Early human migrations

Source: Wikipedia, the free encyclopedia.
Further information: History of human migration and Pre-modern human migration

Putative migration waves out of Africa and back migrations into the continent, as well as the locations of major ancient human remains and archeological sites (López et al., 2015).

Early human migrations are the earliest migrations and expansions of archaic and modern humans across continents. They are believed to have begun approximately 2 million years ago with the early expansions out of Africa by Homo erectus. This initial migration was followed by other archaic humans including H. heidelbergensis, which lived around 500,000 years ago and was the likely ancestor of Denisovans and Neanderthals as well as modern humans. Early hominids had likely crossed land bridges that have now sunk.

Within Africa,

across Europe
about 40,000 years ago.

Early Eurasian Homo sapiens fossils have been found in Israel and Greece, dated to 194,000–177,000 and 210,000 years old respectively. These fossils seem to represent failed dispersal attempts by early Homo sapiens, who were likely replaced by local Neanderthal populations.

The migrating modern human populations are known to have

interbred with earlier local populations, so that contemporary human populations are descended in small part (below 10% contribution) from regional varieties of archaic humans.[note 2]

After the

Austronesian expansion
.

Early humans (before Homo sapiens)

See also: Early expansions of hominins out of Africa

The

Kenyan Rift Valley, where the oldest known stone tools have been found. Stone tools recently discovered at the Shangchen site in China and dated to 2.12 million years ago are claimed to be the earliest known evidence of hominins outside Africa, surpassing Dmanisi in Georgia by 300,000 years.[6]

Homo erectus

Main articles: Homo erectus and Oldowan

Between 2 and less than a million years ago,

Saharan pump, around 1.9 million years ago.[citation needed] Homo erectus dispersed throughout most of the Old World, reaching as far as Southeast Asia. Its distribution is traced by the Oldowan lithic industry, by 1.3 million years ago extending as far north as the 40th parallel (Xiaochangliang
).

Key sites for this early migration out of Africa are Riwat in Pakistan (~2 Ma?[7]), Ubeidiya in the Levant (1.5 Ma) and Dmanisi in the Caucasus (1.81 ± 0.03 Ma, p=0.05[8]).

China shows evidence of Homo erectus from 2.12 mya in Gongwangling, in Lantian county.[9] Two Homo erectus incisors have been found near Yuanmou, southern China, and are dated to 1.7 mya, and a cranium from Lantian has been dated to 1.63 mya. Artefacts from Majuangou III and Shangshazui in the Nihewan basin, northern China, have been dated to 1.6–1.7 mya.[9][10] The archaeological site of Xihoudu (西侯渡) in Shanxi province is the earliest recorded use of fire by Homo erectus, which is dated 1.27 million years ago.[11]

Atapuerca).[12]

Robert G. Bednarik has suggested that Homo erectus may have built rafts and sailed oceans, a theory that has raised some controversy.[13]

After H. erectus

Further information: Neanderthal extinction
Further information: Clactonian, Micoquien, and Mousterian
Spread of Denisovans and Neanderthals after 500,000 years ago.
Known Neanderthal range with separate populations within Europe and the Caucasus (blue), the Near East (orange), Uzbekistan (green), and the Altai region (purple).

One million years after its dispersal, H. erectus was diverging into new species. H. erectus is a

Denisovans
.

H. heidelbergensis, Neanderthals and Denisovans expanded north beyond the

Urals (Komi Republic, 65°01′N 57°25′E / 65.02°N 57.42°E / 65.02; 57.42).[15]

Other archaic human species are assumed to have spread throughout Africa by this time, although the fossil record is sparse. Their presence is assumed based on traces of

admixture with modern humans found in the genome of African populations.[5][16][17][18] Homo naledi, discovered in South Africa in 2013 and tentatively dated to about 300,000 years ago, may represent fossil evidence of such an archaic human species.[19]

Neanderthals spread across the Near East and Europe, while Denisovans appear to have spread across Central and East Asia and to Southeast Asia and Oceania. There is evidence that Denisovans interbred with Neanderthals in Central Asia where their habitats overlapped.[20] Neanderthal evidence has also been found quite late at 33,000 years ago at the 65th latitude of the Byzovaya site in the Ural Mountains. This is far outside of any otherwise known habitat, during a high ice cover period, and perhaps reflects a refugia of near extinction.

Homo sapiens

Dispersal throughout Africa

Further information:
Archaeogenetics of sub-Saharan Africa

Homo sapiens are believed to have emerged in Africa about 300,000 years ago, based in part on thermoluminescence dating of artifacts and remains from Jebel Irhoud, Morocco, published in 2017.[note 4][22] The Florisbad Skull from Florisbad, South Africa, dated to about 259,000 years ago, has also been classified as early Homo sapiens.[23][24][25][26] Previously, the Omo remains, excavated between 1967 and 1974 in Omo National Park, Ethiopia, and dated to 200,000 years ago, were long held to be the oldest known fossils of Homo sapiens.[27]

In September 2019, scientists reported the computerized determination, based on 260

skull shape of the last common human ancestor to anatomically modern humans, representative of the earliest modern humans, and suggested that modern humans arose between 260,000 and 350,000 years ago through a merging of populations in East and South Africa.[28][29]

In July 2019, anthropologists reported the discovery of 210,000 year old remains of a H. sapiens and 170,000 year old remains of a H. neanderthalensis in Apidima Cave in southern Greece, more than 150,000 years older than previous H. sapiens finds in Europe.[30][31][32][33]

Early modern humans expanded to Western Eurasia and Central, Western and Southern Africa from the time of their emergence. While

early expansions to Eurasia appear not to have persisted,[34][20] expansions to Southern and Central Africa resulted in the deepest temporal divergence in living human populations. Early modern human expansion in sub-Saharan Africa appears to have contributed to the end of late Acheulean (Fauresmith) industries at about 130,000 years ago, although very late coexistence of archaic and early modern humans, until as late as 12,000 years ago, has been argued for West Africa in particular.[35]

The ancestors of the modern

haplogroup L1-6 in central/eastern Africa, ancestral to everyone else. There was a significant back-migration of bearers of L0 towards eastern Africa between 120 and 75 kya.[note 6]

Expansion to Central Africa by the ancestors of the

Central African forager populations (African Pygmies) most likely took place before 130,000 years ago, and certainly before 60,000 years ago.[37][38][39][40][note 7]

The situation in

hybridization with H. sapiens in West Africa.[35]

Early northern Africa dispersal

Main article:
Northern Dispersal
Further information: Sahara pump theory
Overview map of the peopling of the world by early modern humans (numbers indicate dates in thousands of years ago [ka]).

Populations of Homo sapiens migrated to the Levant and to Europe[dubious – discuss] between 130,000 and 115,000 years ago, and possibly in earlier waves as early as 185,000 years ago.[note 8]

A fragment of a jawbone with eight teeth found at Misliya Cave has been dated to around 185,000 years ago. Layers dating from between 250,000 and 140,000 years ago in the same cave contained tools of the Levallois type which could put the date of the first migration even earlier if the tools can be associated with the modern human jawbone finds.[42][43]

These early migrations do not appear to have led to lasting colonisation and receded by about 80,000 years ago.[20] There is a possibility that this first wave of expansion may have reached China (or even North America[dubious – discuss][44]) as early as 125,000 years ago, but would have died out without leaving a trace in the genome of contemporary humans.[20]

millennia
before present).

There is some evidence that modern humans left Africa at least 125,000 years ago using two different routes: through the

southern China) teeth, which include a right upper second molar and a left lower second molar, indicate that the molars may be as old as 126,000 years.[55][56]

Since these previous exits from Africa did not leave traces in the results of genetic analyses based on the Y chromosome and on MtDNA, it seems that those modern humans did not survive in large numbers and were assimilated by our major antecessors. An explanation for their extinction (or small genetic imprint) may be the Toba eruption (74,000 years ago), though some argue it scarcely affected human population.[57]

Coastal migration

Main article: Southern Dispersal
Further information: Recent African origin of modern humans, Interbreeding between archaic and modern humans, and Archaic humans in Southeast Asia § Anatomically modern humans in southeast Asia
Overview map of the peopling of the world by early humans during the Upper Paleolithic, following the Southern Dispersal paradigm.

The so-called "recent dispersal" of modern humans took place about 70–50,000 years ago.[58][59][60] It is this migration wave that led to the lasting spread of modern humans throughout the world.

A small group from a population in East Africa, bearing

Persia to the South Asia before 55,000 years ago. Other research supports a migration out of Africa between about 65,000 and 50,000 years ago.[58][64][60] The coastal migration between roughly 70,000 and 50,000 years ago is associated with mitochondrial haplogroups M and N
, both derivative of L3.

Along the way H. sapiens interbred with Neanderthals and Denisovans,[65] with Denisovan DNA making 0.2% of mainland Asian and Native American DNA.[66]

Nearby Oceania

Migrations continued along the Asian coast to Southeast Asia and Oceania, colonising

Mamanwa, a Negrito people in the Philippines, suggesting the interbreeding took place in Eastern Asia where the Denisovans lived.[70][71][72] Denisovans may have crossed the Wallace Line, with Wallacea serving as their last refugium.[73][74] Homo erectus had crossed the Lombok gap reaching as far as Flores, but never made it to Australia.[75]

last glacial maximum
, 20,000 yrs ago and when the sea level was probably more than 110m lower than today.

During this time sea level was much lower and most of

haplogroup C
.

Sequencing of one Aboriginal genome from an old hair sample in Western Australia revealed that the individual was descended from people who migrated into East Asia between 62,000 and 75,000 years ago. This supports the theory of a single migration into Australia and New Guinea before the arrival of Modern Asians (between 25,000 and 38,000 years ago) and their later migration into North America.[77] This migration is believed to have happened around 50,000 years ago, before Australia and New Guinea were separated by rising sea levels approximately 8,000 years ago.[78][79] This is supported by a date of 50,000–60,000 years ago for the oldest evidence of settlement in Australia,[67][80] around 40,000 years ago for the oldest human remains,[67] the earliest humans artifacts which are at least 65,000 years old[81] and the extinction of the Australian megafauna by humans between 46,000 and 15,000 years ago argued by Tim Flannery,[82] which is similar to what happened in the Americas. The continued use of Stone Age tools in Australia has been much debated.[83]

Dispersal throughout Eurasia

Further information: Upper Paleolithic, Mammoth steppe, Interbreeding between archaic and modern humans, and Mousterian
Successive dispersals of
  Homo erectus greatest extent (yellow),
  Homo neanderthalensis greatest extent (ochre) and
  Homo sapiens (red).

The population brought to the

coastal migration appears to have remained there for some time, during roughly 60,000 to 50,000 years ago, before spreading further throughout Eurasia. This dispersal of early humans, at the beginning of the Upper Paleolithic, gave rise to the major population groups of the Old World and the Americas
.

Towards the West, Upper Paleolithic populations associated with mitochondrial haplogroup R and its derivatives, spread throughout Asia and Europe, with a back-migration of M1 to North Africa and the Horn of Africa several millennia ago. [dubious – discuss]

Presence

Neanderthal ancestry, but it seems likely that substantial interbreeding with Neanderthals ceased before 47,000 years ago, i.e. took place before modern humans entered Europe.[85]

There is evidence from

founder effects may have been maximised. The greater diversity amongst African genomes may reflect the extent of African refugia during the Toba incident.[87] However, a recent review highlights that the single-source hypothesis of non-African populations is less consistent with ancient DNA analysis than multiple sources with genetic mixing across Eurasia.[20]

Europe

Main articles:
Early European modern humans, Paleolithic Europe, Aurignacian, Gravettian, and Art of the Upper Paleolithic
Further information: Hominid dispersals in Europe
30,000-year-old cave lion and bison painting found in the Chauvet Cave, France.

The recent expansion of

interbred with Neanderthal populations
to a limited degree. Populations of modern humans and Neanderthal overlapped in various regions such as the Iberian peninsula and the Middle East. Interbreeding may have contributed Neanderthal genes to palaeolithic and ultimately modern Eurasians and Oceanians.

An important difference between Europe and other parts of the inhabited world was the northern latitude. Archaeological evidence suggests humans, whether Neanderthal or Cro-Magnon, reached sites in Arctic Russia by 40,000 years ago.[89]

Cro-Magnon are considered the first anatomically modern humans in Europe. They entered Eurasia by the Zagros Mountains (near present-day Iran and eastern Turkey) around 50,000 years ago, with one group rapidly settling coastal areas around the Indian Ocean and another migrating north to the steppes of Central Asia.[90] Modern human remains dating to 43,000–45,000 years ago have been discovered in Italy[91] and Britain,[92] as well as in the European Russian Arctic from 40,000 years ago.[89][93]

Humans colonised the environment west of the Urals, hunting reindeer especially,[94] but were faced with adaptive challenges; winter temperatures averaged from −20 to −30 °C (−4 to −22 °F) with fuel and shelter scarce. They travelled on foot and relied on hunting highly mobile herds for food. These challenges were overcome through technological innovations: tailored clothing from the pelts of fur-bearing animals; construction of shelters with hearths using bones as fuel; and digging "ice cellars" into the permafrost to store meat and bones.[94][95]

However, from recent research it is believed that the ecological crisis resulting from the eruption in c. 38,000 BC of the super-volcano in the

Phlegrean Fields near Naples, which left much of eastern Europe covered in ash, wiped out both the last Neanderthal and the first Homo Sapiens populations of the early Upper Paleolithic.[96][97] Modern Europeans of today bear no trace of the genomes of the first Homo Sapiens Europeans, but only of those from after the ecological crisis of 38,000 BC.[98] Modern humans then repopulated Europe from the east after the eruption and the ice age that took place from 38,000 to 36,000 BC.[99]

A mitochondrial DNA sequence of two Cro-Magnons from the Paglicci Cave in Italy, dated to 23,000 and 24,000 years old (Paglicci 52 and 12), identified the mtDNA as Haplogroup N, typical of the latter group.[100]

Migration of modern humans into Europe, based on simulation by Currat & Excoffier (2004)[101]
(YBP =
Years before present)
Up to 37,500 YBP
Up to 35,000 YBP
Up to 32,500 YBP
Up to 30,000 YBP
Further information: Neanderthal extinction and Interbreeding between archaic and modern humans

The expansion of modern human population is thought to have begun 45,000 years ago, and it may have taken 15,000–20,000 years for Europe to be colonized.[102][103]

During this time, the Neanderthals were slowly being displaced. Because it took so long for Europe to be occupied, it appears that humans and Neanderthals may have been constantly competing for territory. The Neanderthals had larger brains, and were larger overall, with a more robust or heavily built frame, which suggests that they were physically stronger than modern Homo sapiens. Having lived in Europe for 200,000 years, they would have been better adapted to the cold weather. The anatomically modern humans known as the

Iberian peninsula. Neanderthals disappeared about 40,000 years ago.[104]

From the extent of linkage disequilibrium, it was estimated that the last Neanderthal gene flow into early ancestors of Europeans occurred 47,000–65,000 years BP. In conjunction with archaeological and fossil evidence, interbreeding is thought to have occurred somewhere in Western Eurasia, possibly the Middle East.[85] Studies show a higher Neanderthal admixture in East Asians than in Europeans.[105][106] North African groups share a similar excess of derived alleles with Neanderthals as non-African populations, whereas Sub-Saharan African groups are the only modern human populations with no substantial Neanderthal admixture.[note 10] The Neanderthal-linked haplotype B006 of the dystrophin gene has also been found among nomadic pastoralist groups in the Sahel and Horn of Africa, who are associated with northern populations. Consequently, the presence of this B006 haplotype on the northern and northeastern perimeter of Sub-Saharan Africa is attributed to gene flow from a non-African point of origin.[note 11]

East, Southeast and North Asia

Ancient North Eurasian populations from Siberia were an important genetic contributor to Ancient Native Americans and Eastern European Hunter-Gatherers. Neolithic Iranian farmers and Jōmon people (ancestors of the Ainu people) also received geneflow from ANE-related populations.[109]

"

Neanderthal introgression of 18 genes within the chromosome 3p21.31 region (HYAL region) of East Asians. The introgressive haplotypes were positively selected in only East Asian populations, rising steadily from 45,000 years ago until a sudden increase of growth rate around 5,000 to 3,500 years ago. They occur at very high frequencies among East Asian populations in contrast to other Eurasian populations (e.g. European and South Asian populations). The findings also suggest that this Neanderthal introgression occurred within the ancestral population shared by East Asians and Native Americans.[111]

A 2016 study presented an analysis of the population genetics of the

EDAR gene, dated to c. 35,000 years ago.[note 12][note 13]

Further information:
Ancient North Eurasians

Mitochondrial haplogroups

Japan, by about 35,000 years ago. Parts of these populations migrated to North America during the Last Glacial Maximum
.

A review paper by Melinda A. Yang (in 2022) summarized and concluded that a distinctive "Basal-East Asian population" referred to as 'East- and Southeast Asian lineage' (ESEA); which is ancestral to modern East Asians,

Northern China, but already differentiated and distinct from European-related and Australasian-related lineages, found in other regions of prehistoric Eurasia. The ESEA lineage trifurcated from an earlier East-Eurasian or "eastern non-African" (ENA) meta-population, which also contributed to the formation of Ancient Ancestral South Indians (AASI) as well as to Australasians.[116]

Last Glacial Maximum

Eurasia

Further information: Solutrean and Magdalenian
Schematic illustration of the Beringia migration based on matrilineal genetics: Arrival of Central Asian populations to the Beringian Mammoth steppe c. 25,000 years ago, followed by a "swift peoplling of the Americas"[citation needed] c. 15,000 years ago.

Around 20,000 years ago, approximately 5,000 years after the Neanderthal extinction, the

Indo-European expansion. A Paleolithic site on the Yana River, Siberia, at 71°N, lies well above the Arctic Circle and dates to 27,000 radiocarbon years before present, during glacial times. This site shows that people adapted to this harsh, high-latitude, Late Pleistocene environment much earlier than previously thought.[117]

Americas

Main articles:
Genetic history of Indigenous peoples of the Americas

late glacial maximum.[118][119][120][121] Details of Paleo-Indian migration to and throughout the American continent, including the dates and the routes traveled, are subject to ongoing research and discussion.[122]

Conventional estimates have it that humans reached North America at some point between 15,000 and 20,000 years ago.

Cordilleran ice sheets.[127] Another route proposed is that, either on foot or using primitive boats, they migrated down the Pacific coast to South America as far as Chile.[128] Any archaeological evidence of coastal occupation during the last Ice Age would now have been covered by the sea level rise, up to a hundred metres since then.[129] The recent finding of indigenous Australasian genetic markers in Amazonia supports that a coastal route and subsequent isolation did occur with some migrants.[130]

Holocene migrations

Further information: Pre-modern human migration, Mesolithic, and Linguistic homeland
Prehistoric migration routes for Y-chromosome Haplogroup N lineage following the retreat of ice sheets after the Last Glacial Maximum (22–18 kya).[131]

The

glaciers
were now re-populated.

This period sees the transition from the

China
beginning around 4,000 years ago.

Large-scale migrations of the Mesolithic to Neolithic era are thought to have given rise to the pre-modern distribution of the world's major

Nostratic theory
postulates the derivation of the major language families of Eurasia (excluding Sino-Tibetan) from a single proto-language spoken at the beginning of the Holocene period.

Eurasia

Further information: Genetic history of Europe, Neolithic Revolution, Indo-European migrations, and Proto-Uralic homeland

Evidence published in 2014 from genome analysis of ancient human remains suggests that the modern native populations of Europe largely descend from three distinct lineages: "

Indo-European expansion.[132] The Ancient North Eurasian component was introduced to Western Europe by people related to the Yamnaya culture.[133] Additional ANE ancestry is found in European populations through Paleolithic interactions with Eastern Hunter-Gatherers.[134]

Sub-Saharan Africa

Further information:
Niger-Congo

West-Eurasian back-migrations started in the early

Central Sudanic peoples, were mostly agriculturalists.[136]

The

c. 3900 BCE.[137] The Bantu expansion has spread the Bantu languages to Central, Eastern and Southern Africa, partly replacing the indigenous populations of these regions, including the African Pygmies, Hadza people and San people
. Beginning about 3,000 years ago, it reached South Africa about 1,700 years ago.[138]

Some evidence (including a 2016 study by Busby et al.) suggests admixture from ancient and recent migrations from Eurasia into parts of Sub-Saharan Africa.[139] Another study (Ramsay et al. 2018) also shows evidence that ancient Eurasians migrated into Africa and that Eurasian admixture in modern Sub-Saharan Africans ranges from 0% to 50%, varying by region and generally higher in the Horn of Africa and parts of the Sahel zone, and found to a lesser degree in certain parts of Western Africa, and Southern Africa (excluding recent immigrants).[140]

Indo-Pacific

Further information: Austronesian peoples § Austronesian expansion
Austronesian expansion
.

The first seaborne human migrations were by the

Island Southeast Asia at around 3000 to 1500 BCE. From the Philippines and Eastern Indonesia they colonized Micronesia by 2200 to 1000 BCE.[141][142]

A branch of the Austronesians reached

Rapa Nui by 1000 CE, and New Zealand by 1200 CE.[142][143][144]

In the

Papuans and the Negritos in Island Southeast Asia.[141][142] The Austronesian expansion was the last and the most far-reaching Neolithic human migration event.[145]

Caribbean

The Caribbean was one of the last places in the Americas that were settled by humans. The oldest remains are known from the Greater Antilles (Cuba and Hispaniola) dating between 4000 and 3500 BCE, and comparisons between tool-technologies suggest that these peoples moved across the Yucatán Channel from Central America. All evidence suggests that later migrants from 2000 BCE and onwards originated from South America, via the Orinoco region.[146] The descendants of these migrants include the ancestors of the Taíno and Kalinago (Island Carib) peoples.[147]

Arctic

Main article: Circumpolar peoples
Map showing the decline of the Paleo-Eskimo Dorset culture and expansion of the Thule people (900 to 1500 CE).

The earliest inhabitants of North America's central and eastern Arctic are referred to as the Arctic small tool tradition (AST) and existed c. 2500 BCE. AST consisted of several Paleo-Eskimo cultures, including the Independence cultures and Pre-Dorset culture.[148][149]

The

gradually displaced the Dorset culture.[150][151]

See also

Notes

  1. ^ Based on Schlebusch et al., "Southern African ancient genomes estimate modern human divergence to 350,000 to 260,000 years ago",[1] Fig. 3 (H. sapiens divergence times) and Stringer (2012),[2] (archaic admixture).
  2. ^ Archaic admixture from various sources is known from Europe and Asia (Neanderthals), Southeast Asia and Melanesia (Denisovans) as well as from Western and Southern Africa. The proportion of admixture varies by region, but in all cases has been reported below 10%: In Eurasian mostly estimated at 1–4% (with a high estimate of 3.4–7.3% by Lohse (2014)[3]) in Melanesians estimated at 4–6% (Reich et al. (2010)).[4] Admixture of an unknown archaic hominin in Sub-Saharan African hunter-gatherer populations was estimated at 2% (Hammer et al. (2011)).[5]
  3. Java, is considered the latest known specimen of H. erectus. Formerly dated to as late as 50,000 to 40,000 years ago, a 2011 study pushed back the date of the extinction of H. e. soloensis to 143,000 years ago at the latest, more likely before 550,000 years ago.[14]
  4. ^ "Here we report the ages, determined by thermoluminescence dating, of fire-heated flint artefacts obtained from new excavations at the Middle Stone Age site of Jebel Irhoud, Morocco, which are directly associated with newly discovered remains of H. sapiens. A weighted average age places these Middle Stone Age artefacts and fossils at 315±34 thousand years ago. Support is obtained through the recalculated uranium series with electron spin resonance date of 286±32 thousand years ago for a tooth from the Irhoud 3 hominin mandible."[21]
  5. ^ Estimated split times given in the source cited (in kya): Human-Neanderthal: 530–690, Deep Human [H. sapiens]: 250–360, NKSP-SKSP: 150–190, Out of Africa (OOA): 70–120.[1]
  6. ^ "By ~130 ka two distinct groups of anatomically modern humans co-existed in Africa: broadly, the ancestors of many modern-day Khoe and San populations in the south and a second central/eastern African group that includes the ancestors of most extant worldwide populations. Early modern human dispersals correlate with climate changes, particularly the tropical African "megadroughts" of MIS 5 (marine isotope stage 5, 135–75 ka) which paradoxically may have facilitated expansions in central and eastern Africa, ultimately triggering the dispersal out of Africa of people carrying haplogroup L3 ~60 ka. Two south to east migrations are discernible within haplogroup L0. One, between 120 and 75 ka, represents the first unambiguous long-range modern human dispersal detected by mtDNA and might have allowed the dispersal of several markers of modernity. A second one, within the last 20 ka signalled by L0d, may have been responsible for the spread of southern click-consonant languages to eastern Africa, contrary to the view that these eastern examples constitute relicts of an ancient, much wider distribution."[36]
  7. ^ "We studied the branching history of Pygmy hunter–gatherers and agricultural populations from Africa and estimated separation times and gene flow between these populations. The model identified included the early divergence of the ancestors of Pygmy hunter–gatherers and farming populations ~60,000 years ago, followed by a split of the Pygmies' ancestors into the Western and Eastern Pygmy groups – 20,000 years ago."[41]
  8. ^ Early modern human presence outside of Africa has been proposed to date back to as early as 177,000 years ago.[34]
  9. ^ The authors of Liu (2010) seem to accept that the individual has African recent ascentry, but with Asian archaic human admixture.[50] See also Dennell (2010).[51] Brief comments at [52] and [53]
  10. ^ "We found that North African populations have a significant excess of derived alleles shared with Neandertals, when compared to sub-Saharan Africans. This excess is similar to that found in non-African humans, a fact that can be interpreted as a sign of Neandertal admixture. Furthermore, the Neandertal's genetic signal is higher in populations with a local, pre-Neolithic North African ancestry. Therefore, the detected ancient admixture is not due to recent Near Eastern or European migrations. Sub-Saharan populations are the only ones not affected by the admixture event with Neandertals."[107]
  11. ^ "Of 1,420 sub-Saharan chromosomes, only one copy of B006 was observed in Ethiopia, and five in Burkina Faso, one among the Rimaibe and four among the Fulani and Tuareg, nomad-pastoralists known for having contacts with northern populations (supplementary table S1, Supplementary Material online). B006 only occurrence at the northern and northeastern outskirts of sub-Saharan Africa is thus likely to be a result of gene flow from a non-African source."[108]
  12. ^ Traits affected by the mutation are sweat glands, teeth, hair shaft thickness and breast tissue.[113][114]
  13. EDAR, ADH1B, ABCC1, and ALDH2 in particular. The East Asian types of ADH1B are associated with rice domestication and would thus have arisen after the c. 11,000 years ago.[115]

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Further reading

External links
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