Early human migrations

Early human migrations are the earliest migrations and expansions of archaic and modern humans across continents and are believed to have begun approximately 2 million years ago with the out of Africa migration of 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 both Denisovans and Neanderthals. Early hominids were said to have "crossed land bridges that were eventually covered in water" (History Alive, pub. 2004, TCI).

Within Africa, Homo sapiens dispersed around the time of its speciation, roughly 300,000 years ago.[note 1][3] The recent African origin paradigm suggests that the anatomically modern humans outside of Africa descend from a population of Homo sapiens migrating from East Africa roughly 70–50,000 years ago and spreading along the southern coast of Asia and to Oceania by about 50,000 years ago. Modern humans spread 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.[4][5][6][7][3]

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

After the Last Glacial Maximum, North Eurasian populations migrated to the Americas about 20,000 years ago. Northern Eurasia was peopled after 12,000 years ago, in the beginning Holocene. Arctic Canada and Greenland were reached by the Paleo-Eskimo expansion around 4,000 years ago. Finally, Polynesia was peopled within the past 2,000 years in the last wave of the Austronesian expansion.

Early humans (before Homo sapiens)

The earliest humans developed out of australopithecine ancestors after about 3 million years ago, most likely in Eastern Africa, most likely in the area of the Kenyan Rift Valley, where the oldest known stone tools were 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.[11]

Homo erectus

Between 3 and 2 million years ago, Homo spread throughout East Africa and to Southern Africa (Telanthropus capensis), but not yet to West Africa. Around 1.9 million years ago, Homo erectus migrated out of Africa via the Levantine corridor and Horn of Africa to Eurasia. This migration has been proposed as being related to the operation of the Saharan pump, around 1.9 million years ago. 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?[12]), Ubeidiya in the Levant (1.5 Ma) and Dmanisi in the Caucasus (1.81 ± 0.03 Ma, p=0.05[13]).

China was populated as early as 1.66 Mya based on stone artifacts found in the Nihewan Basin.[14] 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.[15]

Southeast Asia (Java) was reached about 1.7 million years ago (Meganthropus). Western Europe was first populated around 1.2 million years ago (Atapuerca).[16]

Robert G. Bednarik has suggested that Homo erectus may have built rafts and sailed oceans, a theory that has raised some controversy.[17]

After H. erectus

One million years after its dispersal, H. erectus was diverging into new species. H. erectus is a chronospecies and was never extinct, so that its "late survival" is a matter of taxonomic convention. Late forms of H. erectus are thought to have survived until after about 0.5 million ago to 143,000 years ago at the latest,[note 3] with derived forms classified as H. antecessor in Europe around 800,000 years ago and H. heidelbergensis in Africa around 600,000 years ago. H. heidelbergensis in its turn spread across East Africa (H. rhodesiensis) and to Eurasia, where it gave rise to Neanderthals and Denisovans.

H. heidelbergensis, Neanderthals and Denisovans expanded north beyond the 50th parallel (Eartham Pit, Boxgrove 500kya, Swanscombe Heritage Park 400kya, Denisova Cave 50 kya). It has been suggested that late Neanderthals may even have reached the boundary of the Arctic, by c. 32,000 years ago, when they were being displaced from their earlier habitats by H. sapiens, based on 2011 excavations at the site of Byzovaya in the Urals (Komi Republic, 65.02°N 57.42°E / 65.02; 57.42).[19]

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.[10][20][21][22] 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.[23]

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.[24]

Homo sapiens

Human body

Dispersal throughout Africa

Homo sapiens (anatomically modern humans) are assumed to have emerged about 300,000 years ago based on thermoluminescence dating of artefacts and remains from Jebel Irhoud, Morocco, published in 2017.[note 4][26] The Florisbad Skull from Florisbad, South Africa, dated to about 259,000 years ago, has also been classified as early Homo sapiens.[27][28][29][30] 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 anatomically modern humans.[31]

In September 2019, scientists reported the computerized determination, based on 260 CT scans, of a virtual skull shape of the last common human ancestor to modern humans/H. sapiens, 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.[32][33]

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.[5][6][7][3]

Early modern humans expanded to Western Eurasia, Central, Western and Southern Africa from the time of their emergence. While early expansions to Eurasia appear not to have persisted,[34][24] 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 Khoi-San expanded to Southern Africa before 150,000 years ago, possibly as early as before 260,000 years ago,[note 5] 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. 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 West Africa is difficult to interpret due to a sparsity of fossil evidence. Homo sapiens seems to have reached the western Sahelian zone by 130 kya, while tropical West African sites associated with H. sapiens are known only from after 130 kya. Unlike elsewhere in Africa, archaic MSA sites appear to persist until very late, down to the Holocene boundary (12 kya), pointing to the possibility of late survival of archaic humans, and late hybridization with H. sapiens in West Africa.[35]

Early northern Africa dispersal

Populations of H. sapiens migrated to the Levant and to Europe between 130,000 and 115,000 years ago, and possibly in earlier waves as early as 185,000 years ago.[note 8] These early migrations do not appear to have led to lasting colonisation and receded by about 80,000 years ago.[24] There is a possibility that this first wave of expansion may have reached China (or even North America[42]) as early as 125,000 years ago, but would have died out without leaving a trace in the genome of contemporary humans.[24]

There is some evidence that modern humans left Africa at least 125,000 years ago using two different routes: through the Nile Valley heading to the Middle East, at least into modern Israel (Qafzeh: 120,000–100,000 years ago); and a second route through the present-day Bab-el-Mandeb Strait on the Red Sea (at that time, with a much lower sea level and narrower extension), crossing to the Arabian Peninsula and settling in places like the present-day United Arab Emirates (125,000 years ago)[43] and Oman (106,000 years ago),[44] and possibly reaching the Indian Subcontinent (Jwalapuram: 75,000 years ago). Although no human remains have yet been found in these three places, the apparent similarities between the stone tools found at Jebel Faya, those from Jwalapuram and some from Africa suggest that their creators were all modern humans.[45] These findings might give some support to the claim that modern humans from Africa arrived at southern China about 100,000 years ago (Zhiren Cave, Zhirendong, Chongzuo City: 100,000 years ago;[note 9] and the Liujiang hominid (Liujiang County): controversially dated at 139,000–111,000 years ago [50]). Dating results of the Lunadong (Bubing Basin, Guangxi, 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.[51][52] Since these previous exits from Africa did not leave traces in the results of genetic analyses based on the Y chromosome and on MtDNA (which represent only a small part of the human genetic material), 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 impacted human population.[53]

An Asia center of origin and dispersal for the mtDNA haplogroup L3 has also been hypothesized based on the fossil record, the similar coalescence dates of L3 and its Eurasian-distributed M and N derivative clades (~71 kya), the distant location in Southeast Asia of the oldest subclades of M and N, and the comparable age of the paternal haplogroup DE. After an initial Out-of-Africa migration of early anatomically modern humans around 125 kya, fully modern human L3-carrying females are thus proposed to have back-migrated from the maternal haplogroup's place of origin in Eurasia around 70 kya along with males bearing the paternal haplogroup E, which is also proposed to have originated in Eurasia. These new Eurasian lineages are then suggested to have largely replaced the old autochthonous male and female African lineages.[54]

Other research suggests that earlier waves of modern human migration out of Africa predating 70kya mostly became extinct (contributing about 2% to the ancestry only of some Oceanian peoples such as Papuans) and that instead modern Eurasians descend from a single Out-of-Africa migration occurring approximately 50,000-70,000 years ago[55][56][57][58] associated with the origin and expansion of maternal haplogroup L3 from Eastern Africa.[59][60]

Coastal migration

The so-called "recent dispersal" of modern humans has taken place after beginning about 70–50,000 years ago.[61][62][58] 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 mitochondrial haplogroup L3 and numbering possibly fewer than 1,000 individuals,[63][64] crossed the Red Sea strait at Bab el Mandib, to what is now Yemen, after around 75,000 years ago.[65] A recent review has also shown support for the northern route through Sinai/Israel/Syria (Levant).[24] Their descendants spread along the coastal route around Arabia and Persia to the Indian subcontinent before 55,000 years ago. Other research supports a migration out of Africa between about 65,000 and 50,000 years ago.[61][60][58] The coastal migration between roughly 70,000 and 50,000 years ago is associated with mitochondrial haplogroups M and N, both derivative of L3.

A fragment of a jawbone with eight teeth found at Misliya Cave, Israel, 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.[66][67]

Along the way H. sapiens interbred with Neanderthals and Denisovans,[68] with Denisovan DNA making 0.2% of mainland Asian and Native American DNA.[69]

Near Oceania

Migrations continued along the Asian coast to Southeast Asia and Oceania, colonising Australia by around 50,000 years ago.[70][71][72] By reaching Australia, H. sapiens for the first time expanded its habitat beyond that of H. erectus. Denisovan ancestry is shared by Melanesians, Australian Aborigines, and smaller scattered groups of people in Southeast Asia, such as the Mamanwa, a Negrito people in the Philippines, suggesting the interbreeding took place in Eastern Asia where the Denisovans lived.[73][74][75] Denisovans may have crossed the Wallace Line, with Wallacea serving as their last refugium.[76][77] Homo erectus had crossed the Lombok gap reaching as far as Flores, but never made it to Australia.[78]

During this time sea level was much lower and most of Maritime Southeast Asia formed one land mass known as Sunda. Migration continued Southeast on the coastal route to the straits between Sunda and Sahul, the continental land mass of present-day Australia and New Guinea. The gaps on the Weber Line are up to 90 km wide,[79] so the migration to Australia and New Guinea would have required seafaring skills. Migration also continued along the coast eventually turning northeast to China and finally reaching Japan before turning inland. This is evidenced by the pattern of mitochondrial haplogroups descended from haplogroup M, and in Y-chromosome 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.[80] 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.[81][82] This is supported by a date of 50,000–60,000 years ago for the oldest evidence of settlement in Australia,[70][83] around 40,000 years ago for the oldest human remains,[70] the earliest humans artifacts which are at least 65,000 years old[84] and the extinction of the Australian megafauna by humans between 46,000 and 15,000 years ago argued by Tim Flannery,[85] which is similar to what happened in the Americas. The continued use of stone age tools in Australia has been much debated.[86]

Dispersal throughout Eurasia

The population brought to South Asia by 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.

Presence in Europe is certain after 40,000 years ago, possibly as early as 43,000 years ago,[87] rapidly replacing the Neanderthal population. Contemporary Europeans have 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.[88]

There is evidence from mitochondrial DNA that modern humans have passed through at least one genetic bottleneck, in which genome diversity was drastically reduced. Henry Harpending has proposed that humans spread from a geographically restricted area about 100,000 years ago, the passage through the geographic bottleneck and then with a dramatic growth amongst geographically dispersed populations about 50,000 years ago, beginning first in Africa and thence spreading elsewhere.[89] Climatological and geological evidence suggests evidence for the bottleneck. The explosion of Toba, the largest volcanic eruption of the Quaternary, may have created a 1,000 year cold period, potentially reducing human populations to a few tropical refugia. It has been estimated that as few as 15,000 humans survived. In such circumstances genetic drift and founder effects may have been maximised. The greater diversity amongst African genomes may be reflect the extent of African refugia during the Toba incident.[90] 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.[24]


The recent expansion of anatomically modern humans reached Europe around 40,000 years ago from Central Asia and the Middle East, as a result of cultural adaption to big game hunting of sub-glacial steppe fauna.[91] Neanderthals were present both in the Middle East and in Europe, and the arriving populations of anatomically modern humans (also known as "Cro-Magnon" or European early modern humans) 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.[92]

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.[93] Modern human remains dating to 43–45,000 years ago have been discovered in Italy[94] and Britain,[95] as well as in the European Russian Arctic from 40,000 years ago.[92][96]

Humans colonised the environment west of the Urals, hunting reindeer especially,[97] 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.[97][98]

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.[99]

Migration of modern humans into Europe, based on simulation by Currat & Excoffier (2004)[100]
(YBP=Years before present)
Up to 37,500 YBP
Up to 35,000 YBP
Up to 32,500 YBP
Up to 30,000 YBP

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.[101][102]

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 Cro-Magnons, with widespread trade networks, superior technology and bodies likely better suited to running, would eventually completely displace the Neanderthals, whose last refuge was in the Iberian peninsula. After about 25,000 years ago the fossil record of the Neanderthals ends, indicating extinction. The last known population lived around a cave system on the remote south-facing coast of Gibraltar from 30,000 to 24,000 years ago.

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.[88] Studies show a higher Neanderthal admixture in East Asians than in Europeans.[103][104] 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 nomad 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 and North Asia

"Tianyuan Man", an individual who lived in China c. 40,000 years ago, showed substantial Neanderthal admixture. A 2017 study of the ancient DNA of Tianyuan Man found that the individual is related to modern Asian and Native American populations.[107] A 2013 study found 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 suggests that this Neanderthal introgression occurred within the ancestral population shared by East Asians and Native Americans.[108]

A 2016 study presented an analysis of the population genetics of the Ainu people of northern Japan as key to the reconstruction of the early peopling of East Asia. The Ainu were found to represent a more basal branch than the modern farming populations of East Asia, suggesting an ancient (pre-Neolithic) connection with northeast Siberians.[109] A 2013 study associated several phenotypical traits associated with Mongoloids with a single mutation of the EDAR gene, dated to c. 35,000 years ago.[note 12][note 13]

Mitochondrial haplogroups A, B and G originated about 50,000 years ago, and bearers subsequently colonized Siberia, Korea and Japan, by about 35,000 years ago. Parts of these populations migrated to North America during the Last Glacial Maximum.

Last Glacial Maximum


Around 20,000 years ago, approximately 5,000 years after the Neanderthal extinction, the Last Glacial Maximum forced northern hemisphere inhabitants to migrate to several shelters (refugia) until the end of this period. The resulting populations are presumed to have resided in such refuges during the LGM to ultimately reoccupy Europe, where archaic historical populations are considered their descendants. The composition of European populations was later altered by further migrations, notably the Neolithic expansion from the Middle East, and still later the Chalcolithic population movements associated with 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.[113]


Paleo-Indians originated from Central Asia, crossing the Beringia land bridge between eastern Siberia and present-day Alaska.[114] Humans lived throughout the Americas by the end of the last glacial period, or more specifically what is known as the late glacial maximum, no earlier than 23,000 years before present.[114][115][116][117] 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.[118]

The routes of migration are also debated. The traditional theory is that these early migrants moved when sea levels were significantly lowered due to the Quaternary glaciation,[115][118] following herds of now-extinct pleistocene megafauna along ice-free corridors that stretched between the Laurentide and Cordilleran ice sheets.[119] 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.[120] 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.[121] The recent finding of indigenous Australasian genetic markers in Amazonia supports the coastal route hypothesis.[122][123]

Holocene migrations

The Holocene is taken to begin 12,000 years ago, after the end of the Last Glacial Maximum. During the Holocene climatic optimum, beginning about 9,000 years ago, human populations which had been geographically confined to refugia began to migrate. By this time, most parts of the globe had been settled by H. sapiens; however, large areas that had been covered by glaciers were now re-populated.

This period sees the transition from the Mesolithic to the Neolithic stage throughout the temperate zone. The Neolithic subsequently gives way to the Bronze Age in Old World cultures and the gradual emergence of the historical record in the Near East and 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 language families such as the Niger-Congo, Nilo-Saharan, Afro-Asiatic, Uralic, Sino-Tibetan or Indo-European phyla. The speculative Nostratic theory postulates the derivation of the major language families of Eurasia (excluding Sino-Tibetan) from a single proto-languages spoken at the beginning of the Holocene period.


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: "Western Hunter-Gatherers", derivative of the Cro-Magnon population of Europe, Early European Farmers introduced to Europe from the Near East during the Neolithic Revolution and Ancient North Eurasians which expanded to Europe in the context of the Indo-European expansion.[125]

The Afroasiatic Urheimat has been placed in either Africa or Asia.

Sub-Saharan Africa

The Nilotic peoples are thought to be derived from an earlier undifferentiated Eastern Sudanic unity by the 3rd millennium BCE. The development of the Proto-Nilotes as a group may have been connected with their domestication of livestock. The Eastern Sudanic unity must have been considerably earlier still, perhaps around the 5th millennium BCE (while the proposed Nilo-Saharan unity would date to the Upper Paleolithic about 15kya). The original locus of the early Nilotic speakers was presumably east of the Nile in what is now South Sudan. The Proto-Nilotes of the 3rd millennium BCE were pastoralists, while their neighbors, the Proto-Central Sudanic peoples, were mostly agriculturalists.[126]

The Niger-Congo phylum is thought to have emerged around 6,000 years ago in West or Central Africa. Its expansion may have been associated with the expansion of Sahel agriculture in the African Neolithic period, following the desiccation of the Sahara in c. 3900 BCE.[127] The Bantu expansion has spread the Bantu languages to Central, Eastern and Southern Africa, partly replacing the indigenous populations of these regions. Beginning about 3,000 years ago, it reached South Africa about 1,700 years ago.[128]

Some evidence (including a 2016 study by Busby et al.) suggests admixture from ancient amd recent migrations from Eurasia into parts of Sub-Saharan Africa.[129] 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).[130]


The first seaborne human migrations were by the Austronesian peoples originating from Taiwan known as the "Austronesian expansion".[131] Using advanced sailing technologies like catamarans, outrigger boats, and crab claw sails, they built the first sea-going ships and rapidly colonized Island Southeast Asia at around 3000 to 1500 BCE. From the Philippines and Eastern Indonesia they colonized Micronesia by 2200 to 1000 BCE.[131][132]

A branch of the Austronesians reached Island Melanesia between 1600 to 1000 BCE, establishing the Lapita culture (named after the archaeological site in Lapita, New Caledonia, where their characteristic pottery was first discovered). They are the direct ancestors of the modern Polynesians. They ventured into Remote Oceania reaching Vanuatu, New Caledonia, and Fiji by 1200 BCE, and Samoa and Tonga by around 900 to 800 BCE. This was the furthest extent of the Lapita culture expansion. During a period of around 1,500 years, they gradually lost the technology for pottery (likely due to the lack of clay deposits in the islands), replacing it with carved wooden and bamboo containers. Back-migrations from the Lapita culture also merged back Island Southeast Asia in 1500 BCE, and into Micronesia at around 200 BCE. It was not until 700 CE when they started voyaging further into the Pacific Ocean, when they colonized the Cook Islands, the Society Islands, and the Marquesas. From there, they further colonized Hawaii by 900 CE, Rapa Nui by 1000 CE, and New Zealand by 1200 CE.[132][133][134]

In the Indian Ocean, Austronesians from Borneo also colonized Madagascar and the Comoros Islands by around 500 CE. Austronesians remain the dominant ethnolinguistic group of the islands of the Indo-Pacific, and were the first to establish a maritime trade network reaching as far west as East Africa and the Arabian peninsula. They assimilated earlier Pleistocene to early Holocene human overland migrations through Sundaland like the Papuans and the Negritos in Island Southeast Asia.[131][132] The Austronesian expansion was the last and the most far-reaching Neolithic human migration event.[135]


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 4000–3500 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. The descendants of these migrants include the ancestors of the Taíno and Kalinago (Island Carib) peoples.[136]


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.[137][138]

The Inuit are the descendants of the Thule culture, which emerged from western Alaska around AD 1000 and gradually displaced the Dorset culture.[139][140]

See also


  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);[8] in Melanesians estimated at 4–6% (Reich et al. (2010)).[9] Admixture of an unknown archaic hominin in Sub-Saharan African hunter-gatherer popultations was estimated at about 2% (Hammer et al. (2011)).[10]
  3. Homo erectus soloensis, found in 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 its extinction of H. e. soloensis to 143,000 years ago at the latest, more likely before 550,000 years ago.[18]
  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."[25]
  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.[46] See also Dennell (2010).[47] Brief comments at [48] and [49]
  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."[105]
  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."[106]
  12. Traits affected by the mutation are sweat glands, teeth, hair thickness and breast tissue.[110][111]
  13. East Asian genetics shows a number of concentrated alleles suggestive of selection pressures. This concerns the genes 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.[112]


  1. Schlebusch; et al. (3 November 2017). "Southern African ancient genomes estimate modern human divergence to 350,000 to 260,000 years ago". Science. 358 (6363): 652–655. Bibcode:2017Sci...358..652S. doi:10.1126/science.aao6266. PMID 28971970.
  2. Stringer, C (2012). "What makes a modern human". Nature. 485 (7396): 33–35. Bibcode:2012Natur.485...33S. doi:10.1038/485033a. PMID 22552077.
  3. Harvati, Katerina; et al. (10 July 2019). "Apidima Cave fossils provide earliest evidence of Homo sapiens in Eurasia". Nature. 571 (7766): 500–504. doi:10.1038/s41586-019-1376-z. PMID 31292546.
  4. Callaway, Ewen (2018). "Israeli fossils are the oldest modern humans ever found outside of Africa". Nature. 554 (7690): 15–16. Bibcode:2018Natur.554...15C. doi:10.1038/d41586-018-01261-5. PMID 29388957.
  5. Zimmer, Carl (10 July 2019). "A Skull Bone Discovered in Greece May Alter the Story of Human Prehistory – The bone, found in a cave, is the oldest modern human fossil ever discovered in Europe. It hints that humans began leaving Africa far earlier than once thought". The New York Times. Retrieved 11 July 2019.
  6. Staff (10 July 2019). "'Oldest remains' outside Africa reset human migration clock". Phys.org. Retrieved 10 July 2019.
  7. Delson, Eric (10 July 2019). "An early dispersal of modern humans from Africa to Greece – Analysis of two fossils from a Greek cave has shed light on early hominins in Eurasia. One fossil is the earliest known specimen of Homo sapiens found outside Africa; the other is a Neanderthal who lived 40,000 years later". Nature. 571 (7766): 487–488. doi:10.1038/d41586-019-02075-9. PMID 31337897.
  8. Lohse, K; Frantz, L.A.F. (2014). "Neandertal Admixture in Eurasia Confirmed by Maximum-Likelihood Analysis of Three Genomes". Genetics. 196 (4): 1241–1251. doi:10.1534/genetics.114.162396. PMC 3982695. PMID 24532731.
  9. "Genetic history of an archaic hominin group from Denisova Cave in Siberia". Nature. 468 (7327): 1053–1060. doi:10.1038/nature09710. PMID 21179161)
  10. Hammer, M.F.; Woerner, A.E.; Mendez, F.L.; Watkins, J.C.; Wall, J.D. (2011). "Genetic evidence for archaic admixture in Africa". Proceedings of the National Academy of Sciences. 108 (37): 15123–15128. Bibcode:2011PNAS..10815123H. doi:10.1073/pnas.1109300108. PMC 3174671. PMID 21896735.
  11. Zhu, Zhaoyu; Dennell, Robin; Huang, Weiwen; Wu, Yi; Qiu, Shifan; Yang, Shixia; Rao, Zhiguo; Hou, Yamei; Xie, Jiubing; Han, Jiangwei; Ouyang, Tingping (2018). "Hominin occupation of the Chinese Loess Plateau since about 2.1 million years ago". Nature. 559 (7715): 608–612. Bibcode:2018Natur.559..608Z. doi:10.1038/s41586-018-0299-4. PMID 29995848.
  12. Fleagle, J.G.; Shea, J.J.; Grine, F.E.; Baden, A.L.; Leakey, R.E, eds. (2010). Out of Africa I: The First Hominin Colonization of Eurasia. Vertebrate Paleobiology and Paleoanthropology. Springer. ISBN 9789048190355.
  13. Garcia, T.; Féraud, G.; Falguères, C.; de Lumley, H.; Perrenoud, C.; Lordkipanidze, D. (2010). "Earliest human remains in Eurasia: New 40Ar/39Ar dating of the Dmanisi hominid-bearing levels, Georgia". Quaternary Geochronology. 5 (4): 443–451. doi:10.1016/j.quageo.2009.09.012.
  14. R. Zhu et al. (2004), New evidence on the earliest human presence at high northern latitudes in northeast Asia.
  15. Rixiang Zhu; Zhisheng An; Richard Pott; Kenneth A. Hoffman (June 2003). "Magnetostratigraphic dating of early humans in China" (PDF). Earth-Science Reviews. 61 (3–4): 191–361. Bibcode:2003ESRv...61..191A. doi:10.1016/S0012-8252(02)00110-1. Archived from the original (PDF) on 2011-07-24.
  16. Hopkin M (2008-03-26). "Fossil find is oldest European yet". Nature News. doi:10.1038/news.2008.691.
  17. Bednarik RG (2003). "Seafaring in the Pleistocene". Cambridge Archaeological Journal. 13 (1): 41–66. doi:10.1017/S0959774303000039.
    ScienceNews summary
  18. Indriati, E; Swisher CC III; Lepre C; Quinn RL; Suriyanto RA (2011). "The Age of the 20 Meter Solo River Terrace, Java, Indonesia and the Survival of Homo erectus in Asia". PLoS ONE. 6 (6): e21562. Bibcode:2011PLoSO...621562I. doi:10.1371/journal.pone.0021562. PMC 3126814. PMID 21738710.
  19. Callaway, Ewen (12 May 2011). "An Arctic refuge for Neanderthals?". nature.com. Retrieved 2019-05-08.
  20. Xu, D.; et al. (2017). "Archaic Hominin Introgression in Africa Contributes to Functional Salivary MUC7 Genetic Variation". Molecular Biology and Evolution. 34 (10): 2704–2715. doi:10.1093/molbev/msx206. PMC 5850612. PMID 28957509.
  21. Callaway, E. (26 July 2012). "Hunter-gatherer genomes a trove of genetic diversity". Nature. doi:10.1038/nature.2012.11076.
  22. Lachance, J.; Vernot, B.; Elbers, C.C.; Ferwerda, B.; Froment, A.; Bodo, J.M.; et al. (2012). "Evolutionary History and Adaptation from High-Coverage Whole-Genome Sequences of Diverse African Hunter-Gatherers". Cell. 150 (3): 457–469. doi:10.1016/j.cell.2012.07.009. PMC 3426505. PMID 22840920.
  23. Dirks, Paul HGM; Roberts, Eric M.; Hilbert-Wolf, Hannah; Kramers, Jan D.; Hawks, John; Dosseto, Anthony; Duval, Mathieu; Elliott, Marina; Evans, Mary; Grün, Rainer; Hellstrom, John; Herries, Andy IR; Joannes-Boyau, Renaud; Makhubela, Tebogo V.; Placzek, Christa J.; Robbins, Jessie; Spandler, Carl; Wiersma, Jelle; Woodhead, Jon; Berger, Lee R. (2017-05-09). "The age of Homo naledi and associated sediments in the Rising Star Cave, South Africa" (PDF). eLife. 6. doi:10.7554/eLife.24231. PMC 5423772. PMID 28483040.
  24. Lopez, Saioa; van Dorp, Lucy; Hellenthal, Garrett (2016). "Human Dispersal Out of Africa: A Lasting Debate". Evolutionary Bioinformatics. 11s2 (Suppl 2): 57–68. doi:10.4137/EBO.S33489. ISSN 1176-9343. PMC 4844272. PMID 27127403.
  25. David Richter; et al. (8 June 2017). "The age of the hominin fossils from Jebel Irhoud, Morocco, and the origins of the Middle Stone Age". Nature. 546 (7657): 293–296. Bibcode:2017Natur.546..293R. doi:10.1038/nature22335. PMID 28593967.
  26. Smith TM, Tafforeau P, Reid DJ, et al. (April 2007). "Earliest evidence of modern human life history in North African early Homo sapiens". Proceedings of the National Academy of Sciences of the United States of America. 104 (15): 6128–33. Bibcode:2007PNAS..104.6128S. doi:10.1073/pnas.0700747104. PMC 1828706. PMID 17372199.
  27. Stringer, C. (2016). "The origin and evolution of Homo sapiens". Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences. 371 (1698): 20150237. doi:10.1098/rstb.2015.0237. PMC 4920294. PMID 27298468.
  28. Sample, Ian (7 June 2017). "Oldest Homo sapiens bones ever found shake foundations of the human story". The Guardian. Retrieved 7 June 2017.
  29. Hublin, Jean-Jacques; Ben-Ncer, Abdelouahed; Bailey, Shara E.; Freidline, Sarah E.; Neubauer, Simon; Skinner, Matthew M.; Bergmann, Inga; Le Cabec, Adeline; Benazzi, Stefano; Harvati, Katerina; Gunz, Philipp (2017). "New fossils from Jebel Irhoud, Morocco and the pan-African origin of Homo sapiens" (PDF). Nature. 546 (7657): 289–292. Bibcode:2017Natur.546..289H. doi:10.1038/nature22336. PMID 28593953.
  30. Scerri, M.L., et al., "Did Our Species Evolve in Subdivided Populations across Africa, and Why Does It Matter?" Trends in Ecology & Evolution 33(8), August 2018, 582594, doi:10.1016/j.tree.2018.05.005.
  31. Mcdougall, I.; Brown, H.; Fleagle, G. (Feb 2005). "Stratigraphic placement and age of modern humans from Kibish, Ethiopia". Nature. 433 (7027): 733–736. Bibcode:2005Natur.433..733M. doi:10.1038/nature03258. ISSN 0028-0836. PMID 15716951.
  32. Zimmer, Carl (10 September 2019). "Scientists Find the Skull of Humanity's Ancestor — on a Computer - By comparing fossils and CT scans, researchers say they have reconstructed the skull of the last common forebear of modern humans". The New York Times. Retrieved 10 September 2019.
  33. Mounier, Aurélien; Lahr, Marta (2019). "Deciphering African late middle Pleistocene hominin diversity and the origin of our species". Nature Communications. 10 (1): 3406. doi:10.1038/s41467-019-11213-w. PMC 6736881. PMID 31506422.
  34. Hershkovitz, et al. (26 Jan 2018). "The earliest modern humans outside Africa". Science. 359 (6374): 456–459. Bibcode:2018Sci...359..456H. doi:10.1126/science.aap8369. PMID 29371468.
  35. Scerri, Eleanor (2017). "The Stone Age Archaeology of West Africa". African History. doi:10.1093/acrefore/9780190277734.013.137. ISBN 9780190277734.
  36. Rito, T; Richards, MB; Fernandes, V; Alshamali, F; Cerny, V; Pereira, L; Soares, P (2013). "The first modern human dispersals across Africa". PLOS ONE. 8 (11): e80031. Bibcode:2013PLoSO...880031R. doi:10.1371/journal.pone.0080031. PMC 3827445. PMID 24236171.
  37. Jarvis JP, Scheinfeldt LB, Soi S, Lambert C, Omberg L, Ferwerda B, et al. (2012). "Patterns of Ancestry, Signatures of Natural Selection, and Genetic Association with Stature in Western African Pygmies". PLoS Genetics. 8 (4): e1002641. doi:10.1371/journal.pgen.1002641. PMC 3343053. PMID 22570615.
  38. López Herráez D, Bauchet M, Tang K, Theunert C, Pugach I, Li J, et al. (2009). "Genetic Variation and Recent Positive Selection in Worldwide Human Populations: Evidence from Nearly 1 Million SNPs". PLoS ONE. 4 (11): e7888. Bibcode:2009PLoSO...4.7888L. doi:10.1371/journal.pone.0007888. PMC 2775638. PMID 19924308.
  39. Tishkoff SA, Reed FA, Friedlaender FR, Ehret C, Ranciaro A, Froment A, Hirbo JB, Awomoyi AA, Bodo JM, Doumbo O, Ibrahim M, Juma AT, Kotze MJ, Lema G, Moore JH, Mortensen H, Nyambo TB, Omar SA, Powell K, Pretorius GS, Smith MW, Thera MA, Wambebe C, Weber JL, Williams SM (2009). "The Genetic Structure and History of Africans and African Americans". Science. 324 (5930): 1035–1044. Bibcode:2009Sci...324.1035T. doi:10.1126/science.1172257. PMC 2947357. PMID 19407144. (Supplementary Data)
  40. Quintana-Murci, et al. (2008). "Maternal traces of deep common ancestry and asymmetric gene flow between Pygmy hunter–gatherers and Bantu-speaking farmers". PNAS. 105 (5): 1596–601. Bibcode:2008PNAS..105.1596Q. doi:10.1073/pnas.0711467105. PMC 2234190. PMID 18216239. fig. 3.
  41. Patin E, Laval G, Barreiro LB, Salas A, Semino O, Santachiara-Benerecetti S, Kidd KK, Kidd JR, Van der Veen L, Hombert JM, Gessain A, Froment A, Bahuchet S, Heyer E, Quintana-Murci L (2009). "Inferring the Demographic History of African Farmers and Pygmy Hunter–Gatherers Using a Multilocus Resequencing Data Set". PLoS Genetics. 5 (4): e1000448. doi:10.1371/journal.pgen.1000448. PMC 2661362. PMID 19360089.
  42. Holen, Steven R. (2017). "A 130,000-year-old archaeological site in southern California, USA". Nature. 544 (7651): 479–483. Bibcode:2017Natur.544..479H. doi:10.1038/nature22065. PMID 28447646.
  43. Lawler, Andrew (2011). "Did Modern Humans Travel Out of Africa Via Arabia?". Science. 331 (6016): 387. Bibcode:2011Sci...331..387L. doi:10.1126/science.331.6016.387. PMID 21273459.
  44. "Trail of 'Stone Breadcrumbs' Reveals the Identity of One of the First Human Groups to Leave Africa". ScienceDaily. 2011-12-01. Retrieved 2019-05-08.
  45. Bower, Bruce (2011-01-27). "Hints of earlier human exit from Africa". Science News. Retrieved 2019-05-08.
  46. Liu, Wu; et al. (2010). "Human remains from Zhirendong, South China, and modern human emergence in East Asia". Proceedings of the National Academy of Sciences. 107 (45): 19201–19206. Bibcode:2010PNAS..10719201L. doi:10.1073/pnas.1014386107. PMC 2984215. PMID 20974952.
  47. Robin Dennell, Two interpretations of the Zhirendong mandible described by Liu and colleagues, Nature Volume: 468, 25 November 2010, pages: 512–513, doi:10.1038/468512a
  48. Modern Humans Emerged Far Earlier Than Previously Thought, Fossils from China Suggest, ScienceDaily (Oct. 25, 2010) https://www.sciencedaily.com/releases/2010/10/101025172924.htm
  49. Oldest Modern Human Outside of Africa Found: http://news.nationalgeographic.com/news/2010/10/101025-oldest-human-fossil-china-out-of-africa-science/
  50. Shena, Guanjun; et al. (2002). "U-Series dating of Liujiang hominid site in Guangxi, Southern China". Journal of Human Evolution. 43 (6): 817–829. doi:10.1006/jhev.2002.0601. PMID 12473485.
  51. "Lunadong fossils support theory of earlier dispersal of modern man". University of Hawaii at Mānoa. 2014-09-18. Retrieved 2019-05-08.
  52. Callaway, Ewen (2015). "Teeth from China reveal early human trek out of Africa". Nature. doi:10.1038/nature.2015.18566. Retrieved 2015-10-23.
  53. Balter, Michael (2010). "Of Two Minds About Toba's Impact". Science. 327 (5970): 1187–1188. Bibcode:2010Sci...327.1187B. doi:10.1126/science.327.5970.1187-a. PMID 20203021.
  54. Vicente M. Cabrera (2017). "Carriers of mitochondrial DNA macrohaplogroup L3 basic lineages migrated back to Africa from Asia around 70,000 years ago". bioRxiv 233502.CS1 maint: uses authors parameter (link)
  55. Appenzeller (2012).
  56. Finlayson (2009), p. 68.
  57. Pagani, Luca (13 October 2016). "Genomic analyses inform on migration events during the peopling of Eurasia". Nature. 538 (7624): 238–242. Bibcode:2016Natur.538..238P. doi:10.1038/nature19792. PMC 5164938. PMID 27654910.
  58. Haber M, Jones AL, Connel BA, Asan, Arciero E, Huanming Y, Thomas MG, Xue Y, Tyler-Smith C (June 2019). "A Rare Deep-Rooting D0 African Y-chromosomal Haplogroup and its Implications for the Expansion of Modern Humans Out of Africa". Genetics. 212 (4): 1421–1428. doi:10.1534/genetics.119.302368. PMC 6707464. PMID 31196864.
  59. Soares, P; Alshamali, F; Pereira, J. B; Fernandes, V; Silva, N. M; Afonso, C; Costa, M. D; Musilova, E; MacAulay, V; Richards, M. B; Cerny, V; Pereira, L (2011). "The Expansion of mtDNA Haplogroup L3 within and out of Africa". Molecular Biology and Evolution. 29 (3): 915–927. doi:10.1093/molbev/msr245. PMID 22096215.
  60. Vai S, Sarno S, Lari M, Luiselli D, Manzi G, Gallinaro M, Mataich S, Hübner A, Modi A, Pilli E, Tafuri MA, Caramelli D, di Lernia S (March 2019). "Ancestral mitochondrial N lineage from the Neolithic 'green' Sahara". Scientific Reports. 9 (1): 3530. Bibcode:2019NatSR...9.3530V. doi:10.1038/s41598-019-39802-1. PMC 6401177. PMID 30837540.
  61. Posth C, Renaud G, Mittnik M, Drucker DG, Rougier H, Cupillard C, et al. (2016). "Pleistocene Mitochondrial Genomes Suggest a Single Major Dispersal of Non-Africans and a Late Glacial Population Turnover in Europe". Current Biology. 26 (6): 827–833. doi:10.1016/j.cub.2016.01.037. PMID 26853362.
  62. Karmin M, Saag L, Vicente M, Wilson Sayres MA, Järve M, Talas UG, et al. (April 2015). "A recent bottleneck of Y chromosome diversity coincides with a global change in culture". Genome Research. 25 (4): 459–66. doi:10.1101/gr.186684.114. PMC 4381518. PMID 25770088.
  63. Zhivotovsky; Rosenberg, NA; Feldman, MW; et al. (2003). "Features of Evolution and Expansion of Modern Humans, Inferred from Genomewide Microsatellite Markers". American Journal of Human Genetics. 72 (5): 1171–1186. doi:10.1086/375120. PMC 1180270. PMID 12690579.
  64. Stix, Gary (2008). "The Migration History of Humans: DNA Study Traces Human Origins Across the Continents". Retrieved 14 June 2011.
  65. Armitage, Simon J.; Jasim, Sabah A.; Marks, Anthony E.; Parker, Adrian G.; Usik, Vitaly I.; Uerpmann, Hans-Peter (2011). "Hints Of Earlier Human Exit From Africa". Science. 331 (6016): 453–456. Bibcode:2011Sci...331..453A. doi:10.1126/science.1199113. PMID 21273486. Retrieved 2011-05-01.
  66. "Scientists discover oldest known modern human fossil outside of Africa: Analysis of fossil suggests Homo sapiens left Africa at least 50,000 years earlier than previously thought". ScienceDaily. Retrieved 2018-01-27.
  67. Ghosh, Pallab (2018). "Modern humans left Africa much earlier". BBC News. Retrieved 2018-01-27.
  68. Dennell, Robin; Petraglia, Michael D. (2012). "The dispersal of Homo sapiens across southern Asia: how early, how often, how complex?". Quaternary Science Reviews. 47: 15–22. Bibcode:2012QSRv...47...15D. doi:10.1016/j.quascirev.2012.05.002.
  69. Prüfer, Kay; et al. (2013). "The complete genome sequence of a Neanderthal from the Altai Mountains". Nature. 505 (7481): 43–49. Bibcode:2014Natur.505...43P. doi:10.1038/nature12886. PMC 4031459. PMID 24352235.
  70. Bowler, James M.; et al. (2003). "New ages for human occupation and climatic change at Lake Mungo, Australia". Nature. 421 (6925): 837–840. Bibcode:2003Natur.421..837B. doi:10.1038/nature01383. PMID 12594511.
  71. Wood R (2017-09-02). "Comments on the chronology of Madjedbebe". Australian Archaeology. 83 (3): 172–174. doi:10.1080/03122417.2017.1408545. ISSN 0312-2417.
  72. O'Connell JF, Allen J, Williams MA, Williams AN, Turney CS, Spooner NA, et al. (August 2018). "Homo sapiens first reach Southeast Asia and Sahul?". Proceedings of the National Academy of Sciences of the United States of America. 115 (34): 8482–8490. doi:10.1073/pnas.1808385115. PMC 6112744. PMID 30082377.
  73. Callaway, Ewen (22 September 2011), "First Aboriginal genome sequenced", Nature, doi:10.1038/news.2011.551
  74. Reich; et al. (2011), "Denisova Admixture and the First Modern Human Dispersals into Southeast Asia and Oceania", The American Journal of Human Genetics, 89 (4): 516–528, doi:10.1016/j.ajhg.2011.09.005, PMC 3188841, PMID 21944045
  75. Choi, Charles (22 September 2011). "Now-Extinct Relative Had Sex with Humans Far and Wide". LiveScience.
  76. Cooper A.; Stringer C.B. (2013). "Did the Denisovans Cross the Wallace Line". Science. 342 (6156): 321–333. Bibcode:2013Sci...342..321C. doi:10.1126/science.1244869. PMID 24136958.
  77. Salleh, Anna (October 18, 2013). "Humans dated ancient Denisovan relatives beyond the Wallace Line". ABC Science.
  78. First Mariners – Archaeology Magazine Archive. Archive.archaeology.org. Retrieved on 2013-11-16.
  79. "Pleistocene Sea Level Maps". Fieldmuseum.org. Retrieved 2010-09-23.
  80. Rasmussen, M; et al. (Oct 2011). "An Aboriginal Australian genome reveals separate human dispersals into Asia". Science. 334 (6052): 94–98. Bibcode:2011Sci...334...94R. doi:10.1126/science.1211177. PMC 3991479. PMID 21940856.
  81. Hudjashov G, Kivisild T, Underhill PA, et al. (May 2007). "Revealing the prehistoric settlement of Australia by Y chromosome and mtDNA analysis". Proc Natl Acad Sci USA. 104 (21): 8726–8730. Bibcode:2007PNAS..104.8726H. doi:10.1073/pnas.0702928104. PMC 1885570. PMID 17496137.
  82. Wade, Nicholas (2007-05-08). "From DNA Analysis, Clues to a Single Australian Migration". Australia: Nytimes.com. Retrieved 2011-05-01.
  83. Clarkson, Chris; Smith, Mike; Marwick, Ben; Fullagar, Richard; Wallis, Lynley A.; Faulkner, Patrick; Manne, Tiina; Hayes, Elspeth; Roberts, Richard G.; Jacobs, Zenobia; Carah, Xavier; Lowe, Kelsey M.; Matthews, Jacqueline; Florin, S. Anna (June 2015). "The archaeology, chronology and stratigraphy of Madjedbebe (Malakunanja II): A site in northern Australia with early occupation". Journal of Human Evolution. 83: 46–64. doi:10.1016/j.jhevol.2015.03.014. hdl:1773/33254. PMID 25957653.
  84. Clarkson, Chris; Jacobs, Zenobia; Marwick, Ben; Fullagar, Richard; Wallis, Lynley; Smith, Mike; Roberts, Richard G.; Hayes, Elspeth; Lowe, Kelsey; Carah, Xavier; Florin, S. Anna; McNeil, Jessica; Cox, Delyth; Arnold, Lee J.; Hua, Quan; Huntley, Jillian; Brand, Helen E. A.; Manne, Tiina; Fairbairn, Andrew; Shulmeister, James; Lyle, Lindsey; Salinas, Makiah; Page, Mara; Connell, Kate; Park, Gayoung; Norman, Kasih; Murphy, Tessa; Pardoe, Colin (19 July 2017). "Human occupation of northern Australia by 65,000 years ago" (PDF). Nature. 547 (7663): 306–310. Bibcode:2017Natur.547..306C. doi:10.1038/nature22968. hdl:2440/107043. PMID 28726833.
  85. Flannery, Tim (2002), "The Future Eaters: An Ecological History of the Australasian Lands and People" (Grove Press)
  86. Mellars, Paul (11 August 2006). "Going East: New Genetic and Archaeological Perspectives on the Modern Human Colonization of Eurasia". Science. 313 (5788): 796–800. Bibcode:2006Sci...313..796M. doi:10.1126/science.1128402. PMID 16902130.
  87. "Fossil Teeth Put Humans in Europe Earlier Than Thought". The New York Times. 2011-11-02.
  88. Sankararaman, S.; Patterson, N.; Li, H.; Pääbo, S.; Reich, D; Akey, J.M. (2012). "The Date of Interbreeding between Neandertals and Modern Humans". PLoS Genetics. 8 (10): e1002947. arXiv:1208.2238. Bibcode:2012arXiv1208.2238S. doi:10.1371/journal.pgen.1002947. PMC 3464203. PMID 23055938.
  89. Harpending, Henry; Cochran, Gregory (2009). The 10,000 Year Explosion (PDF). Basic Books. p. 214. ISBN 978-0-465-00221-4. Retrieved December 1, 2015.
  90. Ambrose, Stanley (1998). "Late Pleistocene human population bottlenecks, volcanic winter, and differentiation of modern humans". Journal of Human Evolution. 34 (6): 623–651. doi:10.1006/jhev.1998.0219. PMID 9650103.
  91. Oppenheimer, Stephen "Out of Eden: Peopling of the World" (Robinson; New Ed edition (March 1, 2012))
  92. Pavlov, P; Svendsen, JI; Indrelid, S (2001). "Human presence in the European Arctic nearly 40,000 years ago". Nature. 413 (6851): 64–67. Bibcode:2001Natur.413...64P. doi:10.1038/35092552. PMID 11544525.
  93. "Atlas of human journey: 45–40,000". The genographic project. National Geographic Society. 1996–2010. Retrieved 2019-05-08.
  94. Benazzi, S.; Douka, K.; Fornai, C.; Bauer, C.C.; Kullmer, O.; Svoboda, J.F.; Pap, I.; Mallegni, F.; Bayle, P.; Coquerelle, M.; Condemi, S.; Ronchitelli, A.; Harvati, K.; Weber, G.W. (2011). "Early dispersal of modern humans in Europe and implications for Neanderthal behaviour". Nature. 479 (7374): 525–528. Bibcode:2011Natur.479..525B. doi:10.1038/nature10617. PMID 22048311.
  95. Higham, T.; Compton, T.; Stringer, C.; Jacobi, R.; Shapiro, B.; Trinkaus, E.; Chandler, B.; Gröning, F.; Collins, C.; Hillson, S.; o'Higgins, P.; Fitzgerald, C.; Fagan, M. (2011). "The earliest evidence for anatomically modern humans in northwestern Europe". Nature. 479 (7374): 521–524. Bibcode:2011Natur.479..521H. doi:10.1038/nature10484. PMID 22048314.
  96. "Mamontovaya Kurya:an enigmatic, nearly 40000 years old Paleolithic site in the Russian Arctic" (PDF).
  97. Hoffecker, J. (2006). A Prehistory of the North: Human Settlements of the Higher Latitudes. New Jersey: Rutgers University Press. p. 101.
  98. Hoffecker, John F. (2002). Desolate landscapes: Ice-Age settlement in Eastern Europe. New Brunswick: Rutgers University Press. pp. 158–162, 217–233.
  99. Caramelli, D; Lalueza-Fox, C; Vernesi, C; Lari, M; Casoli, A; Mallegni, F; Chiarelli, B; Dupanloup, I; Bertranpetit, J; Barbujani, G; Bertorelle, G (May 2003). "Evidence for a genetic discontinuity between Neandertals and 24,000-year-old anatomically modern Europeans". PNAS. 100 (11): 6593–6597. Bibcode:2003PNAS..100.6593C. doi:10.1073/pnas.1130343100. ISSN 0027-8424. PMC 164492. PMID 12743370.
  100. Currat M, Excoffier L (2004). "Modern Humans Did Not Admix with Neanderthals during Their Range Expansion into Europe". PLoS Biology. 2 (12): e421. doi:10.1371/journal.pbio.0020421. PMC 532389. PMID 15562317.
  101. Maca-Meyer N, González AM, Larruga JM, Flores C, Cabrera VM (2001). "Major genomic mitochondrial lineages delineate early human expansions". BMC Genet. 2 (1): 13. doi:10.1186/1471-2156-2-13. PMC 55343. PMID 11553319.
  102. Currat M, Excoffier L (Dec 2004). "Modern humans did not admix with Neanderthals during their range expansion into Europe". PLoS Biology. 2 (12): e421. doi:10.1371/journal.pbio.0020421. PMC 532389. PMID 15562317.
  103. Meyer, M.; Kircher, M.; Gansauge, M.T.; Li, H.; Racimo, F.; Mallick, S.; et al. (2012). "A High-Coverage Genome Sequence from an Archaic Denisovan Individual". Science. 338 (6104): 222–226. Bibcode:2012Sci...338..222M. doi:10.1126/science.1224344. PMC 3617501. PMID 22936568.
  104. Wall, J.D.; Yang, M.A.; Jay, F.; Kim, S.K.; Durand, E.Y.; Stevison, L.S.; et al. (2013). "Higher Levels of Neanderthal Ancestry in East Asians than in Europeans". Genetics. 194 (1): 199–209. doi:10.1534/genetics.112.148213. PMC 3632468. PMID 23410836.
  105. Sánchez-Quinto, F.; Botigué, L.R.; Civit, S.; Arenas, C.; Ávila-Arcos, M.C.; Bustamante, C.D.; et al. (2012). "North African Populations Carry the Signature of Admixture with Neandertals". PLoS ONE. 7 (10): e47765. Bibcode:2012PLoSO...747765S. doi:10.1371/journal.pone.0047765. PMC 3474783. PMID 23082212.
  106. Yotova, Vania; et al. (2011). "An X-linked haplotype of Neandertal origin is present among all non-African populations". Molecular Biology and Evolution. 28 (7): 1957–1962. doi:10.1093/molbev/msr024. PMID 21266489.
  107. Yang et al., "40,000-Year-Old Individual from Asia Provides Insight into Early Population Structure in Eurasia", Current Biology Volume 27, Issue 20, pp. 3202–3208.e9, 23 October 2017, doi:10.1016/j.cub.2017.09.030.
  108. Ding, Q.; Hu, Y.; Xu, S.; Wang, J.; Jin, L. (2014) [Online 2013]. "Neanderthal Introgression at Chromosome 3p21.31 was Under Positive Natural Selection in East Asians". Molecular Biology and Evolution. 31 (3): 683–695. doi:10.1093/molbev/mst260. PMID 24336922..
  109. Jeong et al., "Deep History of East Asian Populations Revealed Through Genetic Analysis of the Ainu", Genetics. 2016 Jan;202(1):261–272. doi:10.1534/genetics.115.178673.
  110. Kamberov, Yana G (14 February 2013). "Modeling Recent Human Evolution in Mice by Expression of a Selected EDAR Variant". Cell. 152 (4): 691–702. doi:10.1016/j.cell.2013.01.016. PMC 3575602. PMID 23415220.
  111. Wade, Nicholas (14 February 2013). "East Asian Physical Traits Linked to 35,000-Year-Old Mutation". The New York Times. Retrieved 2019-05-06.
  112. Peng, Y (2010). "The ADH1B ARG47His polymorphism in East Asian populations and expansion of rice domestication in history". BMC Evolutionary Biology. 10 (15): 15. doi:10.1186/1471-2148-10-15. PMC 2823730. PMID 20089146.
  113. Pitulko, V.V.; Nikolsky, P.A.; Girya, E.Y.; Basilyan, A.E.; Tumskoy, V.E.; Koulakov, S.A.; Astakhov, S.N.; Pavlova, E.Y.; Anisimov, M.A. (2004). "The Yana RHS Site: Humans in the Arctic Before the Last Glacial Maximum". Science. 303 (5654): 52–56. Bibcode:2004Sci...303...52P. doi:10.1126/science.1085219. PMID 14704419.
  114. Wells, Spencer; Read, Mark (2002). The Journey of Man – A Genetic Odyssey. Random House. pp. 138–140. ISBN 978-0-8129-7146-0.
  115. Fitzhugh, Drs. William; Goddard, Ives; Ousley, Steve; Owsley, Doug; Stanford, Dennis. "Paleoamerican". Smithsonian Institution Anthropology Outreach Office. Archived from the original on 2009-01-05. Retrieved 2009-01-15.
  116. "A DNA Search for the First Americans Links Amazon Groups to Indigenous Australians". Smithsonian Institution. Retrieved 2019-05-08.
  117. Bonatto, S. L.; Salzano, F. M. (1997). "A single and early migration for the peopling of greater America supported by mitochondrial DNA sequence data". Proceedings of the National Academy of Sciences. 94 (5): 1866–1871. Bibcode:1997PNAS...94.1866B. doi:10.1073/pnas.94.5.1866. PMC 20009. PMID 9050871.
  118. "Atlas of the Human Journey-The Genographic Project". National Geographic Society. 1996–2008. Archived from the original on 2011-05-01. Retrieved 2017-01-27.
  119. "The peopling of the Americas: Genetic ancestry influences health". Scientific American. Retrieved 2019-05-08.
  120. Fladmark, K. R. (1979). "Alternate Migration Corridors for Early Man in North America". American Antiquity. 44 (1): 55–69. doi:10.2307/279189. JSTOR 279189.
  121. "68 Responses to "Sea will rise 'to levels of last Ice Age'"". Center for Climate Systems Research, Columbia University. Retrieved 2009-11-17.
  122. accessed 26th October 2015 Genetic studies link indigenous peoples in the Amazon and Australasia Science Daily, July 21, 2015.
  123. Callaway, Ewen (2015). "'Ghost population' hints at long-lost migration to the Americas: Present-day Amazonians share an unexpected genetic link with Asian islanders, hinting at an ancient trek". Nature News. doi:10.1038/nature.2015.18029.
  124. Su, Bing; Ma, Runlin Z.; Zhang, Xiaoming; Peng, Yi; Zhong, Hua; Qi, Xuebin; Shi, Hong (20 June 2013). "Genetic Evidence of an East Asian Origin and Paleolithic Northward Migration of Y-chromosome Haplogroup N". PLOS ONE. 8 (6): e66102. Bibcode:2013PLoSO...866102S. doi:10.1371/journal.pone.0066102. ISSN 1932-6203. PMC 3688714. PMID 23840409.
  125. Gibbons, Ann (4 September 2014). "Three-part ancestry for Europeans". Science. American Association for the Advancement of Science. Archived from the original on 11 October 2014. Retrieved 30 October 2014.
  126. John Desmond Clark, From Hunters to Farmers: The Causes and Consequences of Food Production in Africa, University of California Press, 1984, p. 31
  127. Igor Kopytoff, The African Frontier: The Reproduction of Traditional African Societies (1989), 9–10 (cited afer Igbo Language Roots and (Pre)-History, A Mighty Tree, 2011).
  128. Vansina, J. (1995). "New Linguistic Evidence and 'The Bantu Expansion'". Journal of African History. 36 (2): 173–195. doi:10.1017/S0021853700034101. JSTOR 182309.
  129. Busby, George BJ; Band, Gavin; Si Le, Quang; Jallow, Muminatou; Bougama, Edith; Mangano, Valentina D; Amenga-Etego, Lucas N; Enimil, Anthony; Apinjoh, Tobias (2016). "Admixture into and within sub-Saharan Africa". eLife. 5. doi:10.7554/eLife.15266. ISSN 2050-084X. PMC 4915815. PMID 27324836.
  130. Ramsay, Michèle; Hazelhurst, Scott; Sengupta, Dhriti; Aron, Shaun; Choudhury, Ananyo (2018-08-01). "African genetic diversity provides novel insights into evolutionary history and local adaptations". Human Molecular Genetics. 27 (R2): R209–R218. doi:10.1093/hmg/ddy161. ISSN 0964-6906. PMC 6061870. PMID 29741686.
  131. Meacham, William (1984–1985). "On the improbability of Austronesian origins in South China" (PDF). Asian Perspective. 26: 89–106.
  132. Bellwood, Peter (1991). "The Austronesian Dispersal and the Origin of Languages". Scientific American. 265 (1): 88–93. Bibcode:1991SciAm.265a..88B. doi:10.1038/scientificamerican0791-88. JSTOR 24936983.
  133. Heath, Helen; Summerhayes, Glenn R.; Hung, Hsiao-chun (2017). "Enter the Ceramic Matrix: Identifying the Nature of the Early Austronesian Settlement in the Cagayan Valley, Philippines". In Piper, Philip J.; Matsumara, Hirofumi; Bulbeck, David (eds.). New Perspectives in Southeast Asian and Pacific Prehistory. terra australis. 45. ANU Press. ISBN 9781760460952.
  134. Carson, Mike T.; Hung, Hsiao-chun; Summerhayes, Glenn; Bellwood, Peter (January 2013). "The Pottery Trail From Southeast Asia to Remote Oceania". The Journal of Island and Coastal Archaeology. 8 (1): 17–36. doi:10.1080/15564894.2012.726941.
  135. Lansing, Steve. "Did a butterfly effect change the history of the Pacific?". Stockholm Resilience Centre. Stockholm University. Retrieved 9 November 2019.
  136. Fagan, B.M. (2007). People of the earth: An introduction to world prehistory. Upper Saddle River, NJ: Pearson Prentice Hall
  137. Hoffecker, John F. (2005). A prehistory of the north: human settlement of the higher latitudes. Rutgers University Press. p. 130. ISBN 978-0-8135-3469-5.
  138. Gibbon, pp. 28–31
  139. Rigby, Bruce. "101. Qaummaarviit Historic Park, Nunavut Handbook" (PDF). Archived from the original (PDF) on 2006-05-29. Retrieved 2009-10-02.
  140. Wood, Shannon Raye (April 1992). "Tooth Wear and the Sexual Division of Labour in an Inuit Population" (PDF). Department of Archaeology University of Saskatchewan. Simon Fraser University. Retrieved 2009-10-02.

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