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 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]
Key sites for this early migration out of Africa are Riwat in Pakistan (~2 Ma?), Ubeidiya in the Levant (1.5 Ma) and Dmanisi in the Caucasus (1.81 ± 0.03 Ma, p=0.05).
China was populated as early as 1.66 Mya based on stone artifacts found in the Nihewan Basin. 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.
Spread of Denisovans and Neanderthals after 500,000 years ago
Known Neanderthal range with separate populations in 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 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.
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.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.
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.
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.
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, 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.
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]
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.
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. There is a possibility that this first wave of expansion may have reached China (or even North America[dubious – discuss]) as early as 125,000 years ago, but would have died out without leaving a trace in the genome of contemporary humans.
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) and Oman (106,000 years ago), 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. 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 ). 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. 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.
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.
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 associated with the expansion of maternal haplogroup L3.
The so-called "recent dispersal" of modern humans has taken place after beginning about 70,000 years ago. 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, crossed the Red Sea strait at Bab el Mandib, to what is now Yemen, after around 75,000 years ago. A recent review has also shown support for the northern route through Sinai/Israel/Syria (Levant). Their descendants spread along the coastal route around Arabia and Persia to the Indian subcontinent before 55,000 years ago. 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.
Along the way H. sapiens interbred with Neanderthals and Denisovans, with Denisovan DNA making 0.2% of mainland Asian and Native American DNA.
Migrations continued along the Asian coast to Southeast Asia and Oceania, colonising Australia before 50,000 years ago. 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. Denisovans may have crossed the Wallace Line, with Wallacea serving as their last refugium. Homo erectus had crossed the Lombok gap reaching as far as Flores, but never made it to Australia.
The map shows the probable extent of land and water at the time of the last glacial maximum, 20,000 yrs ago and when the sea level was probably more than 110m lower than today.
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. 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. This is supported by a date of 50,000–60,000 years ago for the oldest evidence of settlement in Australia, around 40,000 years ago for the oldest human remains, the earliest humans artifacts which are at least 65,000 years old and the extinction of the Australian megafauna by humans between 46,000 and 15,000 years ago argued by Tim Flannery, which is similar to what happened in the Americas. The continued use of stone age tools in Australia has been much debated.
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, 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.
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. 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. 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.
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.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.
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. Modern human remains dating to 43–45,000 years ago have been discovered in Italy and Britain, as well as in the European Russian Arctic from 40,000 years ago.
Humans colonised the environment west of the Urals, hunting reindeer especially, 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.
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.
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. Studies show a higher Neanderthal admixture in East Asians than in Europeans. 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]
"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. 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.
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. 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.
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.
Prehistoric migration routes for Y-chromosome Haplogroup N lineage following the retreat of ice sheets after the Last Glacial Maximum (22–18 kya).
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.
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.
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.
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.
The islands of the Pacific were populated between c. 1600 BCE and 1000 CE.
The Lapita people, who got their name from the archaeological site in Lapita, New Caledonia, where their characteristic pottery was first discovered, were an Austronesian-speaking people who settled in Near Oceania (notably the Bismarck Archipelago in Papua New Guinea, and the Solomon Islands) around 1500 BCE, where some intermingling with the existing Papuan population took place. Acquiring long distance voyaging skills, they ventured into 'Remote Oceania', probably settling Vanuatu and New Caledonia around 1200 BCE, then Fiji, Samoa and Tonga. By the beginning of the 1st millennium BCE, this western part of Polynesia was a loose web of thriving populations settled on the islands' coasts and living off the sea. By 0 CE Micronesia was completely colonized; tropical eastern Polynesia, including Tahiti, was probably settled by 700 CE. The last region of Polynesia to be reached was New Zealand, probably by 1300 CE.
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.
^Based on Schlebusch et al., "Southern African ancient genomes estimate modern human divergence to 350,000 to 260,000 years ago",Fig. 3 (H. sapiens divergence times) and Stringer (2012), (archaic admixture).
^Archaic admixture from different 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); in Melanesians estimated at 4–6% (Reich et al. (2010)). Admixture of an unknown archaic hominin in Sub-Saharan African hunter-gatherer popultations was estimated at about 2% (Hammer et al. (2011)).
^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.
^"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."
^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.
^"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."
^"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."
^Early modern human presence outside of Africa has been proposed to date back to as early as 177,000 years ago.
^The authors of Liu (2010) seem to accept that the individual has African recent ascentry, but with Asian archaic human admixture. See also Dennell (2010). Brief comments at  and 
^"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."
^"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."
^Traits affected by the mutation are sweat glands, teeth, hair thickness and breast tissue.
^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.
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^R. Zhu et al. (2004), New evidence on the earliest human presence at high northern latitudes in northeast Asia.
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