ref

“By revealing substantial and previously unknown genetic variation, researchers demonstrate that a sizable portion of human genetic variation, including common variants, remains undiscovered among populations often overlooked in medical genetics. Researchers inferred adaptation signals in the genomes and found an overrepresentation of these signals overlapping immunity genes, irrespective of group or time period. This suggests that immunity genes have been under selection throughout human evolutionary history and across the globe.” ref 

San people

“The compound Khoisan is used to refer to the pastoralist Khoi and the foraging San collectively.  The San refer to themselves as their individual nations, such as ǃKung (also spelled ǃXuun, including the Juǀʼhoansi), ǀXam, Nǁnǂe (part of the ǂKhomani), Kxoe (Khwe and ǁAni), Haiǁom, Ncoakhoe, Tshuwau, Gǁana and Gǀui (ǀGwi), etc. Representatives of San peoples in 2003 stated their preference for the use of such individual group names, where possible, over the use of the collective term San.” ref

“The San peoples (also Saan), or Bushmen, are the members of any of the indigenous hunter-gatherer cultures of southern Africa, and the oldest surviving cultures of the region. Their recent ancestral territories span Botswana, Namibia, Angola, Zambia, Zimbabwe, Lesotho, and South Africa. The San speak, or their ancestors spoke, languages of the Khoe, Tuu, and Kxʼa language families, and can be defined as a people only in contrast to neighboring pastoralists such as the Khoekhoe and descendants of more recent waves of immigration such as the Bantu, Europeans, and Asians.” ref

“In Khoekhoegowab, the term “San” has a long vowel and is spelled Sān. It is an exonym meaning “foragers” and is used in a derogatory manner to describe people too poor to have cattle. Based on observation of lifestyle, this term has been applied to speakers of three distinct language families living between the Okavango River in Botswana and Etosha National Park in northwestern Namibia, extending up into southern Angola; central peoples of most of Namibia and Botswana, extending into Zambia and Zimbabwe; and the southern people in the central Kalahari towards the Molopo River, who are the last remnant of the previously extensive indigenous peoples of southern Africa.” ref

“The designations “Bushmen” and “San” are both exonyms. The San have no collective word for themselves in their own languages. “San” comes from a derogatory Khoekhoe word used to refer to foragers without cattle or other wealth, from a root saa “picking up from the ground” + plural -n in the Haiǁom dialect. “Bushmen” is the older cover term, but “San” was widely adopted in the West by the late 1990s. The term Bushmen, from 17th-century Dutch Bosjesmans, is still used by others and to self-identify, but is now considered pejorative or derogatory by many South Africans.” ref

“The hunter-gatherer San are among the oldest cultures on Earth, and are thought to be descended from the first inhabitants of what is now Botswana and South Africa. The historical presence of the San in Botswana is particularly evident in northern Botswana’s Tsodilo Hills region. San were traditionally semi-nomadic, moving seasonally within certain defined areas based on the availability of resources such as water, game animals, and edible plants. Peoples related to or similar to the San occupied the southern shores throughout the eastern shrubland and may have formed a Sangoan continuum from the Red Sea to the Cape of Good Hope.” ref

“The San kinship system reflects their history as traditionally small mobile foraging bands. San kinship is similar to Inuit kinship, which uses the same set of terms as in European cultures but adds a name rule and an age rule for determining what terms to use. The age rule resolves any confusion arising from kinship terms, as the older of two people always decides what to call the younger. Relatively few names circulate (approximately 35 names per sex), and each child is named after a grandparent or another relative, but never their parents.” ref

“Children have no social duties besides playing, and leisure is very important to San of all ages. Large amounts of time are spent in conversation, joking, music, and sacred dances. Women may be leaders of their own family groups. They may also make important family and group decisions and claim ownership of water holes and foraging areas. Women are mainly involved in the gathering of food, but sometimes also partake in hunting.” ref

“Water is important in San life. During long droughts, they make use of sip wells in order to collect water. To make a sip well, a San scrapes a deep hole where the sand is damp, and inserts a long hollow grass stem into the hole. An empty ostrich egg is used to collect the water. Water is sucked into the straw from the sand, into the mouth, and then travels down another straw into the ostrich egg.” ref

“Traditionally, the San were an egalitarian society. Although they had hereditary chiefs, their authority was limited. The San made decisions among themselves by consensus, with women treated as relative equals in decision making. San economy was a gift economy, based on giving each other gifts regularly rather than on trading or purchasing goods and services. Most San are monogamous, but if a hunter is able to obtain enough food, he can afford to have a second wife as well.” ref

“A set of tools almost identical to that used by the modern San and dating to 42,000 BCE was discovered at Border Cave in KwaZulu-Natal in 2012. Historical evidence shows that certain San communities have always lived in the desert regions of the Kalahari; however, eventually nearly all other San communities in southern Africa were forced into this region. The Kalahari San remained in poverty where their richer neighbours denied them rights to the land. Before long, in both Botswana and Namibia, they found their territory drastically reduced.” ref

“Various Y chromosome studies show that the San carry some of the most divergent (earliest branching) human Y-chromosome haplogroups. These haplogroups are specific sub-groups of haplogroups A and B, the two earliest branches on the human Y-chromosome tree. Mitochondrial DNA studies also provide evidence that the San carry high frequencies of the earliest haplogroup branches in the human mitochondrial DNA tree. This DNA is inherited only from one’s mother. The most divergent (earliest branching) mitochondrial haplogroup, L0d, has been identified at its highest frequencies in the southern African San groups.” ref

“In a study published in March 2011, Brenna Henn and colleagues found that the ǂKhomani San, as well as the Sandawe and Hadza peoples of Tanzania, were the most genetically diverse of any living humans studied. This high degree of genetic diversity hints at the origin of anatomically modern humans. A 2008 study suggested that the San may have been isolated from other original ancestral groups for as much as 50,000 to 100,000 years and later rejoined, re-integrating into the rest of the human gene pool. A DNA study of fully sequenced genomes, published in September 2016, showed that the ancestors of today’s San hunter-gatherers began to diverge from other human populations in Africa about 200,000 years ago and were fully isolated by 100,000 years ago.” ref

ǃKung people

“The ǃKung are one of the San peoples who live mostly on the western edge of the Kalahari desert, Ovamboland (northern Namibia and southern Angola), and Botswana. The names ǃKung (ǃXun) and Ju are variant words for ‘people’, preferred by different ǃKung groups. This band level society used traditional methods of hunting and gathering for subsistence up until the 1970s. Today, the great majority of ǃKung people live in the villages of Bantu pastoralists and European ranchers.” ref

“The ǃKung people of Southern Africa recognize a Supreme Being, ǃXu, who is the Creator and Upholder of life. Like other African High Gods, he also punishes man by means of the weather, and the Otjimpolo-ǃKung know him as Erob, who “knows everything”. They also have animistic and animatistic beliefs, which means they believe in both personifications and impersonal forces. For example, they recall a culture hero named Prishiboro, whose wife was an elephant. Prishiboro’s older brother tricked him into killing his wife and eating her flesh. Her herd tried to kill Prishiboro in revenge, but his brother defeated them.” ref

“Amongst the ǃKung there is a strong belief in the existence of spirits of the dead (llgauwasi) who live immortally in the sky. The llgauwasi can come to the earth and interact with humans. There is no particular connection to personal ancestors but the ǃKung fear the llgauwasi, pray to them for sympathy and mercy as well as call on them in anger.” ref

“The ǃKung practice shamanism to communicate with the spirit world, and to cure what they call “Star Sickness”. The communication with the spirit world is done by a natural healer entering a trance state and running through a fire, thereby chasing away bad spirits. Star Sickness is cured by laying hands on the diseased. Nisa, a ǃKung woman, reported through anthropologist Marjorie Shostak that a healer in training is given a root to help induce trance. Nisa said, “I drank it a number of times and threw up again and again. Finally, I started to tremble. People rubbed my body as I sat there feeling the effect getting stronger and stronger. … Trance-medicine really hurts! As you begin to trance, the nǀum [power to heal] slowly heats inside you and pulls at you. It rises until it grabs your insides and takes your thoughts away.” ref

“Healing rites are a primary part of the ǃKung culture. In the ǃKung state of mind, having health is equivalent to having social harmony, meaning that relationships within the group are stable and open between other people. Any ǃKung can become a healer because it “is a status accessible to all,” but it is a grand aspiration of many members because of its importance. Even though there is no restriction of the power, “nearly half the men and one-third of the women are acknowledged of having the power to heal,” but with the responsibility comes great pain and hardship. To become a healer, aspirants must become an apprentice and learn from older healers. Their training includes the older healer having to “go into a trance to teach the novices, rubbing their own sweat onto the pupils’ centers – their bellies, backs, foreheads, and spines.” “Most of the apprentices have the intentions of becoming a healer but then become frightened or have a lack of ambition and discontinue.” ref

“The ǃKung term for this powerful healing force is nǀum. This force resides in the bellies of men and women who have gone through the training and have become a healer. Healing can be transmitted through the ǃkia dance that begins at sundown and continues through the night. The ǃkia can be translated as “trance” which can give a physical image of a sleeping enchantment. While they dance, “in preparation for entering a trance state to effect a cure, the substance [the nǀum] heats up and, boiling, travels up the healer’s spine to explode with therapeutic power in the brain.” While the healers are in the trance they propel themselves in a journey to seek out the sickness and argue with the spirits. Women on the other hand have a special medicine called the gwah which starts in the stomachs and kidneys. During the Drum Dance, they enter the ǃkia state and the gwah travels up the spine and lodges in the neck.” ref

“In order to obtain the gwah power the women, “chop up the root of a short shrub, boil it into a tea and drink it.” They do not need to drink the tea every time because the power they obtain lasts a lifetime. The community of the ǃKung fully supports the healers and depends heavily on them. They have trust in the healers and the teachers to guide them psychologically and spiritually through life. The ǃKung have a saying: “Healing makes their hearts happy, and a happy heart is one that reflects a sense of community.” Because of their longing to keep the peace between people, their community is tranquil.” ref

“Traditionally, especially among Juǀʼhoansi ǃKung, women generally collect plant foods and water while men hunt. However, these gender roles are not strict and people do all jobs as needed with little or no stigma. Women generally take care of children and prepare food. However, this does restrict them to their homes, since these activities are generally done with, or close to, others, so women can socialize and help each other. Men are also engaged in these activities. Children are raised in village groups of other children of a wide age range. Sexual activities amongst children are seen as natural play for both sexes.” ref

“ǃKung women often share an intimate sociability and spend many hours together discussing their lives, enjoying each other’s company and children. In the short documentary film A Group of Women, ǃKung women rest, talk and nurse their babies while lying in the shade of a baobab tree. This illustrates “collective mothering”, where several women support each other and share the nurturing role. Marriage is the major focus of alliance formation between groups of ǃKung. When a woman starts to develop, she is considered ready for marriage. Every first marriage is arranged. The culture of the ǃKung is “being directed at marriage itself, rather than at a specific man.” ref 

“Even though it does not matter who the man is, the woman’s family is looking for a specific type of man. On the marriage day, the tradition is the “marriage-by-capture” ceremony in which the bride is forcibly removed from her hut and presented to her groom. During the ceremony, the bride has her head covered and is carried and then laid down in the hut while the groom is led to the hut and sits beside the door. The couple stay respectfully apart from each other and do not join the wedding festivities. After the party is over, they spend the night together and the next morning they are ceremonially rubbed with oil by the husband’s mother.” ref

“Girls who are displeased with their parents’ selection may violently protest against the marriage by kicking and screaming and running away at the end of the ceremony. After she has run away, this may result in the dissolution of the arrangement. Half of all first-time marriages end in divorce, but because it is common, the divorce process is not long. Anthropologist Marjorie Shostak generalizes that, “Everyone in the village expresses a point of view” on the marriage and if the couple should be divorced or not. After the village weighs in, they are divorced and can live in their separate huts with their family. Relations between divorced individuals are usually quite amicable, with former partners living near one another and maintaining a cordial relationship. After a woman’s first divorce, she is free to marry a man of her choosing or stay single and live on her own.” ref

“Unlike other complex food-foraging groups, it is unusual for the ǃKung to have a chieftain or headman in a position of power over the other members. These San are not devoid of leadership, but neither are they dependent on it. San groups of the Southern Kalahari have had chieftains in the past; however, there is a somewhat complicated process to gain that position. Chieftainship within these San groups is not a position with the greatest power, as they have the same social status as those members of “aged years”. Becoming chieftain is mostly nominal, though there are some responsibilities the chieftain assumes, such as becoming the group’s “logical head”. This duty entails such roles as dividing up the meat from hunters’ kills; these leaders do not receive a larger portion than any other member of the village. The !Kung people have given name to the Theory of Regal and Kungic Societal Structures due to their peacefulness and egalitarian social structure.” ref

“Kinship is one of the central organizing principles for societies like the ǃKung. Richard Borshay Lee breaks ǃKung kinship principles down into three different sets (Kinship I, Kinship II, and Kinship III or wi). Kinship I follows conventional kin terms (father, mother, brother, sister) and is based on genealogical position. Kinship II applies to name relationships, meaning that people who share the same name (ǃkunǃa) are treated as though they are kin of the same family and are assigned the same kinship term. This is a common occurrence as there are a limited number of ǃKung names. Kung names are also strictly gendered meaning that men and women cannot share the same name. Names are passed down from ancestors according to a strict set of rules, though parents are never allowed to name their children after themselves. Every member of ǃKung society fits into one of these categories, there are no neutral people.” ref

“Hunting can take days of tracking, attacking, and following a wounded beast. The Juǀʼhoansi have rites to prevent arrogance amongst male hunters. When a man kills a beast, he does not take it directly into the settlement, but leaves the body and returns as if he was unsuccessful. An older man will inquire about his hunt and remark upon his failure, to which the hunter must avoid credit and accept humility. The next day, a group will go “see if some small animal was nicked by an arrow.” Upon finding the quarry, the hunter will be reassured of the little value of the kill which is finally returned. Additionally, the kill may belong not to him, but to the person who gave him arrows (man or woman), who then follows rules on how to distribute the meat to everyone in the group.” ref

“Upon returning from a successful hunt, if the kill is transportable, it will be brought back to the village. The ǃKung promote the belief of community well-being, and so the village elders or “those of mature years” will allot meat to the members of the group. The ǃKung also believe in the betterment of their neighbors, so if the kill is too large to move or there is a surplus of meat, word will be spread to villages that are close by to come collect meat for themselves. The ǃKung language, commonly called Ju, is one of the larger click languages and belongs to the Kxʼa language family, loosely categorised in the Khoisan language group (no longer seen as one family). The ǃKung people often communicate only by high-pitched clicks when sneaking up on prey in order to stay undetected.” ref 

Hadza people

“The Hadza, or Hadzabe (Wahadzabe, in Swahili), are a protected hunter-gatherer Tanzanian indigenous ethnic group, primarily based in Baray, an administrative ward within Karatu District in southwest Arusha Region. They live around the Lake Eyasi basin in the central Rift Valley and in the neighboring Serengeti Plateau. As descendants of Tanzania’s aboriginal, pre-Bantu expansion hunter-gatherer population, they have probably occupied their current territory for thousands of years with relatively little modification to their basic way of life until the last century. They have no known close genetic relatives and their language is considered an isolate. Once classified among the Khoisan languages primarily because it has clicks, the Hadza language (Hadzane) is now thought to be an isolate, unrelated to any other language. Hadzane is an entirely oral language.” ref

“One telling of Hadza’s oral history divides their past into four epochs, each inhabited by a different culture. According to this tradition, at the beginning of time, the world was inhabited by hairy giants called the akakaanebee (first ones) or geranebee (ancient ones). The akakaanebee did not possess tools or fire; they hunted game by running it down until it fell dead; they ate meat raw. They did not build houses but slept under trees, as the Hadza do today in the dry season. In older versions of this story, they did not use fire because it was physically impossible in the earth’s primeval state. Younger Hadza, who have been to school, say that the akakaanebee did not know how to use fire.

“In the second epoch, the akakaanebee were succeeded by the xhaaxhaanebee (in-between ones), who were equally gigantic but without hair. Fire could be made and used to cook meat, but animals had grown more wary of humans and had to be chased and hunted by dogs. The xhaaxhaanebee were the first people to use medicines and charms to protect themselves from enemies and initiated the epeme rite. They lived in caves.” ref

“The third epoch was inhabited by the people of hamakwanebee (recent days), who were smaller than their predecessors. They invented bows and arrows, cooked with containers, and mastered the use of fire. They also built huts like those of Hadza today. The people of hamakwanebee were the first of the Hadza ancestors to have contact with non-foraging people, with whom they traded for iron to make knives and arrowheads. They also invented the gambling game lukuchuko. The fourth epoch continues today and is inhabited by the hamayishonebee (those of today). When discussing the hamayishonebee epoch, people often mention specific names and places and can say approximately how many generations ago events occurred.” ref

“The Hadza are not closely related to any other people. The Hadza language was once classified with the Khoisan languages because it has click consonants; however, there is no further evidence they are related. Genetically, the Hadza do not appear to be closely related to Khoisan speakers; even the Sandawe, who live around 150 kilometres (93 mi) away, diverged from the Hadza more than 15,000 years ago. Genetic testing also suggests significant admixture has occurred between the Hadza and Bantu. Minor admixture with Nilotic and Cushitic-speaking populations may have occurred in the last few thousand years. Today, a few Hadza women marry into neighbouring groups such as the Bantu Isanzu and the Nilotic Datooga, but these marriages often fail, and the women and their children return to the Hadza. In previous decades, rape and capture of Hadza women by outsiders seems to have been common. During a famine in 1918–20, some Hadza men were reported as taking Isanzu wives.” ref

“The Hadza’s ancestors have probably lived in their current territory for tens of thousands of years. Hadzaland is about 50 kilometres (31 mi) from Olduvai Gorge, an area sometimes called the “Cradle of Mankind” because of the number of hominin fossils found there, and 40 kilometres (25 mi) from the prehistoric site of Laetoli. Archaeological evidence suggests that the area has been continuously occupied by hunter-gatherers much like the Hadza since at least the beginning of the Later Stone Age, 50,000 years ago. Although the Hadza do not make rock art today, they consider several rock art sites within their territory, probably at least 2,000 years old, to have been created by their ancestors, and their oral history does not suggest they moved to Hadzaland from elsewhere.” ref

“The Hadza population is dominated by haplogroup B2-M112 (Y-DNA). There are also Y-haplogroups haplogroup E-V38(Y-DNA) and haplogroup E-M215(Y-DNA). Until about 500 BCE, Tanzania was exclusively occupied by hunter-gatherers akin to the Hadza. The first agriculturalists to enter the region were Cushitic-speaking cattle herders from the Horn of Africa. Around 500 CE, the Bantu expansion reached Tanzania, bringing populations of farmers with iron tools and weapons. The last major ethnic group to enter the region were Nilotic pastoralists who migrated south from Sudan in the 18th century.” ref

“Each of these expansions of farming and herding peoples displaced earlier populations of hunter-gatherers, who were at a demographic and technological disadvantage and vulnerable to the loss of environmental resources (i.e., foraging areas and habitats for game) to farmland and pastures. Groups such as the Hadza and the Sandawe are remnants of indigenous hunter-gatherer populations that were once much more widespread, and they are under continued pressure from the expansion of agriculture into their traditional lands.” ref

“Farmers and herders appeared in the vicinity of Hadzaland relatively recently. The Isanzu, a Bantu farming people, began living south of Hadzaland around 1850. The pastoralist Iraqw and Datooga were both forced to migrate into the area by the expansion of the Maasai, the former in the 19th century and the latter in the 1910s. The Hadza also have direct contact with the Maasai and with the Sukuma west of Lake Eyasi. The upheavals caused by the Maasai expansion in the late 19th century caused a decline in the Hadza population.” ref

“The Hadza’s interaction with many of these peoples has been hostile. Pastoralists often killed Hadza as reprisals for the “theft” of livestock since the Hadza did not have the notion of animal ownership and would hunt them as they would wild game. The general attitude of neighboring agro-pastoralists towards the Hadza was prejudicial. They viewed them as backward, lacking a “real language,” and made up of the dispossessed of neighboring tribes that had fled into the forest out of poverty or because they committed a crime. Many of these misconceptions were transmitted to early colonial visitors to the region who wrote about the Hadza.” ref

“The Isanzu were hostile to the Hadza at times. Isanzu people may have captured them as part of the slave trade until as late as the 1870s when it was halted by the German colonial government. Later interactions were more peaceful, with the two peoples sometimes intermarrying and residing together, though as late as 1912, the Hadza were reported as being “ready for war” with the Isanzu. Still, folk tales depict the Isanzu as favorable and, at times, heroic, unlike the Iraqw and the cattle-raiding Maasai. Moreover, many goods and customs come from them, and the Hadza myths mention and depict a benevolent influence of the Isanzu in their mythology. The Sukuma and the Hadza had a more amicable relationship. The Sukuma drove their herds and salt caravans through Hadza lands and exchanged old metal tools, which the Hadza made into arrowheads, for the right to hunt elephants in Hadzaland.” ref

“The Hadza are organized into bands or ‘camps’ of 20–30 people. Camps of over a hundred may form during berry season. There is no tribal or other governing hierarchy, and almost all decisions are made by reaching an agreement through discussion. The Hadza trace descent bilaterally (through both paternal and maternal lines), and almost all Hadza people can trace some kin tie to all other Hadza people. Furthermore, the Hadza are egalitarian, so there are no real status differences between individuals. While the elderly receive slightly more respect, all individuals are equal to others of the same age and sex, and compared to strictly stratified societies, women are fairly equal to men. This egalitarianism results in high levels of freedom and self-dependence. When conflict arises, one of the parties involved may voluntarily move to another camp as resolution. Ernst Fehr and Urs Fischbacher point out that the Hadza people “exhibit a considerable amount of altruistic punishment” to organize these tribes. The Hadza live in a communal setting and engage in cooperative child rearing, where many people, both related and unrelated, provide high-quality child care.” ref

“The Hadza move camp for several reasons. Camps can split when individuals move to resolve conflicts. Camps can be abandoned when someone falls ill and dies, as any illness is associated with the place it was contracted. There is also seasonal migration between dry-season refuges, better hunting grounds when water is more abundant, and areas with large numbers of tubers or berry trees when they are in season. If a man kills a particularly large animal, such as a giraffe, far from home, a camp will temporarily relocate to the kill site. Shelters can be built in a few hours, and most of the possessions owned by an individual can be carried on their back.” ref

“The Hadza are predominantly monogamous, though there is no social enforcement of monogamy. After marriage, the husband and wife are free to live where they decide, which may be with the father or mother’s family. This marital residence pattern is called ambilocality and is common among foragers. Specifically among Hadza, there is a slightly higher frequency of married couples living with the mother’s kin than with the father’s kin. Men and women value traits such as hard work when evaluating partners. They also value physical attractiveness, and many of their preferences for attractiveness, such as symmetry, averageness and sexually dimorphic voice pitch,  are similar to preferences found in Western nations.” ref

“While men specialize in procuring meat, honey, and baobab fruit, women specialize in tubers, berries, and greens. This division of labor is relatively consistent, but women will occasionally gather a small animal or egg or collect honey, and men will occasionally bring a tuber or some berries back to camp. Hadza men usually forage individually. During the day, they usually feed themselves while foraging and bring home any additional honey, fruit, or wild game. Women forage in larger parties and usually bring home berries, baobab fruit, and tubers, depending on availability. Men and women also forage cooperatively for honey and fruit; at least one adult male will usually accompany a group of foraging women.” ref

“Women’s foraging technology includes digging sticks, grass baskets for carrying berries, large fabric or skin pouches for carrying items, knives, shoes, other clothing, and various small items held in a pouch around the neck. Men carry axes, bows, poisoned and non-poisoned arrows, knives, small honey pots, fire drills, shoes and apparel, and various small items. A myth depicts a woman harvesting the honey of wild bees, and at the same time, declares that the job of honey harvesting belongs to the men.” ref

“For harvesting honey or fruit from large trees such as the baobab, the Hadza beat pointed sticks into the trunk of the tree to use as ladders. This technique is depicted in a folk tale and documented on film. The Hadza do not follow a formal religion, and it has been claimed that they do not believe in an afterlife.  They offer prayers to Ishoko (the Sun) or to Haine (the moon) during hunts and believe they go to Ishoko when they die. They also hold rituals such as the monthly epeme dance for men at the new moon and the less frequent maitoko circumcision and coming-of-age ceremony for women.” ref

Epeme

“The Hadza people embrace epeme, which can be understood as their concept of manhood, hunting, and the relationship between sexes. “True” adult men are called epeme men, which they become by killing large game, usually in their early 20s. Being an epeme comes with an advantage: only epeme men are allowed to eat certain parts of large game animals, such as warthog, giraffe, buffalo, wildebeest, and lion. The parts of these animals that are typically considered epeme are the kidney, lung, heart, neck, tongue, and genitals. No one besides other epeme men are allowed to be present for the epeme meat-eating. If a man still has not killed a large game animal by his thirties, he will automatically be considered epeme and will be allowed to eat the epeme meat.” ref

“In addition to eating epeme meat, the epeme men participate in an epeme dance. In Jon Yates’s summary of Frank Marlowe’s account, this dance occurs every night when the moon isn’t visible, and must occur in near-complete darkness, with camp-fires extinguished. To begin the ritual, the women separate from the men and sit where they cannot be seen. The men gather behind a tree or hut and prepare for the dance. In the pitch dark, as the women begin to sing, the first man starts to dance. He wears a headdress of dark ostrich feathers, bells on one of his ankles, a rattle in his hand, and a long black cape on his back.” ref

“He stamps his right foot hard on the ground in time with the women’s singing, causing the bells to ring while marking the beat of the music with his rattle. He sings out to the women, who answer in a call and response. As the singing grows in strength, the women rise to join the man, who continues to dance—committing his efforts to a family member, one of the women, a friend, or one of his children. At this point, the child may join the dance as well. After each man has danced the epeme two or three times, the ritual is finished, by which time it is close to midnight. The ritual has been shown to promote social cohesion among the Hadza, and those who share the epeme dance show elevated levels of mutual trust and support.” ref 

Mythological figures with celestial connotations

“Ishoko and Haine are mythological figures who are believed to have arranged the world by rolling the sky and the earth like two sheets of leather and swapping their order to put the sky above us; in the past, the sky was under the earth. These figures are described as making crucial decisions about the animals and humans by choosing their food and environment, giving people access to fire, and creating the capability of sitting. These figures have celestial connotations: Ishoko is a solar figure, and Haine, her husband, is a lunar figure. Uttering Ishoko’s name can be a greeting or a good wish to someone for a successful hunt. The character “Ishoye” seems to be another name for Ishoko. She is depicted in some tales as creating animals, including people. Some of her creatures later turned out to be man-eating giants, disastrous for their fellow giants and people. Seeing the disaster, she killed these giants, saying, “You are not people any longer.” ref

Culture heroes

Indaya, the man who went to the Isanzu territory after his death and returned, plays the role of a culture hero: he introduces customs and goods to the Hadza. The Isanzu people neighbor the Hadza. They are regarded as peaceful, and the Hadza myths mention and depict this benevolent influence of the Isanzu in their mythology. This advantageous view of the Isanzu gives them a place as heroes in Hadza folklore. In some of the mythical stories about giants, it is an Isanzu man who liberates the Hadza from a malevolent giant.” ref

Stories about giants

“The Hadza have many stories about giants, which describe people with superhuman strength and size but otherwise with human weaknesses. They have human needs, eat and drink, and can be poisoned or cheated. One of the giants, Sengani (or Sengane), is depicted as Haine’s helper. As the story goes, Haine gave him the power to rule over the people. In Haine’s absence, the giant endangered people with his decisions. The people resisted his choices, so the giant ordered the lions to attack them. This surprised the humans, who had previously regarded lions as harmless. The people killed the giant in revenge.” ref

“This giant had brothers, rendered as “Ssaabo” and “Waonelakhi” in Kohl-Larsen. Several tales describe the disaster these giants caused by constantly killing and beating the Hadza. The Hadza had to ask for help from neighboring groups, and finally, the giants were tricked and either poisoned or shot to death by poison arrows. Another story tells of a man-eating giant, rendered as “!Esengego” by Kohl-Larsen. He and his family were killed by a benevolent snake, which turned out to be the remedy applied by the goddess Ishoko to liberate people. Ishoko changed the corpses of the giant family into leopards. She prohibited them from attacking people unless an arrow provoked or wounded them.” ref

“Another giant, rendered “!Hongongoschá” by Kohl-Larsen, appears as a different sort of mythological figure. He did not bother the Hadza much in his tales, only secretly stealing small things at night. His nourishment was the flowers of trees (and occasionally stolen vegetables). The people greeted him with great respect, and the giant wished them good luck in hunting. This changed when a boy deliberately injured the giant, and though he attempted to provide goodwill, !Hongongoschá took revenge by killing the boy. Finally, the god Haine determined a course of justice: he warned the people, revealed the boy’s malevolent deed, and changed the giant into a big white clam.” ref

Humanity’s forgotten return to Africa revealed in DNA

“Call it humanity’s unexpected U-turn. One of the biggest events in the history of our species is the exodus out of Africa some 65,000 years ago, the start of Homo sapiens‘ long march across the world. Now a study of southern African genes shows that, unexpectedly, another migration took western Eurasian DNA back to the very southern tip of the continent 3000 years ago. According to conventional thinking, the Khoisan tribes of southern Africa, have lived in near-isolation from the rest of humanity for thousands of years. In fact, the study shows that some of their DNA matches most closely people from modern-day southern Europe, including Spain and Italy.” ref

“Because Eurasian people also carry traces of Neanderthal DNA, the finding also shows – for the first time – that genetic material from our extinct cousin may be widespread in African populations. The Khoisan tribes of southern Africa are hunter-gatherers and pastoralists who speak unique click languages. Their extraordinarily diverse gene pool split from everyone else’s before the African exodus. The unexpected snippets of DNA most resembled sequences from southern Europeans, including Sardinians, Italians and people from the Basque region (see “Back to Africa – but from where?“). Dating methods suggested they made their way into the Khoisan DNA sometime between 900 and 1800 years ago – well before known European contact with southern Africa.” ref

“Archaeological and linguistic studies of the region can make sense of the discovery. They suggest that a subset of the Khoisan, known as the Khoe-Kwadi speakers, arrived in southern Africa from east Africa around 2200 years ago. Khoe-Kwadi speakers were – and remain – pastoralists who make their living from herding cows and sheep. The suggestion is that they introduced herding to a region that was otherwise dominated by hunter-gatherers.” ref

Khoe-Kwadi tribes

“Reich and his team found that the proportion of Eurasian DNA was highest in Khoe-Kwadi tribes, who have up to 14 per cent of western Eurasian ancestry. What is more, when they looked at the east African tribes from which the Khoe-Kwadi descended, they found a much stronger proportion of Eurasian DNA – up to 50 per cent. That result confirms a 2012 study by Luca Pagani of the Wellcome Trust Sanger Institute in Hinxton, UK, which found non-African genes in people living in Ethiopia. Both the 2012 study and this week’s new results show that the Eurasian genes made their way into east African genomes around 3000 years ago. About a millennium later, the ancestors of the Khoe-Kwadi headed south, carrying a weaker signal of the Eurasian DNA into southern Africa.” ref

“The cultural implications are complex and potentially uncomfortably close to European colonial themes. “I actually am not sure there’s any population that doesn’t have west Eurasian [DNA],” says Reich. “These populations were always thought to be pristine hunter-gatherers who had not interacted with anyone for millennia,” says Reich’s collaborator, linguist Brigitte Pakendorf of the University of Lyon in France. “Well, no. Just like the rest of the world, Africa had population movements too. There was simply no writing, no Romans or Greeks to document it.” There’s one more twist to the tale. In 2010 a research team – including Reich – published the first draft genome of a Neanderthal. Comparisons with living humans revealed traces of Neanderthal DNA in all humans with one notable exception: sub-Saharan peoples like the Yoruba and Khoisan.” ref

“That made sense. After early humans migrated out of Africa around 60,000 years ago, they bumped into Neanderthals somewhere in what is now the Middle East. Some got rather cosy with each other. As their descendants spread across the world to Europe, Asia and eventually the Americas, they spread bits of Neanderthal DNA along with their own genes. But because those descendants did not move back into Africa until historical times, most of this continent remained a Neanderthal DNA-free zone. Or so it seemed at the time. Now it appears that the Back to Africa migration 3000 years ago carried a weak Neanderthal genetic signal deep into the homeland. Indeed one of Reich’s analyses, published last month, found Neanderthal traces in Yoruba DNA.” ref 

Y-Chromosome Variation in Southern African Khoe-San Populations Based on Whole-Genome Sequences 

“Abstract: Although the human Y chromosome has effectively shown utility in uncovering facets of human evolution and population histories, the ascertainment bias present in early Y-chromosome variant data sets limited the accuracy of diversity and TMRCA estimates obtained from them. The advent of next-generation sequencing, however, has removed this bias and allowed for the discovery of thousands of new variants for use in improving the Y-chromosome phylogeny and computing estimates that are more accurate. Here, we describe the high-coverage sequencing of the whole Y chromosome in a data set of 19 male Khoe-San individuals in comparison with existing whole Y-chromosome sequence data. Due to the increased resolution, we potentially resolve the source of haplogroup B-P70 in the Khoe-San, and reconcile recently published haplogroup A-M51 data with the most recent version of the ISOGG Y-chromosome phylogeny. Our results also improve the positioning of tentatively placed new branches of the ISOGG Y-chromosome phylogeny. The distribution of major Y-chromosome haplogroups in the Khoe-San and other African groups coincide with the emerging picture of African demographic history; with E-M2 linked to the agriculturalist Bantu expansion, E-M35 linked to pastoralist eastern African migrations, B-M112 linked to earlier east-south gene flow, A-M14 linked to shared ancestry with central African rainforest hunter-gatherers, and A-M51 potentially unique to the Khoe-San.” ref

“The male-specific portion of the Y chromosome (MSY) has long been regarded as an effective tool in the study of human evolutionary history. It has proved useful mainly due to a lack of recombination along its length, making it the longest haplotypic block in the human genome; and its paternal mode of inheritance. The transmission of an intact haplotype from father to son, changing only through mutation, preserves a simpler record of its history and allows us to study the male contribution to the shaping of humanity.” ref

“The mutations found on Y chromosomes sourced from numerous human populations have been used to generate Y-chromosome phylogenies, with well-defined and geographically informative haplogroups, that is, groups of haplotypes that share common ancestors and so present as clades in a phylogeny. The utility of the Y-chromosome phylogeny notwithstanding, the ascertainment bias present when initially sourcing the mutations used to build it, resulted in limitations. The use of predefined sets of variants, limited the ability to generate unbiased estimates of global diversity and accurate estimates of the time to most recent common ancestor (TMRCA). As a solution, short tandem repeat polymorphisms (STRs) have been used for a long time, however, STRs had their own biases and issues. The emergence of next-generation sequencing (NGS) technology, resulted in the discovery of thousands of unbiased variants, which allowed for substantially more accurate estimates of the age the Y-chromosome phylogeny and its haplogroups.” ref

“Although studies usually attempted to balance the global representation of populations in their samples, there did appear to be an underrepresentation of African samples in the final results, with this being especially true of haplogroups A and B. Barbieri et al. (2016) addressed this discrepancy by sequencing a portion of the Y chromosome in 547 Khoe-San- and Bantu-speakers; resulting in much older estimates of the ages of haplogroups A and B and their subclades. Following high-coverage sequencing of the full Y chromosome in a data set of 19 male Khoe-San individuals and comparison with existing full Y-chromosome sequence data, the present study describes the distribution of haplogroups found, in the wider African context. The high level of resolution afforded by a sequencing analysis allowed us to potentially resolve the source of haplogroup B-P70 in the Khoe-San. The use of two slightly differing phylogenies when assigning our haplogroups, allowed for the reconciliation of the haplogroup A-M51 data from Barbieri et al. (2016) with the most recent version of the ISOGG Y-chromosome phylogeny.” ref

Results

“Researchers performed high-coverage whole-genome sequencing of 25 Khoe-San individuals from five different populations. In this study, we discuss the results of the Y-chromosome sequences of the 19 male individuals that were included in the study. After processing and filtering the sequence data (see Materials and Methods), we obtained 5,783 variants; with an average depth of 31×. Once merged with the comparative data from seven additional populations, our total data set contained 7,878 variants from 8.8 Mb (8,800,463 bases) of Y-chromosome sequence in 48 individuals.” ref

Khoe-San Haplogroups

“The major haplogroups found in our sequenced individuals were A-M14 (A1b1a1), A-M51 (A1b1b2a), B-M112 (B2b), E-M2 (E1b1a1), and E-M35 (E1b1b1); and were thus strongly concordant with previous surveys of Y-chromosome variation in Khoe-San populations. The additional resolution provided by sequencing, however, uncovered several new variants especially supporting branches in haplogroups A and B, and allowed for greater clarity regarding the relationships of some subclades and markers. Haplogroup A1b1b2a (A-M51) has often been found to be the most common haplogroup A subclade in southern Africa, though usually found primarily in the Khoe-San or in populations with significant levels of Khoe-San admixture.” ref

“Three previously reported subclades, haplogroups A1b1b2a1a (A-P71), A1b1b2a2 (A-V37), and A1b1b2a1b (A-V306), were found in our data set; and the branching structure was further refined and reconciled with the ISOGG Y phylogeny (ISOGG 2019-2020). Haplogroup A1b1b2a2 is the first to branch off, whereas haplogroups A1b1b2a1a and A1b1b2a1b are united by marker M239. Within haplogroup A1b1b2a1a, all three individuals were also derived at marker P102, with one of them ancestral at marker P291. This is at odds with the current ISOGG phylogeny, which has P291 basal to P102. The three subclades segregated independently among the populations, with A1b1b2a2 found in two !Xun individuals, A1b1b2a1a in three Nama individuals and A1b1b2a1b in two Ju|’hoansi individuals.” ref

Gene Flow into Khoe-San Populations

“Haplogroup E1b1b1b2b2a1 (E-M293) was found in two Khoe-San: one Nama and one !Xun. This E-M35 subclade was previously linked to the movement of pastoralist groups from eastern Africa to southern Africa ∼2,000 years ago. Moreover, the Y-chromosome from the Nama individual belonged to the basal E1b1b1b2b2a1 clade, whereas the !Xun individual belonged to a subclade further derived for markers CTS2104, CTS2297, CTS2553, and Y17343, which were also found in two Sandawe individuals. The two haplogroup B2b (B-M112) chromosomes, found within one Ju|’hoansi and one !Xun, fell within a subclade of haplogroup B2b1a2b2∼ (B-M7592), defined by marker M7591. The closest related Y chromosome belonged to a sister clade within haplogroup B2b1a2b2∼ (B2b1a2b2a∼) and was from a Hadza individual (. Although these Khoe-San B2b chromosomes are known to be derived at marker P8 (unpublished data), this marker has since been removed from recent versions of the ISOGG Y-chromosome phylogeny. The finding that marker M7591 is also derived and appears to be phylogenetically equivalent to markers P70 and P8, allows for the reintroduction of the lineage to the phylogeny.” ref

“The E1b1a1 (E-M2) haplogroup occurs in individuals from many populations. Khoe-San Y chromosomes within haplogroup E1b1a1 are likely to have been introduced by surrounding Bantu-speaker populations. The rapid expansion of E1b1a1 across sub-Saharan Africa makes it difficult to pinpoint its exact sources into the Khoe-San without taking into account historical data.  Within haplogroup E, E1b1b1 was found to have the highest mean branch length; though this may have been due to a lower sample size compared with haplogroup E1b1a.” ref

“The associations of the Khoe-San with other populations in the same major haplogroups also gave some indication of historical interactions and shared ancestry. While haplogroups A1b1a1, A1b1b2a, B2b, and E1b1b1 have long been associated with Khoe-San populations, at this point, it appears haplogroup A1b1b2a may be the only surviving haplogroup in the Khoe-San to have originated autochthonously. The presence of haplogroup A1b1a1 in the Khoe-San in this study and others has been characterized primarily by the more terminal lineages of the haplogroup and lineages ancestral to these have been found in central Africa.” ref

“The majority of haplogroup B2b lineages found so far in Khoe-San populations have fallen into the subclades B-P6 and B-P70, whereas other B2b lineages have been found in central and eastern Africa. Previously, Batini et al. (2011) linked the presence of B-P70 in the Khoe-San to possible gene flow from central African Rainforest hunter-gatherer populations. This was based on the presence and diversity of haplogroup B-P7 (the ancestral lineage to B-P70) in the Rainforest hunter-gatherer populations and low levels of it elsewhere. Our findings, however, place the Khoe-San-specific lineage within a clade (B2b1a2b2∼) together with a Hadza lineage. The TMRCA of haplogroup B2b1a2b2∼ has been estimated to ∼21,000–26,000 years ago.” ref

“Given this relatively deep age, we cannot rule out the possibility that B2b1a2b2∼ was also present in central African Rainforest hunter-gatherer populations, and entered the Khoe-San population. However, our findings together with multiple lines of evidence pointing to gene flow from eastern Africa into southern Africa indicate that B2b1a2b2∼ in the Khoe-San may have come from eastern Africa. Notably, this B2b1a2b2∼ subclade has mainly been found in San populations such as the Ju|’hoansi and !Xun, and not in Khoekhoe herder populations such as the Nama. This is in contrast to the E1b1b1b2b2a1 haplogroup, which is found quite commonly in Nama individuals.” ref

“As the E1b1b1b2b2a1 haplogroup is a marker of the movement of pastoralism from eastern Africa to southern Africa, this would indicate that the arrival of B2b1a2b2∼ in southern Africa was separate from the arrival of pastoralism. Evidence of a gradient of relatedness among eastern African and southern African hunter-gatherers; and while most of the eastern African genomic component present in southern African hunter-gatherers was attributed to the arrival of pastoralism, it appeared that the ancient southern African individuals who did not show the strong signal of eastern African admixture still shared more alleles with eastern Africans than with western Africans; possibly indicating some level of isolation-by-distance. Evidence of gene flow between the Hadza and the Ju|’hoansi, a population shown to have been affected minimally by the recent gene flow from eastern Africa, has also been noted (Schlebusch et al. 2012). These factors may indicate that the presence of haplogroup B2b1a2b2∼ in southern Africa may predate the arrival of pastoralism.” ref 

Genetic substructure and complex demographic history of South African Bantu speakers

“Abstract: South Eastern Bantu-speaking (SEB) groups constitute more than 80% of the population in South Africa. Despite clear linguistic and geographic diversity, the genetic differences between these groups have not been systematically investigated. Based on genome-wide data of over 5000 individuals, representing eight major SEB groups, we provide strong evidence for fine-scale population structure that broadly aligns with geographic distribution and is also congruent with linguistic phylogeny (separation of Nguni, Sotho-Tswana and Tsonga speakers). Although differential Khoe-San admixture plays a key role, the structure persists after Khoe-San ancestry-masking. The timing of admixture, levels of sex-biased gene flow and population size dynamics also highlight differences in the demographic histories of individual groups. The comparisons with five Iron Age farmer genomes further support genetic continuity over ~400 years in certain regions of the country. Simulated trait genome-wide association studies further show that the observed population structure could have major implications for biomedical genomics research in South Africa.” ref

“The archaeological record and rock art evidence trace the presence of San-like hunter-gatherer culture in Southern Africa to at least 20,000–40,000 years ago. Three sets of migration events have dramatically reshaped the genetic landscape of this geographic region in the last two millennia. The first of these was a relatively small scale migration of East African pastoralists, who introduced pastoralism to Southern Africa ~2,000 years ago. This population was subsequently assimilated by local Southern African San hunter-gatherer groups, forming a new population that was ancestral to the Khoekhoe herder populations. Today, Southern African Khoe and San populations collectively refer to hunter-gatherer (San) and herder (Khoekhoe) communities. While Khoe-San groups are distributed over a large geographic area today (spanning the Northern Cape Province of South Africa, large parts of Namibia, Botswana, and Southern Angola), these groups are scattered, small, and marginalized.” ref

“The introduction of pastoralism in the region was closely followed by the arrival of the second set of migrants i.e., the Bantu-speaking (BS) agro-pastoralists. The archaeological record suggests that ancestors of the current-day BS populations undertook different waves of migration instead of a single large-scale movement. The earliest communities spread along the East coast to reach the KwaZulu-Natal South coast by the mid-fifth century CE while the final major episode of settlement is estimated to be around CE 1350. These archaeologically distinct groups gradually spread across present-day South Africa, interacting to various degrees with the Khoe-San groups, eventually giving rise to South Africa’s diverse Bantu-speaking communities. The third major movement into Southern Africa was during the colonial era in the last four centuries when European colonists settled the area. During this period slave trade introduced additional intercontinental gene flow giving rise to complex genomic admixture patterns in current-day Southern African populations.” ref

“South Africa has 11 official languages of which nine are Bantu languages belonging to this family’s South-Eastern branch. Within these nine languages two large subclusters are traditionally distinguished: Nguni (including Zulu, Xhosa, Swazi, and Ndebele) and Sotho-Tswana (including Sotho, Tswana, and Pedi). Venda and Tsonga tend to be seen as independent linguistic entities. A new lexicon-based linguistic phylogeny included in this study broadly confirms the traditionally recognized clusters, but also adds possible insights into how these languages might relate to each other as well as to 60 other Bantu languages.” ref

“While the genetic diversity of Khoe-San and mixed ancestry groups has been widely investigated, the genetic diversity of the SEB-speaking (referred henceforth as SEB) groups has not been systematically investigated. One of the very early studies based on the Y-chromosome and a few autosomal markers, which included almost all the main SEB groups and covered most of the provinces from South Africa, indicated the possibility of genetic structure within the SEB populations. However, many subsequent studies using genome-wide datasets did not investigate genetic differentiation or population structure within SEB groups, which consequently led to its consideration as a group without clear internal substructure. Moreover, studies including multiple SEB groups were often limited in terms of sample size or SEB group diversity.” ref

“The availability of Iron-Age genomes from Southern Africa provided us with the unique opportunity to compare the affinities of present-day SEB groups to populations living in these areas centuries ago. The PCA and PCA-UMAP projecting five Iron-Age Bantu-related genomes (300–700 years old) onto the genetic variation of present-day SEB individuals show these genomes to be on a temporal cline with the older genomes (Pemba, Eland Cave and Newcastle; ranging ~700–450 years ago) appearing closer to the Tsonga and Venda, while more recent genomes (Champagne Castle and Mfongosi; ranging from ~448–300 years ago) occurring closer to the Nguni-speakers. This cline of the Iron-Age genomes also aligns with geographic distribution from North to South, as well as increasing levels of Khoe-San ancestry in them. More ancient genomes from Southern Africa would be required to test whether the trends observed in these Iron-Age genomes are indicative of phases in the movement of groups further south with time, a process marked by concomitant increase of Khoe-San ancestry in the migrants. Interestingly, the wider geographic region of Northern KwaZulu-Natal around Champagne Castle in Central-East South Africa, where the youngest of these Iron-Age genomes was collected, is still dominated by Nguni-speakers, providing support for at least four centuries of genetic continuity in certain regions of South Africa.” ref

Evolutionary Genetics and Admixture in African Populations

“Abstract: As the ancestral homeland of our species, Africa contains elevated levels of genetic diversity and substantial population structure. Importantly, African genomes are heterogeneous: They contain mixtures of multiple ancestries, each of which have experienced different evolutionary histories. In this review, we view population genetics through the lens of admixture, highlighting how multiple demographic events have shaped African genomes. Each of these historical vignettes paints a recurring picture of population divergence followed by secondary contact. First, we give a brief overview of genetic variation in Africa and examine deep population structure within Africa, including the evidence of ancient introgression from archaic “ghost” populations. Second, we describe the genetic legacies of admixture events that have occurred during the past 10,000 years. This includes gene flow between different click-speaking Khoe-San populations, the stepwise spread of pastoralism from eastern to southern Africa, multiple migrations of Bantu speakers across the continent, as well as admixture from the Middle East and Europe into the Sahel region and North Africa. Furthermore, the genomic signatures of more recent admixture can be found in the Cape Peninsula and throughout the African diaspora. Third, we highlight how natural selection has shaped patterns of genetic variation across the continent, noting that gene flow provides a potent source of adaptive variation and that selective pressures vary across Africa. Finally, we explore the biomedical implications of population structure in Africa on health and disease and call for more ethically conducted studies of genetic variation in Africa.” ref

“Compared with the rest of the world, each African genome harbors ∼25% more polymorphisms than each non-African genome. Furthermore, variants that are rare on a global level (<1% frequency) are more frequently found to be common in African populations, that is, there is an excess of variants exclusively found in Africans. Greater numbers of private African alleles are consistent with the out-of-Africa (OOA) model, as substantial numbers of polymorphisms were lost due to serial founder effects. In fact, the genetic variation found outside of Africa is largely a subset of African genetic diversity. Subsets of African genetic variation found outside of Africa also vary by region, indicating that multiple OOA migrations may have occurred.” ref

“Additionally, African populations exhibit a faster decay of LD, leading to shorter haplotypes. This is of biomedical relevance (see below), and it also enables improved fine-mapping of causal variants in genome-wide association studies (GWAS) because casual variants are tagged by fewer other variants. In line with the OOA model, many human populations experienced a major decline in N_e coinciding with the OOA migration 70,000–50,000 years ago. Concurrently, African populations experienced declines in N_e while maintaining consistently larger N_e than non-African populations. Thus, the higher genetic diversity and lower LD in African populations reflect historically larger N_e.” ref

“Hunting and gathering was the predominant subsistence strategy prior to the introduction of agriculture and pastoralism during the Neolithic (i.e., 12,000–6,500 years ago in Africa). Today, only a few traditional hunter–gatherer groups remain that live in small communities. Generally, it is assumed that they have either merged into or were replaced by neighboring agropastoral groups, obscuring some of the ancestral genetic variation and structure. Nevertheless, when accounting for recent admixture, studying the genetics of the traditional hunter–gatherer groups in Africa can provide a snapshot of deep population structure due to their long-term population continuity. Attempts to illuminate the deep population structure in Africa have been further aided by the emergence of ancient DNA from unadmixed hunter–gatherer individuals.” ref

“The remaining traditional hunter–gatherer groups in Africa can be broadly grouped into three major groups: Khoe-San, eastern African hunter–gatherers (EAHG), and rainforest hunter–gatherers (RHG). Khoe-San collectively refers to Khoisan-speaking San hunter–gatherers and Khoekhoe herders, who historically inhabit arid regions in southern Africa. Possibly, Khoe-San were the only inhabitants of southern Africa for much of its prehistory. Khoekhoe herders have adopted a pastoralist lifestyle only recently, likely after admixture with eastern African pastoralists over the past 1,500 years. Similarly, EAHG groups, for example, the click-speaking Hadza and Sandawe in Tanzania and the Chabu in Ethiopia, are traditional foragers, who have practiced a hunter–gatherer lifestyle until recently or are still practicing it. These EAHG groups are more closely related to each other than to other African hunter–gatherer groups. RHG groups comprise genetically diverse populations in equatorial Africa, which are often further subdivided into western (e.g., the Baka) and eastern (e.g., the Mbuti) RHG groups.” ref

“Many hunter–gatherer groups experienced declines in N_e during the Holocene and have small census population sizes today. Nevertheless, African hunter–gatherers have the highest level of genetic diversity of extant populations and represent the most deeply branching human lineages even after accounting for recent admixture. Specifically, the Khoe-San exhibit the highest genetic diversity of all human lineages, with a mean heterozygosity of 1.154 × 10⁻³ compared with 1.09 × 10⁻³ in the Mandenka. As their genetic diversity is still significantly higher after accounting for recent admixture with non–Khoe-San groups, it reflects their historically larger N_e. The lineage leading to the Khoe-San is basal to all other human lineages with an estimated divergence time of 300–200 kya (e.g., the Ju|’Hoan with the lowest level of recent admixture diverged ∼270,000 years ago). Subsequently, the Mbuti (RHG) diverged ∼220,000 years ago from all other human lineages, forming a second basal lineage. These estimates may reflect lower bounds as recent admixture reduces divergence time estimates. For these reasons, assuming regional population continuity, it has been argued for a southern African origin of modern humans, although models involving eastern Africa and/or multiple geographic regions are also debated.” ref

“The genetic relationship between these different hunter–gatherer groups can largely be modeled by an isolation-by-distance model. There appears to be a genetic cline connecting the eastern African Hadza and southern African Khoe-San, as ancient hunter–gatherer genomes from eastern Africa show affinities to extant southern African San and EAHG. For instance, ancient hunter–gatherers genomes from Malawi (∼8,100–2,500 years ago) and Tanzania (∼1,400 years ago) exhibited two-third and one-third San-related ancestry, respectively, suggesting that the San previously occupied a larger geographic area extending into eastern Africa or admixture with a hunter–gatherer group who later gave rise to contemporary San. An additional east–southwest cline was recently identified by the incorporation of six novel genomes of ancient hunter–gatherers from eastern and south–central Africa. Some of these individuals are located closer to ancient and present-day central African RHG in principal component space. This could either suggest deep population structure with EAHG and southern hunter–gatherer groups tracing some of their ancestries to a basal central African RHG lineage or gene flow between southern African and central African foragers, as indicated by a distinct allele-sharing pattern between the !Xun/Ju|’Hoan and Mbuti. However, in general, ancient genomes reveal deep divergence times of eastern, southern, and central African hunter–gatherer groups, indicating little historical gene flow.” ref

“Although recent admixture with agriculturists and pastoralists partially obscures ancestral variation and population structure in traditionally foraging groups, their genomes may still provide exciting glimpses into the deep demographic history of modern humans. The sequencing of more ancient African genomes will likely reveal new complexities of human origins, although the tropical climate is complicating the analysis of ancient DNA in sub-Saharan Africa. Nevertheless, ancient DNA has recently been obtained of 18,000-year-old individuals.” ref

Evidence of Archaic (Ghost) Introgression in Africa

“With the sequencing of genomes of archaic hominins, it has become evident that modern humans interbred with archaic hominins on multiple occasions in Eurasia. Although modern human–Neanderthal interbreeding most likely occurred in Eurasia after the OOA migration (possibly in the Levant), African populations also exhibit signals of Neanderthal admixture—especially northeastern African but also some West African populations. This signal of Neanderthal admixture observed in African genomes is most likely not the result of direct admixture but rather the result of admixture with back-migrating Europeans. This is because most Neanderthal haplotypes are shared with Europeans. However, evidence supporting additional admixture events with unknown archaic hominins—the so-called archaic “ghost” populations—within Africa is also mounting.” ref

“The first evidence for archaic ghost introgression in Africa was obtained by applying S*—an approach that searches for highly divergent haplotypes—to African populations. The time to the most recent common ancestor (TMRCA) of such identified putatively introgressed haplotypes was found to be significantly older than the deepest split of all modern human lineages and similar to the TMRCA of introgressed Neanderthal haplotypes found in Eurasian populations. This suggests that the introgressing archaic ghost lineage diverged approximately at the same time from the modern human lineage as Neanderthals.” ref

“Later studies fitting demographic models to the data or comparing empirical data to simulated data found that models which include archaic admixture in Africa consistently describe the data better than models that do not include archaic admixture. Reassuringly, the different approaches also inferred similar demographic scenarios, involving an archaic lineage that diverged around the same time as the Neanderthal lineage (∼800,000–500,000 years ago) and recurring, low-level admixture as recently as 30,000 years ago. Lorente-Galdos et al. estimated that Khoe-San derive 3.8% (95% CI: 1.7–4.8%), Mbuti 3.9% (95% CI: 1.3–4.9%), and western African populations 5.8% (95% CI: 0.7–9.7%) of their ancestry from an archaic ghost lineage.” ref

“Furthermore, several candidate introgressed genes have been identified. Xu et al. (2017) concluded that a highly divergent haplotype of MUC7 introgressed into modern West Africans from an archaic lineage. This salivary protein has previously been associated with being protective against asthma. However, Durvasula and Sankararaman (2020) did not find evidence for introgression at the MUC7 locus when they applied a novel statistical method (ArchIE) that identifies introgressed segments based on multiple population genetics statistics to western African genomes. Using ArchIE, they identified a set of possibly adaptively introgressed genes that are at high frequencies in West Africans (99.9th percentile of putatively introgressed allele frequencies): NF1, MTFR2, HSD17B2, KCN1P4, and TRPS1.” ref

“Despite the evidence for archaic admixture, it cannot be ruled out that deep population structure confounds the inference of archaic ghost introgression in Africa (Ragsdale et al. 2022; Fan et al. 2023). For instance, Ragsdale et al. (2022) recently found that a structured model with two stems, that is, two weakly differentiated Homo populations connected by gene flow over evolutionary time, can also explain the observed signals of archaic ghost introgression in Africa. However, the possibility of archaic ghost admixture is also supported by fossil records from across Africa, indicating that modern humans spatially and temporally overlapped with hominins exhibiting archaic features. Thus, there were ample opportunities for admixture between modern humans and archaic hominins.” ref

Pervasive Admixture in Africa during the Past 10,000 Years

“In conjunction with archeological and linguistic studies, genetic studies of contemporary humans and ancient remains have painted a complex pattern of human history in Africa, as many African populations are connected by gene flow. Most contemporary African groups share some of their ancestries with groups from different geographic regions. Nevertheless, different genetic ancestries tend to cluster geographically, with deserts and rainforests acting as major barriers to gene flow. In the following subsections, we discuss major migration events that have shaped population structure in Africa during the past 10,000 years. We start with discussing admixture events in the deeper past and move to admixture events closer to the present day.” ref

“The Khoe-San are basal to all other human lineages with an estimated divergence time of 300,000–200,000 years ago. Although these populations are traditional foragers, some Khoe-San groups have recently adopted (agro-)pastoralist lifestyle. Initial studies leveraging autosomal genotyping data and mitochondrial DNA (mtDNA) (Barbieri et al. 2014) suggested differentiation between Khoe-San populations living north and south of the Kalahari Desert, an area that was dominated by lake Makgadikgadi during prehistoric times (i.e., > 10,000 years ago). An additional central Khoe-San–related ancestry component has been identified in more recent studies that leveraged bigger and more diverse data sets.” ref

“Notably, these three ancestry components correlate with geography but not linguistics or present-day subsistence strategy. The Kx`a-speaking Ju|’Hoan and !Xun and the Khoekhoe-speaking Hai||om are representative of the North Khoe-San ancestry component, the Khoekhoe-speaking Nama and Tuu-speaking ‡Khomani and Karretije are representative of the South Khoe-San ancestry component, and all remaining Khoe-San population are representative of the central Khoe-San ancestry component. Interestingly, the pairwise genetic divergences of these three components were found to be similar (i.e., similar fixation index [F_ST] values), and the divergence time was estimated to be ∼25,000 years ago (95% CI: 18,000–32,000 years ago).” ref

“Although most of the genetic variation among Khoe-San populations is explained under an isolation-by-distance model, there is evidence of modest admixture between the three Khoe-San–related ancestry components. In formal admixture tests (f₃–analysis), the ‡Khomani (southern component) showed significant evidence of admixture with Taa populations (central), and the Ju|’Hoan (northern) showed significant signs of admixture with the !Xun (northern) and the Naro (central). Additionally, the Naro (central) showed evidence of admixture with the Ju|’Hoan (northern) and another population characterized by the Central Khoe-San component (e.g., Taa or |Gui).” ref

“However, none of the populations characterized by the central Khoe-San component showed significant evidence of being a mixture between northern and southern Khoe-San groups. Using SpaceMix analyses, found additional evidence for gene flow from the Ju|’Hoan (northern) into the ‡Hoan (central), from the |Gui/Xade San (central) into the Naro (central), and from an undefined Khoe-San population into the Nama (southern). Note that these tests do not definitively establish admixture between specific populations—the actual historical gene flow may have involved other related populations. It has been argued that this gene flow must have occurred within the last 10,000 years ago after the prehistoric lake Makgadikgadi dried up.” ref

Complex Spread of Pastoralism in Eastern and Southern Africa

“Recent genetic studies paint a complex picture of population continuity and admixture in eastern Africa since the introduction of pastoralism in northeastern Africa some 8,000 years ago. Using DNA from ancient individuals from Kenya and Tanzania, it has been proposed that herding and farming spread in multiple steps into eastern Africa. First, in northeastern Africa, admixture between a population related to contemporary Nilo-Saharan speakers (e.g., the Dinka or Nuer) and a population related to modern groups from northern Africa or the Levant created a group of “early northeastern pastoralists.” This group then migrated to eastern Africa and admixed with local foragers ∼4,000 years ago, receiving ∼20% ancestry from a group related to a 4,500-year-old ancient individual from the Mota cave in Ethiopia that is genetically similar to the isolated, Afro-Asiatic–speaking Aari and present-day Afro-Asiatic speakers.” ref

“Given the high genetic affinity of a pastoralist individual who lived 4000 years ago in northern Sudan with ancient individuals from Kenya and Tanzania, it has been argued that this initial dispersal of northeastern pastoralists into East Africa occurred rapidly. Lastly, this group received another pulse of gene flow from a population related to Nilo-Saharan–speaking Dinka in Sudan ∼2,200 years ago, that is, during the Iron Age. Based on varying amounts of Mota-related and Dinka-related ancestry in ancient individuals from the Democratic Republic of Congo, Uganda, and Botswana, it has been argued that a model with repeated, unidirectional gene flow from east African forager groups and Nilo-Saharan–speaking groups into the “early northeastern pastoralist” group provides a better fit. However, with the currently available data, it is impossible to distinguish between multiple waves of migration and complex population structure.” ref 

Mbuti people

“The Mbuti people, or Bambuti, are one of several indigenous pygmy groups in the Congo region of Africa. Their languages are Central Sudanic languages and Bantu languages. Bambuti are pygmy hunter-gatherers, and are one of the oldest indigenous people of the Congo region of Africa. The Bambuti are composed of bands which are relatively small in size, ranging from 15 to 60 people. The Bambuti population totals about 30,000 to 40,000 people. Many Batwa in various parts of the Democratic Republic of the Congo (DRC) also call themselves Bambuti. The Bambuti are primarily hunter-gatherers.” ref

“The Bambuti have an elaborate system of food allowances and restrictions; foods that are restricted are called kweri. Food sources yielded by the forest are non-kweri animals for meat consumption, root plants, palm trees, and bananas; and in some seasons, wild honey. Yams, legumes, beans, peanuts, hibiscus, amaranth, and gourds are consumed. The Bambuti use large nets, traps, and bows and arrows to hunt game. Women and children sometimes assist in the hunt by driving the prey into the nets. Both sexes gather and forage. Each band has its own hunting ground, although boundaries are hard to maintain.  The Mbuti call the forest “mother” and “father” as the mood seizes them, because, like their parents, the forest gives them food, shelter, and clothing, which are readily made from abundant forest materials.” ref

“The Bantu villagers produce many items that the hunter-gatherers trade some of their products for. They often obtain iron goods, pots, wooden goods, and basketry, in exchange for meat, animal hides, and other forest goods. Bushmeat is a particularly frequently traded item. They will also trade to obtain agricultural products from the villagers through barter. Hunting is usually done in groups, with men, women, and children all aiding in the process. Women and children are not involved if the hunting involves the use of a bow and arrow, but if nets are used, it is common for everyone to participate. In some instances, women may hunt using a net more often than men. The women and the children herd the animals to the net, while the men guard the net. Everyone engages in foraging, and women and men both take care of the children.” ref

“Women are in charge of cooking, cleaning and repairing the hut, and obtaining water. The kin-based units work together to provide food and care for the young. It is easier for men to lift the women into the trees for honey. According to a study published in 1987, based on fieldwork and data gathered between 1974 and 1985, the Mbuti restrict some 40% of the over 500 species of plants and animals they gather and hunt, including some 85% of the animals. The kweri animals are thought to cause disease and disorder, especially to young children; restrictions are gradually relaxed as one ages. The Bambuti tend to follow a patrilineal descent system, and their residences after marriage are patrilocal. However, the system is rather loose. The only type of group seen amongst the Bambuti is the nuclear family. Kinship also provides allies for each group of people.” ref

“Sister exchange is the common form of marriage. Based on reciprocal exchange, men from other bands exchange sisters or other females to whom they have ties. In Bambuti society, bride wealth is not customary. There is no formal marriage ceremony: a couple are considered officially married when the groom presents his bride’s parents with an antelope he alone has hunted and killed. Polygamy does occur, but at different rates depending on the group, and it is not very common. The sexual intercourse of married couples is regarded as an act entirely different from that of unmarried partners, for only in marriage may children be conceived.” ref

“Bambuti societies have no ruling group or lineage, no overlying political organization, and little social structure. The Bambuti are an egalitarian society in which the band is the highest form of social organization. Leadership may be displayed for example on hunting treks. Men become leaders because they are good hunters. Owing to their superior hunting ability, leaders eat more meat and fat and fewer carbohydrates than other men. Men and women basically have equal power. Issues are discussed and decisions are made by consensus at fire camps; men and women engage in the conversations equivalently. If there is a disagreement, misdemeanor, or offense, then the offender may be banished, beaten, or scorned. In more recent times the practice is to remove the offender from the forest and have them work for private landowners for little to no pay.” ref

“Everything in the Bambuti life is centered on the forest. They consider the forest to be their great protector and provider and believe that it is a sacred place. They sometimes call the forest “mother” or “father”. An important ritual that impacts the Bambuti’s life is referred to as molimo. After events such as death of an important person in the tribe, molimo is noisily celebrated to wake the forest, in the belief that if bad things are happening to its children, it must be asleep. As with many Bambuti rituals, the time it takes to complete a molimo is not rigidly set; instead, it is determined by the mood of the group. Food is collected from each hut to feed the molimo, and in the evening the ritual is accompanied by the men dancing and singing around the fire.” ref

“Women and children must remain in their huts with the doors closed. These practices were studied thoroughly by British anthropologist Colin Turnbull, known primarily for his work with the tribe. “Molimo” is also the name of a trumpet the men play during the ritual. Traditionally, it was made of wood or sometimes bamboo, but Turnbull also reported the use of metal drainpipes. The sound produced by a molimo is considered more important than the material it is made out of. When not in use, the trumpet is stored in the trees of the forest. During a celebration, the trumpet is retrieved by the youth of the village and carried back to the fire.” ref

Genetic history of Africa

“Africans’ genetic ancestry is largely partitioned by geography and language family, with populations belonging to the same ethno-linguistic groupings showing high genetic homogeneity and coherence. Gene flow, consistent with both short- and long-range migration events followed by extensive admixture and bottleneck events, have influenced the regional genetic makeup and demographic structure of Africans. The historical Bantu expansion had lasting impacts on the modern demographic make up of Africa, resulting in a greater genetic and linguistic homogenization. Genetic, archeologic, and linguistic studies added extra insight into this movement: “Our results reveal a genetic continuum of Niger–Congo speaker populations across the continent and extend our current understanding of the routes, timing and extent of the Bantu migration.” ref

“Overall, different African populations display genetic diversity and substructure, but can be clustered in distinct but partially overlapping groupings:

• Khoisan or ‘South African hunter-gatherers’ from Southern Africa represented by the Khoisan peoples; they are associated with the deepest divergence (c. 270,000 years ago) of human genetic diversity, forming a distinct cluster of their own. They subsequently diverged into a Northern and Southern subgroup, c. 30,000 years ago.
• ‘Central African hunter-gatherers’ or ‘Rain forest hunter-gatherers‘ (Pygmies) of Central Africa, represented by the Biaka and Mbuti; associated with another deep divergence (c. 220,000 years ago). They subsequently diverged into an Eastern and Western subgroup, c. 20,000 years ago.
• “Ancestral Eurasians” represent the ancestral population of modern Eurasians shortly before the Out-of-Africa expansion; they are inferred to have diverged from other African populations, most likely somewhere in Northeast Africa, c. 70,000 years ago.
• The various Afroasiatic-speakers from Northern Africa and the Horn of Africa, are suggested to have diverged from other African groups c. 50,000 years ago, but currently insufficient data and geneflow from other groups complicate an accurare estimation of the divergence date. Afroasiatic-speaking populations display variable amounts of West Eurasian (primarily Natufian-like, but also Neolithic Anatolian and Iranian) admixtures, with the remainder being primarily associated with Nilotic-like ancestry. They also display affinity for the Paleolithic North African Taforalt specimens of the Iberomaurusian culture.
• ‘Eastern African hunter-gatherers‘, represented by Hadza, Sandawe, Omotic-speakers, and the ancient Mota specimen; their phylogenetic relationship to other populations is not clear, but they display affinity to modern East and West African populations, and harbor Khoesan-like geneflow along a Northeast to Southwest cline, as well as later (West) Eurasian admixtures, but at lower amounts than among Afroasiatic-speakers.
• “Ancient East Africans” or “Ancestral West/East Africans” associated with the common ancestor of modern Niger-Congo and Nilo-Saharan-speakers originated around 28,000 years ago, likely in the Nile Valley region. They subsequently diverged at c. 18,000 years ago into the ancestors of West and West-Central African Niger-Congo and Bantu-speakers, and into the East African Nilo-Saharan/Nilotic-speakers. They represent the dominant and most widespreaded ancestry component of modern Africa, and are associated with relative recent population expansions linked to agriculture and pastoralist lifestyles. Genetic data indicates affinity for older hunter-gatherer groups in East Africa, but their exact relationship remains unclear. There is evidence for limited geneflow (9-13%) from a human ghost lineage, referred to as ‘West African foragers’ with a deeper or equally deep divergence time than ‘Khoisan hunter-gatherers’, into modern West Africans.
• Austronesian-speaking Malagasy people in Madagascar have received significant East/Southeast Asian admixture associated with the Austronesian expansion, with the remainder ancestry being primarily associated with West-Central and East African components. The estimated date of geneflow between these sources is c. 2,200 years ago.” ref

“The term ‘indigenous Africans‘ refers to the populations with primarily indigenous (non-Eurasian) ancestries, consisting of Niger–Congo speakers, Nilo-Saharan speakers, the divergent and diverse Khoisan grouping, as well as of several unclassified or isolated ethnolinguistic groupings (see unclassified languages of Africa). The origin of the Afroasiatic languages remains disputed, with some proposing a Middle Eastern origin, while others support an African origin with varying degrees of Eurasian and African components. The Austronesian languages originated in southern East Asia, and later expanded outgoing from the Philippines.” ref

“The Niger–Congo languages probably originated in or near the area where these languages were spoken prior to Bantu expansion (i.e. West Africa or Central Africa). Its expansion may have been associated with the expansion of agriculture, in the African Neolithic period, following the desiccation of the Sahara in c. 3500 BCE or around 5,500 years ago. Proto-Niger-Congo may have originated about 10,000 years before present in the “Green Sahara” of Africa (roughly the Sahel and southern Sahara), and that its dispersal can be correlated with the spread of the bow and arrow by migrating hunter-gatherers, which later developed agriculture.” ref

“Although the validity of the Nilo-Saharan family remains controversial, the region between Chad, Sudan, and the Central African Republic is seen as a likely candidate for its homeland prior to its dispersal around 10,000–8,000 BCE or around 12,000 to 10,000 years ago. The Southern African hunter-gatherers (Khoisan) are suggested to represent the autochthonous hunter-gatherer population of southern Africa, prior to the expansion of Bantu-speakers from Western/Central Africa and East African pastoralists. Khoisan show evidence for Bantu-related admixture, ranging from nearly ~0% to up to ~87.1%. Low levels of West Eurasian ancestry (European or Middle Eastern) are found in Khoe–Kwadi Khoesan-speakers. It could have been acquired indirectly by admixture with migrating pastoralists from East Africa. This hypothesis of gene flow from eastern to southern Africa is further supported by other genetic and archaeological data documenting the spread of pastoralism from East to South Africa.” ref

“Significant Eurasian admixture is found in Northern Africa, and among specific ethnic groups of the Horn of Africa, Northern Sudan, the Sahel region, as well as among the Malagasy people of Madagascar. Various genome studies found evidence for multiple prehistoric back-migrations from various Eurasian populations and subsequent admixture with native groups. West Eurasian-associated geneflow arrived to Northern Africa during the Paleolithic (30,000 to 15,000 years ago), followed by other pre-Neolithic and Neolithic migration events. Genetic data on the Taforalt samples “demonstrated that Northern Africa received significant amounts of gene-flow from Eurasia predating the Holocene and development of farming practices”. Medieval geneflow events, such as the Arab expansion also left traces in various African populations. Pickrell et al. (2014) indicated that Western Eurasian ancestry eventually arrived through Northeast Africa (particularly the Horn of Africa) to Southeast Africa and Southern Africa.” ref

“Ramsay et al. (2018) also found evidence for significant Western Eurasian admixture in various parts of Africa, from both ancient and more recent migrations, being highest among populations from Northern Africa, and some groups of the Horn of Africa: In addition to the intrinsic diversity within the continent due to population structure and isolation, migration of Eurasian populations into Africa has emerged as a critical contributor to the genetic diversity. These migrations involved the influx of different Eurasian populations at different times and to different parts of Africa. Comprehensive characterization of the details of these migrations through genetic studies on existing populations could help to explain the strong genetic differences between some geographically neighboring populations.” ref

“This distinctive Eurasian admixture appears to have occurred over at least three time periods with ancient admixture in central west Africa (e.g., Yoruba from Nigeria) occurring between ~7,500 and 10,500 years ago, older admixture in east Africa (e.g., Ethiopia) occurring between ~2,400 and 3,200 years ago and more recent admixture between ~0.15 and 1.5 years ago in some east African (e.g., Kenyan) populations. Subsequent studies based on LD decay and haplotype sharing in an extensive set of African and Eurasian populations confirmed the presence of Eurasian signatures in west, east and southern Africans. In the west, in addition to Niger-Congo speakers from The Gambia and Mali, the Mossi from Burkina Faso showed the oldest Eurasian admixture event ~7,000 years ago. In the east, these analyses inferred Eurasian admixture within the last 4000 years in Kenya.” ref

“There is no definitive agreement on when or where the original homeland of the Afroasiatic language family existed. Some have suggested that they were spread by people with largely West-Eurasian ancestry during the Neolithic Revolution, towards Northern Africa and the Horn of Africa, outgoing from the Middle East, specifically from the Levant. Others argue that the first speakers of Proto-Afroasiatic were based in Northeast Africa because that region includes the majority of the diversity of the Afroasiatic language family and has very diverse groups in close geographic proximity, which is sometimes considered a telltale sign for a linguistic geographic origin.” ref

“A subset of the Proto-Afroasiatic population would have migrated to the Levant during the late Paleolithic, merging with local West-Eurasians and resulting in a population which would later give rise to Natufian culture, associated with the early development of agriculture and early Afroasiatic languages, or specifically pre-proto-Semitic. In addition, Y-haplogroup sub-lineage E-M215 (also known as “E1b1b) and its derivative E-M35 are quite common among Afroasiatic speakers, and southwestern Ethiopia is a plausible source of these haplogroups. Under this African model, the linguistic group and carriers of this lineage would have arisen and dispersed together from Northeast Africa in the Mesolithic, plausibly having already developed subsistence patterns of pastoralism and intensive plant usage and collection.” ref

“The Near-Eastern agriculturalist hypothesis does not account for the domestication of plants endemic to the Horn of Africa such as teff, ensete, and Niger seed, nor does it account for the lack of evidence of intrusive agricultural populations or for the growing of wheat, barley, or sorghum in that region prior to 3000 BCE or around 5,000 years ago. According to historian and linguist Christopher Ehret, the form of intensive plant collection practiced by the Proto-Afroasiatic population in Northeast Africa may have been a precursor to the other agricultural practices that would later independently develop in the Fertile Crescent and the Horn of Africa. While many studies conducted on Horn of Africa populations estimate a West-Eurasian admixture event around 3,000 years ago.” ref

“Hodgson et al. (2014) found a distinct West-Eurasian ancestral component among studied Afroasiatic-speaking groups in the Horn of Africa (and to a lesser extent in North Africa and West Asia), most prevalent among the ethnic Somali. This ancestral component dubbed “Ethio-Somali” is most closely related to the “Maghrebi” (peaking in Tunisians) component and is believed to have diverged from other non-African ancestries around 23,000 years ago, and migrated back to Africa prior to developing agriculture (12,000–23,000 years ago) from the Near East. This population would have crossed via the Sinai Peninsula and then split into two, with one branch continuing west across North Africa and the other heading south into the Horn of Africa. The authors propose that the “Ethio-Somali” component may have been a substantial ancestral component of the Proto-Afroasiatic-speaking population.” ref

“Later migration from Arabia into the HOA beginning around 3,000 years ago would explain the origin of the Ethiosemitic languages at this time. An mtDNA analysis by Gandini et al. (2016) has produced additional evidence in support of a pre-agricultural back-migration from West-Eurasia into the Horn of Africa with an estimated date of arrival into the Horn of Africa in the early Holocene, possibly as a result of obsidian exchange networks across the Red Sea. Hodgson et al. also confirmed the existence of an ancestral component indigenous to the Horn of Africa – “Ethiopic” or “Omotic” (Pagani et al.) – which is most prevalent among speakers of the Omotic branch of Afroasiatic in southwestern Ethiopia. This lineage is associated with that of a 4,500 year-old fossil (Mota) found in a cave in southwestern Ethiopia, which has high genetic affinity to modern Ethiopian groups, especially the endogamous blacksmith caste of the Omotic Aari people.” ref

“Like Mota, Aari blacksmiths do not show evidence for admixture with West-Eurasians, demonstrating a degree of population continuity in this region for at least 4,500 years. In a comparative analysis of Mota’s genome referencing modern populations, Gallego et al. (2016) concluded that the divergence of Omotic from other Afroasiatic languages may have resulted from the relative isolation of its speakers from external groups. In an analysis of 68 Ethiopian ethnic groups, Lopez et al. (2021) revealed that several groups belonging to the three AA classifications of Cushitic, Omotic and Semitic show high genetic similarity to each other on average. Furthermore, the Nilo-Saharan speakers in the southwest shared more recent ancestry with Bantu and Nilotics, in contrast Afro-Asiatic speakers in the northeast shared more recent ancestry with Egyptians and other West Eurasians. The data also supported widespread recent intermixing among various ethnic groups.” ref

“Neolithic agriculturalists, who may have resided in Northeast Africa and the Near East, may have been the source population for lactase persistence variants, including –13910*T, and may have been subsequently supplanted by later migrations of peoples. The Sub-Saharan West African Fulani, the North African Tuareg, and European agriculturalists, who are descendants of these Neolithic agriculturalists, share the lactase persistence variant –13910*T. While shared by Fulani and Tuareg herders, compared to the Tuareg variant, the Fulani variant of –13910*T has undergone a longer period of haplotype differentiation. The Fulani lactase persistence variant –13910*T may have spread, along with cattle pastoralism, between 9686 and 7534 years ago, possibly around 8500 years ago; corroborating this timeframe for the Fulani, by at least 7500 years ago, there is evidence of herders engaging in the act of milking in the Central Sahara.” ref

“Haplogroup R1b-V88 is thought to have originated in Europe and migrated into Africa with farmers or herders in the Neolithic period, c. 5500 BCE or around 7,500 years ago. R1b-V88 is found at a high frequency among Chadic speaking peoples such as the Hausa, as well as in Kanembu, Fulani, and Toubou populations. West African hunter-gatherers, in the region of western Central Africa (e.g., Shum Laka, Cameroon), particularly between 8000 and 3000 years ago, were found to be related to modern Central African hunter-gatherers (e.g., Baka, Bakola, Biaka, Bedzan). Genetically, African pygmies have some key difference between them and Bantu peoples.” ref

“Throughout Sub-Saharan Africa, genetic adaptation (e.g., rs334 mutation, Duffy blood group, increased rates of G6PD deficiency, sickle cell disease) to malaria has been found among Sub-Saharan Africans, which may have initially developed in 7,300 years ago. Sub-Saharan Africans have more than 90% of the Duffy-null genotype. In the rainforests of Central Africa, genetic adaptation for non-height-related factors (e.g., immune traits, reproduction, thyroid function) and short stature (e.g., EHB1 and PRDM5 – bone synthesis; OBSCN and COX10 – muscular development; HESX1 and ASB14 – pituitary gland’s growth hormone production/secretion) has been found among rainforest hunter-gatherers.” ref

“As of 19,000 years ago, Africans, bearing haplogroup E1b1a-V38, likely traversed across the Sahara, from east to west. At Mota, in Ethiopia, an individual, estimated to date to the 5th millennium years ago, carried haplogroups E1b1 and L3x2a. The individual of Mota is genetically related to groups residing near the region of Mota, and in particular, are considerably genetically related to the Aari people, especially the blacksmith caste of that group. At Ol Kalou, in Nyandarua County, Kenya, a pastoralist of the Pastoral Neolithic carried haplogroups E1b1b1b2b2a1/E-M293 and L3d1d. At Kokurmatakore, in Marsabit County, Kenya, a pastoralist of the Pastoral Iron Age carried haplogroups E1b1b1/E-M35 and L3a2a.” ref

“At Nyarindi Rockshelter, in Kenya, there were two individuals, dated to the Later Stone Age (3500 years ago); one carried haplogroup L4b2a and another carried haplogroup E (E-M96, E-P162). At Lukenya Hill, in Kenya, there were two individuals, dated to the Pastoral Neolithic (3500 years ago); one carried haplogroups E1b1b1b2b (E-M293, E-CTS10880) and L4b2a2b, and another carried haplogroup L0f1. At Hyrax Hill, in Kenya, an individual, dated to the Pastoral Neolithic (2300 years ago), carried haplogroups E1b1b1b2b (E-M293, E-M293) and L5a1b. At Molo Cave, in Kenya, there were two individuals, dated to the Pastoral Neolithic (1500 years ago); while one had haplogroups that went undetermined, another carried haplogroups E1b1b1b2b (E-M293, E-M293) and L3h1a2a1.” ref

“Throughout Sub-Saharan Africa, genetic adaptation (e.g., rs334 mutation, Duffy blood group, increased rates of G6PD deficiency, sickle cell disease) to malaria has been found among Sub-Saharan Africans, which may have initially developed in 7,300 years ago. Sub-Saharan Africans have more than 90% of the Duffy-null genotype. In the highlands of Ethiopia, genetic adaptation (e.g., rs10803083, an SNP associated with the rate and function of hemoglobin; BHLHE41, a gene associated with circadian rhythm and hypoxia response; EGNL1, a gene strongly associated with oxygen homeostasis in mammals) to hypoxia and low atmospheric pressure has been found among the Amhara people, which may have developed within the past 5000 years. In Tanzania, genetic adaptation (e.g., greater amount of amylase genes than in African populations that consume low-starch foods) has been found in the Hadza people due to a food diet that especially includes consumption of tubers.” ref

“From the region of Kenya and Tanzania to South Africa, eastern Bantu-speaking Africans constitute a north to south genetic cline; additionally, from eastern Africa to toward southern Africa, evidence of genetic homogeneity is indicative of a serial founder effect and admixture events having occurred between Bantu-speaking Africans and other African populations by the time the Bantu migration had spanned into South Africa. Three Later Stone Age hunter-gatherers carried ancient DNA similar to Khoisan-speaking hunter-gatherers. Prior to the Bantu migration into the region, as evidenced by ancient DNA from Botswana, East African herders migrated into Southern Africa. Out of four Iron Age Bantu agriculturalists of West African origin, two earlier agriculturalists carried ancient DNA similar to Tsonga and Venda peoples and the two later agriculturalists carried ancient DNA similar to Nguni people; this indicates that there were various movements of peoples in the overall Bantu migration, which resulted in increased interaction and admixing between Bantu-speaking peoples and Khoisan-speaking peoples.” ref 

Strong Maternal Khoisan Contribution to the South African Coloured Population: A Case of Gender-Biased Admixture

“Abstract: The study of recently admixed populations provides unique tools for understanding recent population dynamics, socio-cultural factors associated with the founding of emerging populations, and the genetic basis of disease by means of admixture mapping. Historical records and recent autosomal data indicate that the South African Coloured population forms a unique highly admixed population, resulting from the encounter of different peoples from Africa, Europe, and Asia. However, little is known about the mode by which this admixed population was recently founded. Here we show, through detailed phylogeographic analyses of mitochondrial DNA and Y-chromosome variation in a large sample of South African Coloured individuals, that this population derives from at least five different parental populations (Khoisan, Bantus, Europeans, Indians, and Southeast Asians), who have differently contributed to the foundation of the South African Coloured. In addition, our analyses reveal extraordinarily unbalanced gender-specific contributions of the various population genetic components, the most striking being the massive maternal contribution of Khoisan peoples (more than 60%) and the almost negligible maternal contribution of Europeans with respect to their paternal counterparts. The overall picture of gender-biased admixture depicted in this study indicates that the modern South African Coloured population results mainly from the early encounter of European and African males with autochthonous Khoisan females of the Cape of Good Hope around 350 years ago.” ref 

R1b1b (R-V88)

“R-V88 (R1b1b): the most common forms of R1b found among males native to Sub-Saharan Africa, also found rarely elsewhere. R1b1b (PF6279/V88; previously R1b1a2) is defined by the presence of SNP marker V88, the discovery of which was announced in 2010 by Cruciani et al.  Apart from individuals in southern Europe and Western Asia, the majority of R-V88 was found in the Sahel, especially among populations speaking Afroasiatic languages of the Chadic branch. Based on a detailed phylogenic analysis, D’Atanasio et al. (2018) proposed that R1b-V88 originated in Europe about 12,000 years ago and crossed to North Africa between 8,000 and 7,000 years ago, during the ‘Green Sahara‘ period. R1b-V1589, the main subclade within R1b-V88, underwent a further expansion around 5,500 years ago, likely in the Lake Chad Basin region, from which some lines recrossed the Sahara to North Africa.” ref

“Marcus et al. (2020) provide strong evidence for this proposed model of North to South trans-Saharan movement: The earliest basal R1b-V88 haplogroups are found in several Eastern European Hunter Gatherers close to 11,000 years ago. The haplogroup then seemingly spread with the expansion of Neolithic farmers, who established agriculture in the Western Mediterranean by around 7,500 years ago. R1b-V88 haplogroups were identified in ancient Neolithic individuals in Germany, central Italy, Iberia, and, at a particularly high frequency, in Sardinia. A part of the branch leading to present-day African haplogroups (V2197) was already derived in Neolithic European individuals from Spain and Sardinia, providing further support for a North to South trans-Saharan movement. European autosomal ancestry, mtDNA haplogroups, and lactase persistence alleles have also been identified in African populations that carry R1b-V88 at a high frequency, such as the Fulani and Toubou. The presence of European Neolithic farmers in Africa is further attested by samples from Morocco dating from c. 5400 BCE or around 7,400 years ago onwards.” ref

“Studies in 2005–08 reported “R1b*” at high levels in Jordan, Egypt and Sudan. Subsequent research by Myres et al. (2011) indicates that the samples concerned most likely belong to the subclade R-V88. According to Myres et al. (2011), this may be explained by a back-migration from Asia into Africa by R1b-carrying people. As can be seen in the above data table, R-V88 is found in northern Cameroon in west central Africa at a very high frequency, where it is considered to be caused by a pre-Islamic movement of people from Eurasia.” ref 

Hausa people

“The Hausa (autonyms for singular: Bahaushe, Bahaushiya; plural: Hausawa and general: Hausa; exonyms: Ausa; Ajami: مُتَنٜىٰنْ هَوْسَا / هَوْسَاوَا) are a native ethnic group in West Africa. They speak the Hausa language, which is the second most spoken language after Arabic in the Afro-Asiatic language family. The Hausa are a culturally homogeneous people based primarily in the Sahelian and the sparse savanna areas of southern Niger and northern Nigeria respectively, numbering around 86 million people, with significant populations in Benin, Cameroon, Ivory Coast, Chad, Central African Republic, Togo, Ghana, as well as smaller populations in Sudan, Eritrea, Equatorial Guinea, Gabon, Senegal, Gambia.” ref

“Predominantly Hausa-speaking communities are scattered throughout West Africa and on the traditional Hajj route north and east traversing the Sahara, with an especially large population in and around the town of Agadez. Other Hausa have also moved to large coastal cities in the region such as Lagos, Port Harcourt, Accra, Abidjan, Banjul and Cotonou as well as to parts of North Africa such as Libya over the course of the last 500 years. The Hausa traditionally live in small villages as well as in precolonial towns and cities where they grow crops, raise livestock including cattle as well as engage in trade, both local and long distance across Africa. They speak the Hausa language, an Afro-Asiatic language of the Chadic group. The Hausa aristocracy had historically developed an equestrian based culture. Still a status symbol of the traditional nobility in Hausa society, the horse still features in the Eid day celebrations, known as Ranar Sallah (in English: the Day of the Prayer).” ref

“Daura is the cultural center of the Hausa people. The town predates all the other major Hausa towns in tradition and culture. The Hausa have, in the last 500 years, criss-crossed the vast landscape of Africa in all its four corners for varieties of reasons ranging from military service, long-distance trade, hunting, performance of hajj, fleeing from oppressive Hausa feudal kings as well as spreading Islam. The homeland of Hausa people is Hausaland (“Kasar Hausa”), situated in Northern Nigeria and Southern Niger. However, Hausa people are found throughout Africa and Western Asia. Daura, in northern Nigeria, is the oldest city of Hausaland. The Hausa of Gobir, also in northern Nigeria, speak the oldest surviving classical vernacular of the language.” ref

“The Hausa Kingdoms were independent political entities in what is now Northern Nigeria. The Hausa city states emerged as southern terminals of the Trans-Saharan caravan trade. Like other cities such as Gao and Timbuktu in the Mali Empire, these city states became center of long-distance trade. By the 12th century CE, the Hausa were becoming one of Africa’s major trading powers, competing with Kanem-Bornu and the Mali Empire. According to a Y-DNA study by Hassan et al. (2008), about 47% of Hausa in Sudan carry the West Eurasian haplogroup R1b. The remainder belong to various African paternal lineages: 15.6% B, 12.5% A and 12.5% E1b1a. A small minority of around 4% are E1b1b clade bearers, a haplogroup which is most common in North Africa and the Horn of Africa.” ref

“A more recent study on Hausa of Arewa (Northern Nigeria) revealed similar results: 47% E1b1a, 5% E1b1b, 21% other Haplogroup E (E-M33, E-M75…), 18% R1b and 9% B. In terms of overall ancestry, an autosomal DNA study by Tishkoff et al. (2009) found the Hausa to be most closely related to Nilo-Saharan populations from Chad and South Sudan. This suggests that the Hausa and other modern Chadic-speaking populations originally spoke Nilo-Saharan languages, before adopting languages from the Afroasiatic family after migration into that area thousands of years ago:

From K = 5-13, all Nilo-Saharan speaking populations from southern Sudan, and Chad cluster with west-central Afroasiatic Chadic-speaking populations. These results are consistent with linguistic and archeological data, suggesting a possible common ancestry of Nilo-Saharan speaking populations from an eastern Sudanese homeland within the past ≈10,500 years, with subsequent bi-directional migration westward to Lake Chad and southward into modern day southern Sudan, and more recent migration eastward into Kenya and Tanzania ≈3,000 ya (giving rise to Southern Nilotic speakers) and westward into Chad ≈2,500 years ago (giving rise to Central Sudanic speakers) (S62, S65, S67, S74). A proposed migration of proto-Chadic Afroasiatic speakers ≈7,000 years ago from the central Sahara into the Lake Chad Basin may have caused many western Nilo-Saharans to shift to Chadic languages (S99). Our data suggest that this shift was not accompanied by large amounts of Afroasiatic16 gene flow. Analyses of mtDNA provide evidence for divergence ≈8,000 years ago of a distinct mtDNA lineage present at high frequency in the Chadic populations and suggest an East African origin for most mtDNA lineages in these populations (S100).” ref

“A study from 2019 that genotyped 218 unrelated males from the Igbo, Hausa and Yoruba tribes using X-STR analysis, found that when studying the genetic affinity, no significant differences were detected. It supported a homogeneity of Nigerian ethnic groups for X-chromosome markers. In 2024, a paper similarly found homogeneity in the Yoruba, Igbo and Hausa in Nigeria for X-Chromosomes (mtDNA). However, differences in the Hausa were found for the Y-Chromosome, where they had more paternal lineages associated with Afro-Asiatic speakers, while the Yoruba and Igbo were paternally related to other Niger-Congo speaking groups. Specifically, in the 135 Yoruba and 134 Igbo males, E-M2 was seen at high rates of 90%. In contrast, the 89 Hausa males had E-M2 at 43%, and frequencies for R1b-V88 at 32%, A 9%, E1a 6%, B 5%, and another 5% being made of other lineages.” ref

“The Hausa cultural practices stand unique in Nigeria and have withstood the test of time due to strong traditions, cultural pride as well as an efficient precolonial native system of government. Consequently, and in spite of strong competition from western European culture as adopted by their southern Nigerian counterparts, have maintained a rich and particular mode of dressing, food, language, marriage system, education system, traditional architecture, sports, music and other forms of traditional entertainment. The Hausa language, a member of Afroasiatic family of languages, has more first-language speakers than any other African language. It has around 50 million first-language speakers, and close to 30 million second-language speakers.” ref

“The main Hausa-speaking area is northern Nigeria and southern Niger. Hausa is also widely spoken in northern Ghana, Cameroon, Chad, and Ivory Coast as well as among Fulani, Tuareg, Kanuri, Gur, Shuwa Arab, and other Afro-Asiatic, Niger-Congo, and Nilo-Saharan speaking groups. Maguzanci, an African Traditional Religion, was practised extensively before Islam. In the more remote areas of Hausaland, the people continue to practise Maguzanci. Closer to urban areas, it is not as common, but with elements still held among the beliefs of urban dwellers. Practices include the sacrifice of animals for personal ends, but it is not legitimate to practise Maguzanci magic for harm. People of urbanized areas tend to retain a “cult of spirit possession,” known as Bori. It incorporates the old religion’s elements of African Traditional Religion and magic.” ref

“Hausa animism, Maguzanci or Bori is a pre-Islamic traditional religion of the Hausa people of West Africa that involves magic and spirit possession. While only a part of the Hausa people (mostly within urban elites) converted to Islam before the end of the 18th century, most of the adherents of the religion did the same between the jihad started by the Islamic reformer Usman dan Fodio around 1800 and the middle of the 20th century, while a small minority converted to Christianity. Religious affiliation to this traditional religion is virtually nonexistent at the beginning of the 21st century; however, Hausa animism and Islam among Hausa people have coexisted for centuries, and some practices related to animism carry on locally.” ref

“When discussing “bori”, there is a distinction to be made between the beliefs of the Manguzawa (which consist of pagan or heavily pagan influenced Hausa religion), the general belief in spirits and animism that persists even among Muslim Hausa, and the possession-trance group that puts on dance performances and ceremonies. Bòòríí is a Hausa noun, meaning the spiritual force that resides in physical things, and is related to the word for local distilled alcohol (borassa) as well the practice of medicine (boka).^[2] The Bori religion is both an institution to control these forces, and the performance of an “adorcism” (as opposed to exorcism) ritual, dance and music by which these spirits are controlled and by which illness is healed.” ref

“Spirits are called bori, iska (plu. iskoki), or aljan. Iska has a non-Muslim connotation, so many Muslim Hausa prefer to use the term aljan, which comes from the Arabic word jinn. Possessing spirits in the possession-trance group are called the spirit(s) “on (your) head”. Possession-trance group members are called yan bori (children of the bori), dam bori (son of the bori), yar bori (daughter of the bori), doki (horse) for male devotees, and godiya (mare) for female devotees. Attendants who help the yam bori but don’t trance are called masu kiwo (herders or grooms), and help them get dressed and keep the spectators safe. Musicians may generally be called maroka, though more specific terms exist. Female musicians are called zabiyoyi. A mawaki is a vocalist. A mabushi is a wind instrument player. A makadi is a drummer. The female chorus at bori events is called the Yan Kwarya whether they are professional musicians or not.” ref

“An aspect of the traditional Maguzawa Hausa people‘s religious traditions, Bori became a state religion led by ruling-class priestesses among some of the late precolonial Hausa Kingdoms. When Islam started making inroads into Hausaland in the 11th century, certain aspects of the religion such as idol worship were driven underground. The cult of Tsumbubura in the then-Sultanate of Kano and many other similar Bori cults were suppressed, but Bori survived in “spirit-possession” cults by integrating some aspects of Islam. The Bori spirit possession priestesses maintained nominal influence over the Sultanates that replaced the earlier Animist kingdoms. Priestesses communed with spirits through ecstatic dance ritual, hoping to guide and maintain the state’s ruling houses. A corps of Bori priestesses and their helpers was led by royal priestess, titled the Inna, or “Mother of us all”. The Inna oversaw this network, which was not only responsible for protecting society from malevolent forces through possession dances, but which provided healing and divination throughout the kingdom.” ref

There are many spirits connected to people, animals, plants, and big rocks. The two personal (“friendly”^[16]) bori are like the qarin, which does not come into being until after the person it is attached to is born, as that is when a person’s sex is known (one of these qarin-like spirit is of the opposite sex). All these- people, animals, plants, and big rocks- have a permanent soul (quruwa), two attendant angels, and a bori of the same sex. There are other bori not directly connected to living people, such as those which are or are inspired by Muslim saints, well known jinn, embodiments of other tribes, ancestors, the spirits of infants, totems (such as animals), and gods.” ref

“The bori are like humans, but they are not human, and they are not visible in human cities. They are considered both above humans, in heaven (because they are sometimes conflated with angels), and below humans in the earth. The bori, like people, keep cattle, though this does not prevent them from bothering human herds. Many of the bori belong to 12 families or “houses”. Bori are considered to be like the people they live among. The names, characteristics, favorite resting places, appropriate sacrifices, and more of the spirits form the basis of ritual practice. The rosters of known Bori spirits and their behaviors tend to remain the same, and the spirits stay relevant, though their genealogies may change. Spirits are often divided into “white” or “black”, which mostly overlaps with the categories of “Muslim” and “pagan”, and the categories of “town” and “bush”. However, a Muslim spirit may be called black if it causes certain afflictions, such as paralysis. The black/white and Muslim/pagan distinction seems to be Muslim in origin, while the town/bush distinction seems to have been native, and location based identification is more prominent.” ref

“In Kano, Jangare, the city in which the Bori live, is ruled by a “Court of the East”, with Sarkin Aljan as the head, and is a centralized authority compared to others’ accounts of a more mild feudal organization. In Ningi, he is the ultimate head and divides authority with the heads of the other houses. Everyone agrees that there are 12 houses, but they do not agree on who rules or is part of them. Houses are generally said to be ruled by a first born, and spirits are grouped by ethnicity, occupation, and descent. This becomes complicated as relational terms used for spirits are vague, and spirits practice a complex system of child avoidance and adoption. The basic information is contained in praise-epithets, songs, and spirit behavior at ceremonies, but is interpreted differently. Though these interpretations vary between different people, each person is consistent about their own recounting. The houses are linked by adoption, clientship, office, kinship, and affinity. For example, the Fulani house is linked to all other houses via marriage, and occasionally through the marriages of the offspring of brothers and half brothers. The other spirits tend to endogamy.” ref

“These spirits can cause illness and are placated with offerings, sacrifices, dances, and possession rites where dancers specially prepare to ensure being “ridden” has no ill effects. Their permission must be asked before constructing buildings, and neglect and unintentional slights may anger them. They can be entreated to help in tasks, such as finding treasure, and with solving fertility issues. In the latter case, the bori ask God’s permission to intervene. The bori are everywhere, but are more concentrated near temples, within which they can be imprisoned. Certain bori may prefer to stay in specific areas, such as drains. Specific bori are associated with causing specific ailments. They are also associated with specific plants and specific types of soil, which are used medically to cure spirit ailments. They are associated with specific songs, and typically have more than one. The spirits will not ride their mounts without music, and this music may be as simple as hand clapping.” ref

Malady and manifestation are the two main methods of communication the bori have. The former allows them to communicate their anger at being ignored or offended. The latter allows them self-expression, and in the case of one origin story of the bori, functions as a family reunion. Incense attracts the bori, and they do not like iron. Fire is not a bori, and bori do not like fires or live in them, as it would burn them. However, the bori can simply go over fires, so fire is not a ward against them. It is considered good to give as much of an offering as one can afford, because the bori love the generous and take care of them. The bori get sustenance from blood, and sacrifices are quick ways to get their attention. The blood must be collected in one spot on the ground, preferably near a tree a particular bori is known to rest in.” ref

“Precautions are taken so that unfriendly bori do not possess fetuses. One method to protect newborns is to buy a black hen at around 7 months of pregnancy, and to keep it in the house until the baby is born. It is thought that any bori lingering will possess it and lie in wait for the birth. It is then set free in the Jewish quarter to get rid of the unfriendly bori. This method is borrowed from Arabs. A young child may be protected by their mother calling them Angulu (vulture, which bori are thought to find disgusting, though this is also the name of a bori), and acting as though she’d be glad of her child was gone, as bori take children to punish their mothers.” ref

“If a person yawns without covering their mouth, they must spit afterwards, as doing so may accidentally cause one to let in a bori. Sneezing is thought to expel a bori that has entered someone without their knowledge, and this is part of why a person gives thanks to God after they sneeze. For this reason, the bori do not like pepper. The sound of laughter attracts the bori, the merriment of laughing excites them, and the open mouth, just as with yawning, allows them entry.” ref

“One story of the creation of the bori spirits says that God created everything, and at first the bori did not exist. However, some people did wicked things, and God turned some of these people into half men-half fish, and the rest were turned into bori. They were further cursed to stay in the same state; old bori never die, and young bori never age to become old. Another story is similar to the Ethiopian and Omani story of the origin of zār spirits; a Hausa family with many children tried to hide half of them from God. This angered God, who turned them into hungry spirits that can only be appeased with blood sacrifices. Possession in this context is a family reunion that restores health and balance (lafiya), and spirit and human are complementary opposites.” ref

“For the bori possession-trance group, affliction by spirits, even if it was caused by one committing a transgression, is a mark that someone was chosen by the spirits to become a horse for the spirits or gods. The illnesses they treat include clumsiness, impotence, infertility, rashes, boils, gastrointestinal trouble, headaches, insanity, leprosy, and paralysis. As of the 1910s, totemism had limited importance and recognition. When it was recognized, each clan had a totem, regarded as sacred, which was connected to a patron bori. Both would be referred to as “kan gida” (head of the house). Children inherited the bori of their fathers, though they may also honor their mother’s. Women kept their totems even after marriage, and husbands had the choice of if they would allow her to sacrifice near their home (which was more common) or if she needed to return to her father’s home. One was free to marry someone with the same or different totem.” ref

“The totem tree (connected to either the bori or the totem animal) was never cut, and the totem animal was never eaten. The totem animal was only allowed to killed around harvest time by the chief men of the clan. They would smear the blood on their faces, particularly the forehead (associated with the bori). The head of the animal was sundried and put in the chief’s home until it was replaced next year. The rest was buried. Everyone would bathe at least three days before, and was absintent until a day or two after the ritual. Accidentally killing the totem at other times was not punished. Intentionally killing the totem would result in death, potentially caused by the totem’s bori. Eating it, even accidentally, would cause illness. A bori ceremony may be held a few days after the ritual totem killing.” ref

“Incense may be used to summon totems, and different incenses are used for different animals. Most totem animals appear in bori dances. As of the 1910s in Tunis and Tripoli, there were bori houses (temples) with appointed priestesses, and a chief priest and priestess of West African origin. The priestess must be able to speak Hausa so she can direct performances, and she must be abstinent. She was usually a widow or divorced. The chief priest does not need to speak Hausa, and must be honest and of good judgement. Neither position is hereditary.” ref

“One may form a contract with a bori by sacrificing an animal associated with it. This is commonly done to remedy a misfortune or illness caused by the spirit, or to gain its favor and permission before undertaking certain actions. The spirit drinks the sacrificial blood and the meat is given by alms (sadaka) to Quran scholars and students, the homeless and destitute, or devotees and musicians. The recipient, upon being told what they’re being given is alms, customarily says “May Allah grant your wish.” The witness is a crucial part of the sacrifice, and has been emphasized so much that instead of being given raw meat as alms, one may also be given cooked meat or meatless food. In essence, alms for the bori now include grains and vegetables alongside meat.” ref

“Once one has human permission to build a house, they go to the building site and offer a sacrifice. This will always involve a white hen and a red rooster (only the bori Kuri and Mai-Inna accept these). If one can afford it, they also sacrifice a male goat, and if one is wealthy, they sacrifice a bull (all bori accept either). The blood is spilled on the ground for the bori. The future homeowner and friends eat the flesh. Another hen and rooster sacrifice is done when one moves in. The same is done when building a farm, though the goat is more optional. When moving into an already built house, one sacrifices a hen on the threshold. Similarly, when digging a well, a person would have a diviner go to the desired area, and they would use charms to point out a good dig site. The digger would sacrifice two foul and start digging.” ref

“In Nigeria, as part of the home building, one may set apart a building where incense offerings were done each Thursday, which summons the bori from anywhere in the world. Two fowl were sacrificed on anniversaries of the home building. By the 1910s, this practice had ceased among Nigerian Hausa Muslims, and was not relevant to Hausa Muslims in Tunis and Tripoli, as they were not allowed to build their own homes. Grace is said before and after meals, but thanks is not given to the bori during this. Different issues regarding bori may be resolved in different ways. For example, a bori may cause a false pregnancy where a woman gains weight for 9 or more months.” ref

“This usually happens because a jealous woman or disappointed lover entreated a bori to do so, and can be hard to solve as bori lie about their identity to diviners, making it hard to know which is responsible. Another instance is if a woman struggles to conceive, she serks help from a boka or mallam. She burns incense for three days in a row, and breathes it in as she prays to God, Mohammed, Kuri, and other bori. This process may be intended to clear her of evil influence. An unwed girl’s male bori may cause her period to stop suddenly to keep her from marrying and leaving him. When this happens, attempts are made to placate the bori in other ways.” ref

“A child who cries all the time is afflicted by a bori (usually Sa’idi) and the curative method is to hold the child over incense until it quiets. To protect a child from the Yayan Jiddari, ground nuts and sweets are placed by their head for three nights. After this, the treats are taken to a Mai-Bori, who places them in a pot for a few days. They will be eaten by the bori and vanish. If a childdoesnt develop properly, a Mai-Bori or Boka is consulted to find out the bori responsible, and the bori is sacrificed to. If a saint (marabout) is involved, the mother and other womem of the house may take the child to their tomb. There, they light a candle, burn incense, and rub the child with either the blood of a sacrificed white cock or with dirt from near the tomb. After this, another candle is lit and more incense is burnt. A gift is given to the tomb’s caretaker as well.” ref

“The two types of rituals done by the Yan Bori are girka (the initiation ritual), and the periodic ceremonies. The latter is necessary after going through the former, but the Yan Bori do not penalize wayward members; scorning the bori, including by not going to periodic ceremonies, causes a resurgence in symptoms. The exception is wasam bori (play of the bori) a type of periodic ceremony that usually does not involve possession-trance. Girka may be done in addition to herbal cures, functioning as a cure and as part of initiation into the bori possession-trance tradition. The word “girka” is related to the word for “boiling”, and as such evokes both the idea of preparing traditional medicine and the heat felt in the bodies of mounts experiencing genuine trance. The girka ceremony involves the patient leaving their former residence, especially if it is with their relatives, for in-patient treatment.” ref

“Generally, they do not return to live with their relatives after the ceremony is done. A “father” or “mother of girka” and their female assistant oversees all arrangements prior and during the girka, and the assistant acts as a nurse and cooks for the patient. An uwal saye may also be involved. Patients do not bring many personal items during treatment and only wear a body wrapper. Medicines start more broadly, in association with families of spirits known to cause certain problems, and become specific as the specific spirit(s) become known. Some medicines also protect the mounts from any strain they may experience while being ridden, as some spirits can be physically demanding. When musicians come during the girka process, the assistant prepares the patient by laying out a grass mat facing to Jangare and leads them around it.” ref

“She then seats them, facing east, and places a special amulet in their hands and around their neck. She then ties their big toes and thumbs together and covers them in ehite cloth. She takes a seat to their right, facing north. This process signifies the submission of the mount to the spirits and sets them at ease. For 6–13 days, musicians will play 3 times a day for the patient, starting with Sarkin Makada and ending with Nakada. The first three days are dedicated to the songs for the pagan “black” spirits. During this, a wand made from Calotropis procera may be used to coax the spirits to dismount by pointing them away from the patient’s head. Through this process the spirits on their head are made peace with, and the patient gains skill in behaving as a mount.” ref

“An uwal saye (trainer) will know herbal spirit medicine, and cure the patient who has been diagnosed with a spirit causing their affliction accordingly. The uwal saye trains the new people in the dress, behavior, and personality of the spirits; knowledge of the spirit world (Jangare, the city of spirits^[13]); and what illness they cause. This training is mostly kept secret. Acting as a mount is considered a learned skill. The final stage of the cure is acting in a dance as mount during the kwanan zauen (the night spent sitting up), also called wasa na hira (entertainment for chatting). This is a ceremony where the patient is presented as a “horse of the gods”, and they are publicly ridden for the first time and recognized as a mount. This takes place on the 7th day, if the former stage only took the usual 6 days.” ref

“Experienced mounts who share the spirit dance alongside the new one, showing them the finer points of the performance. After this, the new mount bathes, gives gifts to the assistant that helped, and sponsors a sacrifice to their primary spirit. The items used during initiation are gifted by the new mount to the assistant as well, as they are polluted and unfit for use by the mount. This establishes relations with the spirit, and is considered self administered.” ref

“This creates a lifelong relationship of mutual benefit; the patient is healed and can help others as a medium, and the spirit can express itself in performances. Possession-trance is important as a cure, since it is thought that if the bori are the origin of a malady, they must also be the origin of its cure. Part of this type of cure is acceptance by the patient to be involved in the possession-trance group, and belief in its practices. This is the primary form of initiation, and after this, the new devotee can treat others with similar spirit afflictions. Possession is viewed negatively, as the possessed no longer behaves normally. However, initiation into the Yan Bori turns possession into a more positive state. In instances where it is deemed impossible to cure someone, the individual is still encouraged to participate in Yan Bori events, and will be supported by the group.” ref

“It is generally inappropriate for a musician to be an active Yan Bori member. As such a musician who has taken up the nanyle of boka cannot instruct his patients in trance as he has likely not done it, and it would be improper for him to do so. Curing rituals organized by musicians as such mostly focus on curing the patient, and it is thought that musicians will be able to keep the spirit dormant via recognition of their power and regular sacrifices. As such, the patient will not need to trance. The Yan Bori regard these cures as incomplete or worse, but outsiders view them as an effective alternative to full initiation. The patient is still considered a “horse of the gods”, but is not expected to trance. Instead they give gifts to other mounts when they trance.” ref

“When organizing an event, the host gives a gift to a Yan Bori member or musician they have an affinity with, who shares this gift with musicians and trancers. By taking a share in this gift, one is obligated to participate in the ceremony. This gift may consist of kola nuts, candy, or money. Occasionally a participant will accept the invitation without gift if the intermediary or the host has a relationship with them, and may use this as a future bit of leverage or to satisfy a standing social debt. Another gift is usually given to the musicians by the host upon arrival to further coax them to set up to perform. This may also be replaced by an obligation, as with the invitation gift.” ref

“Attendants to bori dances dress in their best things. During these ceremonies, songs and praise-epithets are sung to and about the spirits by maroka (musicians). These are not always the same songs and praise-epithets, and may vary in the same performance. This variation helps create a full picture of the spirit. The vocal music honors and calls the spirits, while the instrumental music induces trance and controls the flow of time at the ceremony. The first song played at bori ceremonies, both in urban and in many rural areas, is one praising Allah.” ref

“The ideal maroka speaks clearly, enunciates well, and knows many stories and praise-epithets about spirits (sometimes knowing these about almost 300 different spirits). The spirits can’t be called without their music. The maroka are professional musicians, and typically are not themselves possessed. Their music groups usually consist of a leader, chorus, praise-shouter, and an optional vocalist. All the maroka sit, except for the praise-shouter, who walks in front of them, praising the spirits and announcing gifts. When an audience member or participant wishes to address the gathering, they take the praise-shouter to the side. The praise-shouter shouts the proceedings to a halt and says the praises and genealogy of the speaker, who he calls mai magana (owner of speech). They tell the praise-shouter their message quietly and he shouts it out. He is paid 1/10 of a naira for this as of the 1970s.” ref

“The maroka‘s music induces trance in the “mounts” (performing possessed) with a gradually increasing tempo, and then the spirits are called down. The mounts sit before the musicians, sometimes covered with a cloth, and move in intensity with the music. They have entered trance when they collapse or begin moving as the spirit. The mounts consciously try to disconnect and provide themselves as a vessel for the spirits. Auditory and visual intensity are used to induce trance, and this technique is also found in Hamadsha rituals. At events where different pieces of music are played for different spirits, mounts often prepare to be mounted by multiple spirits; at events a spirit only takes one mount at a time, and it is possible for a mount to be mounted, take a break, and return to be mounted again. If a spirit wishes to leave a performance, it will thank the musicians and have its mount sit down. The musicians play a short piece for it, and the spirit dismounts (often signaled by the mount sneezing).” ref

“The dancer who is possessed (“ridden”) is called a horse. During possession, only the spirit speaks, and the human is not held responsible for what occurs during possession-trance. Possession-trance is characterized by the mounts acting out the speech and behavior of specific spirits. The Yan Bori recognize that possession-trances may be authentic or fraudulent, with the latter being a result of an inept mount. Relatedly, because proper possession-trance is a learned skill, it is expected that new devotees will be less proficient and spectacular. True trance is marked by intense sweating and high body temperature. Amnesia regarding what occurred during trance is common. When one wants a bori to enter, such as at this time, one does not say “thanks be to God”. Two mounts of the same spirit may have it manifest differently.” ref

“Events without trance and with simultaneously induced trance from one song are structurally similar. The main difference is in the latter case the host must present a gift of kola nuts, candy, and cash and ask the musicians to begin. The mounts come over, and if they accept a portion of the gift, they will trance. Most women in the possession-trance group do not attend the public ceremonies, instead going to secluded, private ceremonies overseen by other women. The music there is performed by women on a kwarya.” ref

“Bori cures provide safety and support for marginalized Hausa, as well as entertainment in the form of dances. These ceremonies may be considered theater, sacred, or both by outsiders. Many first become involved with bori ceremonies and groups because of illnesses that could not be cured. Bori dance ceremonies may be done as often as once a week to maintain health and the relationship between human and spirit. These periodic performances promote solidarity among the possession-trance group, reaffirm their obligations, and separate and bond together different groups affiliated with the possession-trance rites. They are frequent during the first half of the dry season and gradually drop in frequency in anticipation of the rainy season, where few ceremonies and initiations are done. Performances pick up again after the harvest. Performances are not done during Ramadan.” ref

“The Yan Dandu often hang around the brothels associated with the possession-trance group for social and economic reasons, and attend the public dance ceremonies. They may dance at these themselves, though they typically do not trance. They give small gifts of money to trancing mounts, especially if Dan Galadima is present. One of the ceremonies a Magajiyar Bori may preside over is one where she invites others to her compound to witness her be ridden by as many of her spirits as possible, ideally all of them. She is the only active medium here (though accidental trances may occur from others). Each spirit is greeted on arrival, and wished well on its journey out. The spirits may give gifts of kola nuts, sweets, vegetables, perfume, and money. The audience may also give money by pressing it to the forehead of the Magajiyar Bori and ketting it fall to the ground. Guests also give the musicians in attendance money.” ref

“Different rituals associated with the Yan Bori may be classified by the instruments used, wfich devotees themselves do to some extent. These distinct ways of calling a spirit would be kidan kwarya (drumming on calabashes), kidan garaya (strumming a two stringed lute), kidan goge (bowing a one stringed lute), and busan sarewa (blowing a whistle). They may also be classified by venue and participants. The kidan amada has songs played on kwarya by non-professional female musicians. They are usually secluded, held inside a compound, and largely attended by women. Kidan bori can refer to any ritual or ceremony where bori are called, but usually refers to public performances with mixed gender audiences where the garaya, goge, and sarewa are played by professional male musicians. These events have different protocol for gift giving. Small amounts of money given to mounts are done at both without formality.” ref

“The act of gift giving, which is one of the more obvious displays of generosity in Hausa culture, has added significance in accordance with the giver and receiver’s respective positions in the social hierarchy when done publicly. In the context of the Yan Bori, the gift implies separateness between the giver and receiver, its content is determined by the givers position in relation to the possession-trance group and broader Hausa society, and underlines the ties between separate and unequal social levels as the gift cannot be directly repaid in kind. This is because exchanges between equals do not often occur at ceremonies (the exception being spirits giving each other ritual objects their attendants have to retrieve at the end). The repayment for gifts of money is via services, such as participation in the possession-trance group and witnessing a ceremony. Repayment, especially between those who are not equals, comes in a form different from the original gift. Money begets services, and sacrifices beget divine favor.” ref

“While the mount embodies a spirit, those not experiencing trance have the opportunity to give the spirit gifts. For some, such as other mounts of the same spirit, this may be more of an obligation. Not doing so may anger the spirit and cause their ailment to reoccur. For others, they may do this to get answers for questions. Most gifts are money, and most gifts end up as property of the mount. Large gifts given at public rituals must be announced by a praise shouter. A stop is called for, where the mounts in trance stand “at ease” and the musicians stop playing. The name of the gift giver, their genealogy, and praise-epithets are said. Them the gifts are described in a way that’s accurate, but that inflates their important and quantity. For this service the praise-shouter is given a smaller additional gift of money.” ref

“Gifts to those not directly involved in a performance can done discreetly, but those for spirits, mounts, or performing musicians should be brought forward and displayed at the feet of the seated musicians. Different shoes and hats are done for each recipient to make distribution easier. All gifts given to dancers that don’t trance are given by them to the musicians. Usually all gifts to the maroka are pooled and divided up according to rank, but if one favors a particular musician, they can catch the their eye and say something indicating they should step to the side, where the secret gift is given. If spotted by a praise shouter, the gift will be made public knowledge. When the event ends, participants gather around the musicians for the first distribution. The musicians share is not counted yet, but each praise shouter takes a small amount from it.” ref

“The leader of the musicians then counts the money and itemizes the other gifts given to the mounts, announcing the amount each spirit was given. The mount comes forward and collects it, giving one third of it to the musicians as a thank you, and another third to their attendant or other assistant. Black male goats were at one point preferred sacrifices. However, chickens, guinea fowl, and pigeons were also used, though turkey and ducks were not. Sacrificial animals are selected based on breed, color, sex, and so on, in order to correspond to the appropriate spirit.” ref

“The principal leader of Bori possession-trance groups in a region is the Sarkin Bori (chief of the Bori). This role is usually filled by a man. He is nominally the authority over all other Bori mediums in a region, and hosts any mediums traveling through an area. He organizes performances, sending gifts and invites to the performers. He is given a portion of the performance proceeds, and may be responsible for distributing them. This role is chosen by the possession-trance group based on an individual’s experience as a medium and their popularity, and confirmed via the turbaning ceremony used for political offices in general Hausa society. The actual power and role of the Sarkin Bori varies regionally. The Sarkin Bori and other male leaders are in charge of public possession-trance ceremonies.” ref

“The Magajiyar Bori (heiress of the Bori) is always a woman. Technically the position is subordinate to the Sarkin, but individual Magajiyar Bori may have equal or greater renown. Some may oversee districts as large as the district the Sarkim Bori, but most oversee smaller districts. She may also be the head of a brothel, and her district may be the area surrounding it. The Magajiyar Bori and female leaders in charge of secluded ceremonies. Because of the public/private distinction, while many think of possession-trance groups as having an single vertical hierarchy, it is more accurate to describe them as parallel. The possession-trance group may borrow titles associated with the emirate structure to describe itself. These titles are used because they are a familiar system for organizing.” ref

“The musicians (maroka) preserve the group’s oral tradition, and are thought of as mediators between the mounts and their spirits. Some musicians also are boka, or are astrologers who do horoscopes. Semi urban musicians all tend to divide their time between music and another occupation, such as farming, though they will primarily be seen as maroka. Yan Bori musicians may double as boka, diviners (which in practice may amount to being a counselor), and sellers of Bori paraphernalia. In order to be a boka, one must also be a Yan Bori initiate. Normally, the Sarkin Bori is consulted as a boka, so musicians only fill this role when local leadership is weak or nonexistent. Being a diviner does not conflict with Bori beliefs, which focus on the past and present.” ref

“Divining is not particularly connected to Bori beliefs, and as such it can be easier for Muslim Hausa to admit to believing in astrology than the Bori. Both Maguzawa and Hausa Muslims primarily employ musicians for spiritual purposes at possession-trance ceremonies. In the Hausa social structure, which is primarily based on occupation for men, musicians, praise singers, and praise shouters are that bottom, and at the bottom of them are the possession-trance musicians. Others affiliated with the group are not ranked by their group affiliation.” ref

“Musician groups internally rank themselves, either with titles or more casually. The first is less common. One such example of titles being used had the leader of a female musician group be turbaned with the title Sarkin kidan Kwarye (Chief of beating the kwarya). Her assistants were Majidadin Kwarya (Majidadi of Kwarya) and Wazirin Kwarya (Waziri of Kwarya). As with non-musical Yan Bori positions, music groups elect leaders for the title. It could be that formal titles are mostly used by musicians associated with royal courts, which is not the case for most musicians associated with the Yan Bori.” ref

“It is more common for the leader of a musician group to be called by his personal name or nickname, and the instrument he plays- such as “(Name) Mai Garaya”, meaning “(Name) who plays the garaya”. The leader manages the musicians, and the event host may only deal with the leader. If the Yan Bori leaders in a region are weak or absent, it may the responsibility of the leading musician that does Yan Bori performances to invite the mounts to perform at events. This especially the case for performances at weddings and namings.” ref

“The music leader is expected to be generous with dividing the money earned at performances. They must choose carefully not to performances with small or stingy crowds, and ideally should take pay cuts to prevent friction in the group if the earrings were small. A normal way of dividing performance money would be 50% for the leader, and the remaining 50% divided evenly among the other musicians. Apprentices occasionally appear in music groups, and receive smaller shares of money.” ref

“Prospective musicians are mainly the sons or nephews of musicians (through maternal uncles), with the former being more common. In the latter case, having maternal relatives that are musicians is not the primary reason one becomes a musician, and more so is a justification as most Hausa don’t think anyone would choose to be a musician. Further, musicians affiliated with the Yan Bori usually have family members who were, or are themselves initiates. The latter is less common, as it is not seen as socially acceptable for a musician to trance or do other things typical of a Yan Bori. Being a musician is a potential alternative to being a trancer, though it is not commonly taken.” ref

“Being a Yan Bori musician is about more than being able to play certain instrument or tune. They believe in the power of their music and the spirits. The latter puts them in conflict with mainstream Muslim beliefs (though many Hausa privately believe in the Bori). This conflict and stereotypes about them make the maroka sensitive about their public perception. Learning about the Bori starts young. Urban Yan Bori musicians often live in compounds with the Yan Bori and their sympathizers or nearby. Both groups are usually poor and ostracized for their beliefs, and are as such drawn together. Collectively they teach their children about the Bori and to respect them, often at evening gossip sessions in or near compounds.” ref

“In addition to music and information about the Bori, young musicians are taught acceptable behavior in broader society, being more deferential and using more flattery due to their status. Other Hausa see this as insincere, but musicians that don’t behave accordingly are criticized. Yan Bori musicians have more deviants than musicians in general, and behave accordingly. For example, it is proper to visit Bori patrons, but if one’s finances aren’t in chaos, a Yan Bori musician will stay home instead of making an expected visit to emphasize their independence. The Yan Bori musicians are confident that the patron will call upon them and the Yan Bori regardless once the spirit bothers them again.” ref

“Women and men in the Yan Bori keep somewhat separate. Many are also unmarried, and therefore don’t live with spouses. Instead they live with each other in compounds under the leadership of a senior, who may also be a devotee. These compounds may also function as brothels. The higher status a leader has, the more people usually live with them in a compound. Superiors in Hausa society give money and material gifts to inferiors who repays with services, with exception of a compound leader collecting rent from inferior devotees in their compound. These compounds are centers for Yan Bori activity and places for mediums to socialize. Even devotees who live alone or with family visit the compound after the evening meal.” ref

“Those with spirits of the same generation and house are considered equals. Gifts between equals among the Yan Bori are repaid in kind; money for money, food for food, cloth for cloth. Gifts between devotees of the same sex are more common than cross sex gifts. New devotees are typically the mounts of lower ranked, junior spirits. Senior devotees are typically ridden by spirits of the senior generation, house leaders, or Sarkin Aljan. The Yan Bori are expected to own objects to demonstrate possession-trance for each spirit on their heads, but only have complete costumes for a few. The materials are expensive, and most spirits only get a few damaru (medicine belts), a zane (body cloth), and a few other items. Items can be borrowed from seniors, but it is considered better to own them. Usually the makers and more common sellers of such items are initiates, and this is true of the musicians that sell them.” ref

“Achievements within the Yan Bori do not increase one’s general social status, but do increase their ritual status. While broader Hausa society may often primarily select leaders based on status, genealogy, and the personal reasons of an emir, all Yan Bori leaders and the leaders of the closely associated music groups are selected based on their achievements and skill. High cult status is marked by the ability to give, and doing so validates one’s status in the eyes of others. The seniority of a Yan Bori member is based on their ability to act convincingly and skillfully as a mount. Sex, age, time since initiation, and wealth do not matter in this respect. As one gains seniority one is called to publicly act as mount for the more important, senior spirits on their head. It is unusual for any new initiate to start with the more important spirits. One must prove and hone their skill first. Devotees whose spirits “travel together” often become friends.” ref

Human Y chromosome haplogroup R-V88: a paternal genetic record of early mid Holocene trans-Saharan connections and the spread of Chadic languages

“Abstract: Although human Y chromosomes belonging to haplogroup R1b are quite rare in Africa, being found mainly in Asia and Europe, a group of chromosomes within the paragroup R-P25^* are found concentrated in the central-western part of the African continent, where they can be detected at frequencies as high as 95%. Phylogenetic evidence and coalescence time estimates suggest that R-P25^* chromosomes (or their phylogenetic ancestor) may have been carried to Africa by an Asia-to-Africa back migration in prehistoric times. Here, we describe six new mutations that define the relationships among the African R-P25^* Y chromosomes and between these African chromosomes and earlier reported R-P25 Eurasian sub-lineages.” ref

“The incorporation of these new mutations into a phylogeny of the R1b haplogroup led to the identification of a new clade (R1b1a or R-V88) encompassing all the African R-P25^* and about half of the few European/west Asian R-P25^* chromosomes. A worldwide phylogeographic analysis of the R1b haplogroup provided strong support to the Asia-to-Africa back-migration hypothesis. The analysis of the distribution of the R-V88 haplogroup in >1800 males from 69 African populations revealed a striking genetic contiguity between the Chadic-speaking peoples from the central Sahel and several other Afroasiatic-speaking groups from North Africa. The R-V88 coalescence time was estimated at 9,200–5,600 years ago, in the early mid Holocene. We suggest that R-V88 is a paternal genetic record of the proposed mid-Holocene migration of proto-Chadic Afroasiatic speakers through the Central Sahara into the Lake Chad Basin, and geomorphological evidence is consistent with this view.” ref

“The Sahara, covering about one third of the African continent from the Atlantic Ocean to the Red Sea, is the earth’s most extensive desert. Over the past thousands of years, the Sahara has undergone dramatic climatic oscillations including arid phases, during which it was largely uninhabitable, and humid episodes, which transformed the desert into a fertile landscape. After a hyper-arid period about 23,000–14,500 years ago, the Saharan region experienced a monsoonal moist climate, characterized by increased rainfall.” ref

“During the Holocene Climatic Optimum (about 10,000–5,000 years ago), a few thousand years after the beginning of the humid period, flora and fauna repopulated the desert, and a mosaic of savannah and woodland became well established throughout much of the Sahara. At the same time, the Sahara was home to giant lakes, the largest of which, the paleolake Megachad, may have possibly covered an area of at least 400 000 km², more than the Caspian Sea, the biggest lake on earth today. This greening scenario was interrupted by a number of arid episodes, and at about 5,000–6,000 years ago, the region experienced a rapid onset of dryer conditions. These marked the beginning of a shift towards permanent aridity, with variations in the distribution and timing of these changes between the eastern and central/western Sahara. Human–environment interactions in the Sahara have been greatly influenced by these climate fluctuations.” ref

“Close links between climatic variations and prehistoric human occupation of the Sahara during the early mid Holocene (10,000–5,000 years ago) are documented by archeological and paleoanthropological evidence. However, genetic studies have been limited and mainly focused on uniparental markers and the role of the Nile basin as a corridor for human movements between northeastern and eastern Africa. There have only been a few high-resolution analyses to date regarding the distribution of Y-specific haplogroups in the African continent. The emerging picture indicates a clear differentiation between central/western sub-Saharan and northern African populations. Haplogroup E-DYS271, which accounts for >70% of the Y chromosomes in most of the populations south of the Sahara, is found on an average at a frequency of 2–3% in Northern Africa, whereas haplogroups J-M304, E-M81, and E-M78, which on the whole account for 50–90% of the northern African male-specific region of the Y chromosome (MSY) gene pool, have been only rarely observed in west/central sub-Saharan Africa.” ref

“A group of chromosomes of potential interest to past trans-Saharan connections is the paragroup R1b1^* (R-P25^*). Cruciani et al found this paragroup (at that time defined as haplogroup 117, or R-M173^*(xSRY₁₀₈₃₁, M18, M73, M269)) to be present at high frequencies (up to 95%) in populations from northern Cameroon. The same paragroup was only rarely observed in other sub-Saharan African regions, and not observed at all in western Eurasia. Subsequent studies dealing with the MSY diversity in Africa have confirmed the presence of R-P25^*(xM269) in northern Cameroon at high frequencies and, at lower frequencies (mean 5%, range 0–20%), of R-P25^* immediately south of Cameroon, in several populations from Gabon. Interestingly, chromosomes of haplogroup R-P25/R-M173, ancestral for M269 as well as for other ‘Eurasian’ downstream markers, have been found to be present in northern Africa (1% in Algeria, 4% in Tunisia, and 2–4% in Egypt). The presence of R-P25 Y chromosomes has also been reported in population groups from the Sudan; however, as no internal markers were typed, the sub-haplogroup affiliation of these chromosomes remains undefined.” ref

“To shed some light on the past demographic processes that determined the present distribution of R-P25^* in Africa, we searched for new MSY mutations refining the phylogeny of haplogroup R1b, and surveyed a wide range of African populations (>1800 males from 69 populations) for the presence of the R1b haplogroup. More than 3500 subjects from Europe and Asia were also analyzed for the same haplogroup to obtain a better insight into the Asia-to-Africa back migration associated with this haplogroup. All the African R1b chromosomes, with the exception of one eastern- and a few northern-African R-M269 chromosomes, turned out to belong to the haplogroup R-V88. About one third of the African R-V88 chromosomes carried mutation V69, which was not observed outside Africa. The large majority of R1b chromosomes from western Eurasia carried, as expected, the M269 mutation; only five R-V88 chromosomes were observed, three of which carried distinctive mutations (M18, V35, and V7). The rare R1b chromosomes observed in Asia were either R-M73 or R-M269. The R-P25^* paragroup was only found in five subjects from Europe (3), western Asia (1), and eastern Asia (1).” ref

“According to the phylogeography of macro-haplogroup K-M9 (which contains haplogroup R1b), an ancient Asia-to-Africa back migration has been hypothesized to explain the puzzling presence of R-P25^* in sub-Saharan Africa. This hypothesis is strongly supported by the present data. In the revised Y chromosome phylogeny, there are 119 lineages in the macro-haplogroup K-M9 (which includes haplogroups K1-K4 and L to T). Of these lineages, only two have been observed in sub-Saharan Africa at appreciable frequencies: T-M70 and R-V88. Both haplogroups have also been observed in Europe and western Asia. If the presence of R1b chromosomes in Africa was not because of a back migration, we would have to assume that all the mutations that connect M9 with V88 in the MSY phylogeny (>50 mutations) originated in Africa.” ref

“Under this scenario, we should assume that all the K-M9 lineages that are now found outside sub-Saharan Africa have survived extinction, whereas those which should have accumulated in Africa are now extinct (with the exception of T-M70 and R-V88) and this is an unlikely scenario. We obtained the time estimate of the Asia-to-Africa back migration by using the variation associated with seven tetranucleotide microsatellites and the ASD method. As an upper limit, we used the coalescence time of the R-M343/P25 haplogroup (12,900 years ago, 95% CI=11,600–14,300 years ago, under a conservative scenario of constant population size), which, on the basis of the accumulated nucleotide and microsatellite diversity, most likely originated outside Africa. The coalescence time of the seemingly African-specific haplogroup R-V69 (6,000 years ago, 95% CI=4,200–8,200 years ago, under the hypothesis of an expanding population) was used as a lower limit.” ref

“Within Africa, the highest frequencies of the R-V88 haplogroup (and its commonest sub-clade, R-V69) were observed in the central Sahel (northern Cameroon, northern Nigeria, Chad, and Niger). Immediately south of this region (southern Cameroon and southern Nigeria), frequencies drastically dropped to 0.0–4.8%. The central Sahel is characterized by a strong linguistic fragmentation with populations speaking languages belonging to three of the four linguistic families of Africa (Afroasiatic, Niger-Congo, and Nilo-Saharan). When the linguistic affiliation of the populations from the central Sahel was also taken into account, a clear-cut divide was observed between those speaking Afroasiatic languages (including the Berber-speaking Tuareg, the Semitic Arab Shuwa, and Chadic-speaking populations from northern Cameroon) and the other populations (Mann–Whitney test P=1.4 × 10⁻³), with Chadic-speaking populations mostly contributing to this difference. It is worth noting that, if the finding of 20% R-V88 chromosomes among the Hausa is representative, this population, encompassing by far more people than all other Chadic speakers, also encompasses the highest absolute number of V88 carriers.” ref

“In contrast to prior studies on nuclear (mostly autosomal) ins/del and microsatellite markers, the Chadic are distinguished from the Nilo-Saharan-speaking populations at the Y chromosome variation level. Repeated assimilations of Nilo-Saharan females over generations may account for these conflicting signals. Among the Niger-Congo-speaking populations, the frequency of the haplogroup R-V88 ranged between 0.0 and 66.7%. Outside central Africa, haplogroup R-V88 was only observed in Afroasiatic-speaking populations from northern Africa, with frequencies ranging from 0.3% in Morocco, to 3.0% in Algeria, and to 11.5% in Egypt, where a particularly high frequency (26.9%) was observed among the Berbers from the Siwa Oasis. Although the presence of the haplogroup R-V88 at non-negligible frequencies in some Niger-Congo-speaking populations from the central Sahel can be accounted for by Chadic admixture favored by geographic contiguity, the presence of this haplogroup both in northern Africa and the central Sahel is especially intriguing given that >1500 km across the Sahara separate the two regions. The expansion time for the haplogroup R-V88 in Africa, under two different population models (see Materials and methods), was found to be 9,200–5,600 years ago (95% CI=7.6–10,800 and 4,700–6,600 years ago, respectively).” ref

“Diverse hypotheses have been proposed to explain the process by which proto-Chadic speakers arrived to the Lake Chad region. Ehret has put forward a model for Afroasiatic languages with a primary division between the Omotic languages of Ethiopia and an Erythraean subgroup. This, in turn, has been subdivided into Cushitic and North Erythraean, the latter including Berber, Semitic, Ancient Egyptian, and Chadic. In his opinion, around 7,000 years ago proto-Chadic Afroasiatic speakers may have moved west through the Central Sahara and then farther south into the Lake Chad Basin. Blench, in turn, suggested that speakers of proto-Cushitic–Chadic language migrated east-to-west from the Middle Nile to the Lake Chad, and recent mtDNA data support this view.” ref 

“However, in contrast to the mtDNA, a strong connection between Chadic and other Afroasiatic populations from Northern Africa is revealed by the Y chromosome data. This finding would indicate the trans-Saharan a more likely scenario than the inter-Saharan hypothesis, at least as far as the male component of gene pool is concerned. In this view, it is tempting to speculate that the Y chromosome haplogroup R-V88 represents a preserved genetic record of gene flow along the same axis as the proposed spread of proto-Chadic languages. Indeed, geomorphological evidence4 from the paleolakes that existed in the Sahara during the mid-Holocene indicates that these lakes could have covered an area as large as about 10% of the Sahara, providing an important corridor for human migrations across the region.” ref

“In summary, our data indicate a significant male contribution from northern Africa (and ultimately Asia) to the gene pool of the central Sahel. The trans-Saharan population movements resulting in this genetic pattern would seem to mirror the spread of the proto-Chadic languages, and most likely took place during the early mid Holocene, a period when giant paleolakes may have provided a corridor for human migrations across what is now the Sahara desert.” ref 

Chad Genetic Diversity Reveals an African History Marked by Multiple Holocene Eurasian Migrations 

“Understanding human genetic diversity in Africa is important for interpreting the evolution of all humans, yet vast regions in Africa, such as Chad, remain genetically poorly investigated. Here, we use genotype data from 480 samples from Chad, the Near East, and southern Europe, as well as whole-genome sequencing from 19 of them, to show that many populations today derive their genomes from ancient African-Eurasian admixtures. We found evidence of early Eurasian backflow to Africa in people speaking the unclassified isolate Laal language in southern Chad and estimate from linkage-disequilibrium decay that this occurred 4,750–7,200 years ago.” ref

“It brought to Africa a Y chromosome lineage (R1b-V88) whose closest relatives are widespread in present-day Eurasia; we estimate from sequence data that the Chad R1b-V88 Y chromosomes coalesced 5,700–7,300 years ago. This migration could thus have originated among Near Eastern farmers during the African Humid Period. We also found that the previously documented Eurasian backflow into Africa, which occurred ∼3,000 years ago and was thought to be mostly limited to East Africa, had a more westward impact affecting populations in northern Chad, such as the Toubou, who have 20%–30% Eurasian ancestry today. We observed a decline in heterozygosity in admixed Africans and found that the Eurasian admixture can bias inferences on their coalescent history and confound genetic signals from adaptation and archaic introgression.” ref

“African genetic diversity is still incompletely understood, and vast regions in Africa remain genetically undocumented. Chad, for example, makes up ∼5% of Africa’s surface area, and its central location, connecting sub-Saharan Africa with North and East Africa, positions it to play an important role as a crossroad or barrier to human migrations. However, Chad has been little studied at a whole-genome level, and its position within African genetic diversity is not well known. With 200 ethnic groups and more than 120 indigenous languages and dialects, Chad has extensive ethnolinguistic diversity.¹ It has been suggested that this diversity can be attributed to Lake Chad, which has attracted human populations to its fertile surroundings since prehistoric times, especially after the progressive desiccation of the Sahara starting ∼7,000 years ago.” ref

“Important questions about Africa’s ethnic diversity are the relationships among the different groups and the relationships between cultural groups and existing genetic structures. In the present study, we analyzed four Chadian populations with different ethnicities, languages, and modes of subsistence. Our samples are likely to capture recent genetic signals of migration and mixing and also have the potential to show ancestral genomic relationships that are shared among Chadians and other populations. An additional major question relates to the prehistoric Eurasian migrations to Africa: what was the extent of these migrations, how have they affected African genetic diversity, and what present-day populations harbor genetic signals from the ancient migrating Eurasians?” ref

“Researchers have previously reported evidence of gene flow from the Near East to East Africa ∼3,000 years ago, as well as subsequent selection in Ethiopians on non-African-derived alleles related to light skin pigmentation. A recent attempt to quantify the extent of such backflow into Africa more generally, by using ancient DNA (aDNA), suggested that the impact of the Eurasian migration was mostly limited to East Africa. However, previous studies using mitochondrial DNA and the Y chromosome in populations from the Chad Basin found some with an East African or Mediterranean and Eurasian influence, and analysis based on genome-wide data found a non-African component (suggested to be from East Africa) in central Sahelian populations. Thus, studying diverse Chadian populations on a whole-genome level presents an opportunity to shed more light on the history of African-Eurasian mixtures, including whether or not selection after admixture is a widespread phenomenon in Africa and how the historical events in Chad are related to events that have occurred elsewhere in Africa and the Near East.” ref

“In this work, we present a genetic dataset of 480 Chadian, Near Eastern, and European individuals genotyped at 2.5 million SNPs, in addition to high-coverage whole-genome sequences from 19 of these individuals. From Chad, we studied (1) the Toubou, who are nomads from northern Chad and speak a Nilo-Saharan language; (2) the Sara, who are a sedentary population from southern Chad and also speak a Nilo-Saharan language; (3) the Laal speakers, a population of just ∼750 individuals who speak an unclassified language isolate and live in southern Chad; and (4) an urban population from the capital city of N’Djamena. In addition to the Chadians, we included Greek, Lebanese, and Yemen samples whose location and history suggest that they might be informative about early African-Eurasian migrations. We used this dataset to advance our understanding of human genetic diversity in Africa and neighboring regions by focusing on population migration and mixing and how the admixture process has shaped present-day genetic variation.” ref

African Pygmies

“The African Pygmies (or Congo Pygmies, variously also Central African foragers, “African rainforest hunter-gatherers” (RHG) or “Forest People of Central Africa”) are a group of ethnicities native to Central Africa, mostly the Congo Basin, traditionally subsisting on a forager and hunter-gatherer lifestyle. They are divided into three roughly geographic groups:

• The western Bambenga, or Mbenga (Cameroon, Gabon, Republic of the Congo, Central African Republic),
• the eastern Bambuti, or Mbuti, of the Congo basin (DRC)
• the central and southern Batwa, or Twa (Rwanda, Burundi, DRC, Tanzania, Uganda, Zambia, Angola and Namibia). The more widely scattered (and more variable in physiology and lifestyle) Southern Twa are also grouped under the term Pygmoid.” ref

“They are notable for, and named for, their short stature (described as “pygmyism” in anthropological literature). They are assumed to be descended from the original Middle Stone Age expansion of anatomically modern humans to Central Africa, albeit substantially affected by later migrations from West Africa, from their first appearance in the historical record in the 19th century limited to a comparatively small area within Central Africa, greatly decimated by the prehistoric Bantu expansion, and to the present time widely affected by enslavement at the hands of neighboring Bantu, Ubangian and Central Sudanic groups.” ref

“Most contemporary Pygmy groups are only partially foragers and partially trade with neighboring farmers to acquire cultivated foods and other material items; no group lives deep in the forest without access to agricultural products. A total number of about 900,000 Pygmies were estimated to be living in the central African forests in 2016, about 60% of this number in the Democratic Republic of Congo. The number does not include Southern Twa populations, who live outside of the Central Africa forest environment, partly in open swamp or desert environments.” ref

“Additionally, West African hunter-gatherers may have dwelled in western Central Africa earlier than 32,000 years ago and dwelled in West Africa between 16,000 and 12,000 years ago until as late as 1000 years ago or some period of time after 1500 CE. West African hunter-gatherers, many of whom dwelt in the forest–savanna region, were ultimately acculturated and admixed into larger groups of West African agriculturalists, akin to the migratory Bantu-speaking agriculturalists and their encounters with Central African hunter-gatherers.” ref

“African Pygmies are often assumed to be the direct descendants of the Middle Stone Age hunter-gatherer peoples of the central African rainforest. Genetic evidence for the deep separation of Congo Pygmies from the lineage of West Africans and East Africans, as well as admixture from archaic humans, was found in the 2010s. The lineage of African Pygmies is strongly associated with mitochondrial (maternal line) haplogroup L1, with a divergence time between 170,000 and 100,000 years ago. They were partially absorbed or displaced by later immigration of agricultural peoples of the Central Sudanic and Ubangian phyla beginning after about 5,500 years ago, and, beginning about 3,500 years ago, by the Bantu, adopting their languages.” ref

“Dembner (1996) reported a universal “disdain for the term ‘pygmy'” among the Pygmy peoples of Central Africa: the term is considered a pejorative, and people prefer to be referred to by the name of their respective ethnic or tribal groups, such as Bayaka, Mbuti and Twa. There is no clear replacement for the term “Pygmy” in reference to the umbrella group. A descriptive term that has seen some use since the 2000s is “Central African foragers”. Regional names used collectively of the western group of Pygmies are Bambenga (the plural form of Mbenga), used in the Kongo language, and Bayaka (the plural form of Aka/Yaka), used in the Central African Republic. The Congo Pygmy speak languages of the Niger–Congo and Central Sudanic language families. There has been significant intermixing between the Bantu and Pygmies.” ref

“There are at least a dozen Pygmy groups, sometimes unrelated to each other. They are grouped in three geographical categories:

• the western Bambenga (Mbenga) of Cameroon and Gabon, the Bayaka (Aka and Baka), the Bakola or Bakoya (Gyele and Kola), and the Bongo. These groups are speakers of Bantu and Ubangian languages
• The Bambuti (Mbuti) of the Ituri Rainforest, speakers of Bantu and Central Sudanic languages
• the widely scattered Batwa:
• the Great Lakes Twa of the Great Lakes, speakers of the Bantu Rundi and Kiga languages.
• the “Pygmoid” Southern Twa, not always included in the term “Pygmy”, as they tend to be somewhat taller (male average above 155 cm). Subgroups include the Echuya Twa, Mongo Twa, Lukanga Twa and Kafwe Twa.” ref

 “Substantial non-Bantu and non-Ubangian substrates have been identified in Aka and in Baka, respectively, on the order of 30% of the lexicon. Much of this vocabulary is botanical, deals with honey harvesting, or is otherwise specialized for the forest and is shared between the two western Pygmy groups. This substrate has been suggested as representing a remnant of an ancient “western Pygmy” linguistic phylum, dubbed “Mbenga” or “Baaka”. However, as substrate vocabulary has been widely borrowed between Pygmies and neighboring peoples, no reconstruction of such a “Baaka” language is possible for times more remote than a few centuries ago. An ancestral Pygmy language has been postulated for at least some Pygmy groups, based on the observation of linguistic substrates. According to Merritt Ruhlen (1994), “African Pygmies speak languages belonging to either the Nilo-Saharan or the Niger–Kordofanian family. It is assumed that Pygmies once spoke their own language(s), but that, through living in symbiosis with other Africans, in prehistorical times, they adopted languages belonging to these two families.” ref

“Roger Blench (1997, 1999) criticized the hypothesis of an ancestral “Pygmy language”, arguing that even if there is evidence for a common ancestral language rather than just borrowing, it will not be sufficient to establish a specifically “Pygmy” origin rather than any of the several potential language isolates of (former) hunter-gatherer populations that ring the rainforest. He argued that the Pygmies do not form the residue of a single ancient stock of Central African hunter-gatherers, but that they are rather descended from several neighboring ethno-linguistic groups, independently adapting to forest subsistence strategies. Blench adduced the lack of clear linguistic and archaeological evidence for the antiquity of the African Pygmies, that the genetic evidence, at the time of his writing, was inconclusive, and that there is no evidence of the Pygmies having a hunting technology distinctive from that of their neighbors. He argued that the short stature of Pygmy populations can arise relatively quickly (in less than a few millennia) under strong selection pressures.” ref

“West African hunter-gatherers may have spoken a set of presently extinct Sub-Saharan West African languages. In the northeastern region of Nigeria, Jalaa, a language isolate, may have been a descending language from the original set(s) of languages spoken by West African hunter-gatherers. Genetic studies have found evidence that African Pygmies are descended from the Middle Stone Age peopling of Central Africa, with a separation time from West and East Africans of the order 130,000 years. African Pygmies in the historical period have been significantly displaced by, and assimilated to, several waves of Niger–Congo and Nilo-Saharan speakers, of the Central Sudanic, Ubangian, and Bantu phyla.” ref

“Genetically, African pygmies have some key differences between them and Bantu peoples. African pygmies’ uniparental markers display the most ancient divergence from other human groups among anatomically modern humans, second only to those displayed among some Khoisan populations. Researchers identified an ancestral and autochthonous lineage of mtDNA shared by Pygmies and Bantus, suggesting that both populations were originally one, and that they started to diverge from common ancestors around 70,000 years ago. After a period of isolation, during which current phenotype differences between Pygmies and Bantu farmers accumulated, Pygmy women started marrying male Bantu farmers (but not the opposite). This trend started around 40,000 years ago, and continued until several thousand years ago. Subsequently, the Pygmy gene pool was not enriched by external gene influxes.” ref

“Mitochondrial haplogroup L1c is strongly associated with pygmies, especially with Bambenga groups. L1c prevalence was variously reported as: 100% in Ba-Kola, 97% in Aka (Ba-Benzélé), and 77% in Biaka, 100% of the Bedzan (Tikar), 97% and 100% in the Baka people of Gabon and Cameroon, respectively, 97% in Bakoya (97%), and 82% in Ba-Bongo. Mitochondrial haplogroups L2a and L0a are prevalent among the Bambuti. Patin, et al. (2009) suggest two unique, late Pleistocene (before 60,000 years ago) divergences from other human populations, and a split between eastern and western pygmy groups about 20,000 years ago.” ref

Ancient DNA

“Ancient DNA was able to be obtained from two Shum Laka foragers from the early period of the Stone to Metal Age, in 8000 years ago, and two Shum Laka foragers from the late period of the Stone to Metal Age, in 3000 years ago.” ref

“The mitochondrial DNA and Y-Chromosome haplogroups found in the ancient Shum Laka foragers were Sub-Saharan African haplogroups. Two earlier Shum Laka foragers were of haplogroup L0a2a1 – broadly distributed throughout modern African populations – and two later Shum Laka foragers were of haplogroup L1c2a1b – distributed among both modern West and Central African agriculturalists and hunter-gatherers. One earlier Shum Laka forager was of haplogroup B and one later Shum Laka forager haplogroup B2b, which, together, as macrohaplogroup B, is distributed among modern Central African hunter-gatherers (e.g., Baka, Bakola, Biaka, Bedzan).” ref

“The autosomal admixture of the four ancient Shum Laka forager children was ~35% Western Central African hunter-gatherer and ~65% Basal West African – or, an admixture composed of a modern western Central African hunter-gatherer unit, a modern West African unit, existing locally before 8000 years ago, and a modern East African/West African unit likely from further north in the regions of the Sahel and Sahara.” ref

“The two earlier Shum Laka foragers from 8000 years ago and two later Shum Laka foragers from 3000 years ago show 5000 years of population continuity in region.  Yet, modern peoples of Cameroon are more closely related to modern West Africans than to the ancient Shum Laka foragers. Modern Cameroonian hunter-gatherers, while partly descended, are not largely descended from the Shum Laka foragers, due to the apparent absence of descent from Basal West Africans.” ref

“The Bantu expansion is hypothesized to have originated in a homeland of Bantu-speaking peoples located around western Cameroon, a part of which Shum Laka is viewed as being of importance in the early period of this expansion. By 3000 years ago, the Bantu expansion is hypothesized to have already begun. Yet, the sampled ancient Shum Laka foragers – two from 8000 years ago and two from 3000 years ago – show that most modern Niger–Congo speakers are greatly distinct from the ancient Shum Laka foragers, thus, showing that the ancient Shum Laka people were not the ancestral source population of the modern Bantu-speaking peoples.” ref

“While Southern African hunter-gatherers are generally recognized as being the earliest divergent modern human group, having diverged from other groups around 250,000– 200,000 years ago, as a result of the sampling of the ancient Shum Laka foragers, Central African hunter-gatherers are shown to have likely diverged at a similar time, if not even earlier.” ref

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