Early Human Migrations

3/6/2021 Early human migrations - Wikipedia
Early human migrations

Early human migrations are the earliest
migrations and expansions of archaic and
modern humans across continents. They are
believed to have begun approximately
2 million years ago with the early expansions
out of Africa by Homo erectus. This initial
migration was followed by other archaic
humans including H. heidelbergensis, which
lived around 500,000 years ago and was the
likely ancestor of Denisovans and
Neanderthals as well as modern humans. Early
hominids likely "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 Putative migration waves out of Africa and back migrations
300,000 years ago.[note 1][3] The recent African into the continent, as well as the locations of major ancient
origin paradigm suggests that the anatomically human remains and archeological sites (López et
modern humans outside of Africa descend al.2015).
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 earlier local
populations, so that contemporary human populations are descended in small part (below 10%
contribution) from regional varieties of archaic humans.[note 2]
After the 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.
Contents
Early humans (before Homo sapiens)
Homo erectus
After H. erectus
Homo sapiens
Dispersal throughout Africa
Early northern Africa dispersal
Coastal migration
https://en.wikipedia.org/wiki/Early_human_migrations#Near_Oceania 1/26
Near Oceania
Dispersal throughout Eurasia
Europe
East and North Asia
Last Glacial Maximum
Eurasia
Americas
Holocene migrations
Eurasia
Sub-Saharan Africa
Indo-Pacific
Caribbean
Arctic
See also
Notes
References
Further reading
External links
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 2 and less than a million years ago, Homo spread throughout East Africa and to Southern
Africa (Telanthropus capensis), but not yet to West Africa. Around 1.8 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 shows evidence of Erectus from 2.12 Mya in Gongwangling in Lantian county.[14] Two Homo
erectus incisors have been found near Yuanmou, south China, and are dated to 1.7 Mya, and a
cranium from Lantian has been dated to 1.63 Ma. Artefacts from Majuangou III and Shangshazui in
the Nihewan basin, north China, have been dated to 1.6–1.7 Ma. [15] Artifacts dated to as early as 1.66
Mya were found in the Nihewan Basin.[16] 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.[17]
Southeast Asia (Java) was reached about 1.7 million years ago (Meganthropus). Western Europe was
first populated around 1.2 million years ago (Atapuerca).[18]
Robert G. Bednarik has suggested that Homo erectus may have built rafts and sailed oceans, a theory
that has raised some controversy.[19]
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 Spread of Denisovans and
Africa around 600,000 years ago. H. heidelbergensis in its turn Neanderthals after 500,000 years
spread across East Africa (H. rhodesiensis) and to Eurasia, where ago
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 Known Neanderthal range with
Urals (Komi Republic, 65.02°N 57.42°E).[21] separate populations in Europe and
the Caucasus (blue), the Near East
Other archaic human species are assumed to have spread (orange), Uzbekistan (green), and
throughout Africa by this time, although the fossil record is the Altai region (purple)
sparse. Their presence is assumed based on traces of admixture
with modern humans found in the genome of African
populations.[10][22][23][24] 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.[25]
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.[26] Neanderthal
evidence has also been found quite late as of 33,000 years ago at the almost 65th latitude of the
Byzovaya site in the Ural Mountains. This is far outside of otherwise known habitat, during a high ice
cover period, and perhaps reflects a refugia of near extinction.
Homo sapiens
Dispersal throughout Africa
Homo sapiens are believed to have emerged in

Africa about 300,000 years ago, based in part on
thermoluminescence dating of artefacts and
remains from Jebel Irhoud, Morocco, published in
2017.[note 4][28] The Florisbad Skull from Florisbad,
South Africa, dated to about 259,000 years ago,
has also been classified as early Homo Early modern human migrations based on the
sapiens.[29][30][31][32] Previously, the Omo distribution of mitochondrial haplogroups.
remains, excavated between 1967 and 1974 in Omo National Park, Ethiopia, and dated to 200,000
years ago, were long held to be the oldest known fossils of Homo sapiens.[33]
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 anatomically modern humans,
representative of the earliest modern humans, and suggested that modern humans arose between
260,000 and 350,000 years ago through a merging of populations in East and South Africa.[34][35]
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 and Central, Western and Southern Africa from
the time of their emergence. While early expansions to Eurasia appear not to have persisted,[36][26]
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.[37]
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.[39][40][41][42][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 130,000 years ago, while tropical West African
sites associated with H. sapiens are known only from after 130,000 years ago. Unlike elsewhere in
Africa, archaic Middle Stone Age sites appear to persist until very late, down to the Holocene
boundary (12,000 years ago), pointing to the possibility of late survival of archaic humans, and late
hybridization with H. sapiens in West Africa.[37]
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]
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.[44][45]
These early migrations do not appear to have led to lasting colonisation and receded by about 80,000
years ago.[26] There is a possibility that this first wave of expansion may have reached China (or even
North America[46]) as early as 125,000 years ago, but would have died out without leaving a trace in
the genome of contemporary humans.[26]
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 Palestine (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[47][48] and settling in places like the present-day United Arab Emirates (125,000
years ago)[49] and Oman (106,000 years ago),[50] 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.[51] 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 [56]). 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.[57][58] 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.[59]
Coastal migration
The so-called "recent dispersal" of modern

humans has taken place after beginning about
70–50,000 years ago.[60][61][62] 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 Overview map of the peopling of the world by early
at Bab el Mandib, to what is now Yemen, after humans during the Upper Paleolithic, following to the
around 75,000 years ago. [65] A recent review Southern Dispersal paradigm.
has also shown support for the northern route
through Sinai/Israel/Syria (Levant).[26] 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.[60][66][62] The coastal migration between roughly 70,000 and 50,000 years
ago is associated with mitochondrial haplogroups M and N, both derivative of L3.
Along the way H. sapiens interbred with Neanderthals and Denisovans,[67] with Denisovan DNA
making 0.2% of mainland Asian and Native American DNA.[68]
Near Oceania
Migrations continued along the Asian coast to Southeast Asia and Oceania, colonising Australia by
around 50,000 years ago.[69][70][71] 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.[72][73][74] Denisovans may have crossed the Wallace Line, with Wallacea serving as
their last refugium.[75][76] Homo erectus had crossed the Lombok gap reaching as far as Flores, but
never made it to Australia.[77]
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,[78] 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 The map shows the probable extent of
sample in Western Australia revealed that the individual was land and water at the time of the last
descended from people who migrated into East Asia between glacial maximum, 20,000 yrs ago and
when the sea level was probably more
62,000 and 75,000 years ago. This supports the theory of a
than 110m lower than today.
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.[79] 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.[80][81] This is supported by a date
of 50,000–60,000 years ago for the oldest evidence of settlement in Australia,[69][82] around 40,000
years ago for the oldest human remains,[69] the earliest humans artifacts which are at least 65,000
years old[83] and the extinction of the Australian megafauna by humans between 46,000 and 15,000
years ago argued by Tim Flannery,[84] which is similar to what happened in the Americas. The
continued use of Stone Age tools in Australia has been much debated.[85]
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,[86]
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.[87]
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.[88] 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 reflect the extent of
African refugia during the Toba incident.[89] 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.[26]
Europe
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.[90] 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.[91]
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.[92] Modern human remains dating to 43–45,000 years ago have been
discovered in Italy[93] and Britain,[94] as well as in the European Russian Arctic from 40,000 years
ago.[91][95]
Humans colonised the environment west of the Urals, hunting reindeer especially,[96] 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.[96][97]
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.[98]
Migration of modern humans into Europe, based on simulation by Currat & Excoffier (2004)[99]
(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.[100][101]
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.[87] Studies show a higher Neanderthal admixture in East Asians than in
Europeans.[102][103] North African groups share a similar excess of derived alleles with Neanderthals
as non-African populations, whereas Sub-Saharan African groups are the only modern human
populations with no substantial Neanderthal admixture.[note 10] The Neanderthal-linked haplotype
B006 of the dystrophin gene has also been found among nomadic pastoralist groups in the Sahel and
Horn of Africa, who are associated with northern populations. Consequently, the presence of this
B006 haplotype on the northern and northeastern perimeter of Sub-Saharan Africa is attributed to
gene flow from a non-African point of origin.[note 11]
East 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.[106] 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 suggest that this Neanderthal
introgression occurred within the ancestral population shared by East Asians and Native
Americans.[107]
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.[108] 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
Eurasia
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 Schematic illustration of the
earlier than previously thought.[112] Beringia migration based on
matrilineal genetics: Arrival
of Central Asian
Americas populations to the Beringian
Mammoth steppe c. 25,000
Paleo-Indians originated from Central Asia, crossing the Beringia land years ago, followed by a
bridge between eastern Siberia and present-day Alaska.[113] Humans "swift peoplling of the
lived throughout the Americas by the end of the last glacial period, or Americas" c. 15,000 years
more specifically what is known as the late glacial maximum, no earlier ago.
than 23,000 years before present.[113][114][115][116] 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.[117]
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,[114][117] following herds
of now-extinct pleistocene megafauna along ice-free corridors that stretched between the Laurentide
and Cordilleran ice sheets.[118] 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.[119] 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.[120] The recent finding of indigenous Australasian genetic
markers in Amazonia supports the coastal route hypothesis.[121][122]
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.
Eurasia
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.[124]
The Afroasiatic Urheimat has been placed in

either Africa or Asia.
Prehistoric migration routes for Y-chromosome
Sub-Saharan Africa Haplogroup N lineage following the retreat of ice
sheets after the Last Glacial Maximum (22–18
The Nilotic peoples are thought to be derived kya).[123]
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.[125]
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.[126] 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.[127]
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.[128] 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).[129]
Indo-Pacific
The first seaborne human migrations were by the

Austronesian peoples originating from Taiwan
known as the "Austronesian expansion".[130]
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 Chronological map of the Austronesian expansion
Indonesia they colonized Micronesia by 2200 to
1000 BCE.[130][131]
A branch of the Austronesians reached Island Melanesia between 1600 and 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.[131][132][133]
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.[130][131] The Austronesian expansion was the last and the most far-reaching Neolithic human
migration event.[134]
Caribbean
The Caribbean was one of the last places in the Americas that were settled by humans. The oldest
remains are known from the Greater Antilles (Cuba and Hispaniola) dating between 4000 and 3500
BCE, and comparisons between tool-technologies suggest that these peoples moved across the
Yucatán Channel from Central America. All evidence suggests that later migrants from 2000 BCE and
onwards originated from South America, via the Orinoco region. The descendants of these migrants
include the ancestors of the Taíno and Kalinago (Island Carib) peoples.[135]
Arctic
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.[136][137]
The Inuit are the descendants of the Thule culture, which emerged from western Alaska around AD
1000 and gradually displaced the Dorset culture.[138][139]
See also
List of countries and islands by first human settlement
Middle Paleolithic
Quaternary extinction event
Timeline of human evolution
Timeline of maritime migration and exploration
Notes
1. Based on Schlebusch et al., "Southern African ancient genomes estimate modern human
divergence to 350,000 to 260,000 years ago",[1] Fig. 3 (https://d2ufo47lrtsv5s.cloudfront.net/conte
nt/sci/early/2017/09/27/science.aao6266/F3.large.jpg) (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.[20]
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."[27]
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."[38]
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."[43]
8. Early modern human presence outside of Africa has been proposed to date back to as early as
177,000 years ago.[36]
9. The authors of Liu (2010) seem to accept that the individual has African recent ascentry, but with
Asian archaic human admixture.[52] See also Dennell (2010).[53] Brief comments at [54] and [55]
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."[104]
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."[105]
12. Traits affected by the mutation are sweat glands, teeth, hair thickness and breast tissue.[109][110]
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.[111]
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Further reading
Demeter F, Shackelford LL, Bacon AM, Duringer P, Westaway K, Sayavongkhamdy T, Braga J,
Sichanthongtip P, Khamdalavong P, Ponche JL, et al. (2012). "Anatomically modern human in
Southeast Asia (Laos)" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3437904). Proceedings of
the National Academy of Sciences. 109 (36): 14375–14380. Bibcode:2012PNAS..10914375D (htt
ps://ui.adsabs.harvard.edu/abs/2012PNAS..10914375D). doi:10.1073/pnas.1208104109 (https://d
oi.org/10.1073%2Fpnas.1208104109). PMC 3437904 (https://www.ncbi.nlm.nih.gov/pmc/articles/
PMC3437904). PMID 22908291 (https://pubmed.ncbi.nlm.nih.gov/22908291).
Diamond, Jared, Guns, germs and steel. A short history of everybody for the last 13'000 years,
1997.
Reich, David (2018). Who We Are And How We Got Here – Ancient DNA and the New Science of
the Human Past. Pantheon Books. ISBN 978-1-101-87032-7.Diamond, Jared (20 April 2018). "A
Brand-New Version of Our Origin Story" (https://www.nytimes.com/2018/04/20/books/review/davi
d-reich-who-we-are-how-we-got-here.html). The New York Times. Retrieved 23 April 2018.
Veeramah, Krishna R.; Hammer, Michael F. (4 February 2014). "The impact of whole-genome
sequencing on the reconstruction of human population history". Nature Reviews Genetics. 15 (3):
149–162. doi:10.1038/nrg3625 (https://doi.org/10.1038%2Fnrg3625). PMID 24492235 (https://pu
bmed.ncbi.nlm.nih.gov/24492235). S2CID 19375407 (https://api.semanticscholar.org/CorpusID:1
9375407).
External links
Journey of Mankind – Genetic Map (http://www.bradshawfoundation.com/journey/) – Bradshaw
Foundation
Prehistoric Human Migration From Africa to World Video May 2015 (http://www.businessinsider.co
m/prehistoric-human-migration-from-africa-animated-map-2015-5)
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Early human migrations

Within Africa, Homo sapiens dispersed

Early humans (before Homo sapiens)

One million years after its dispersal, H. erectus was diverging

H. heidelbergensis, Neanderthals and Denisovans expanded

Dispersal throughout Africa

Homo sapiens are believed to have emerged in

Early northern Africa dispersal

The so-called "recent dispersal" of modern

A small group from a population in East

Dispersal throughout Eurasia

Up to 37,500 YBP Up to 35,000 YBP Up to 32,500 YBP Up to 30,000 YBP

East and North Asia

Last Glacial Maximum

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

Evidence published in 2014 from genome

The Afroasiatic Urheimat has been placed in

The first seaborne human migrations were by the

nt/sci/early/2017/09/27/science.aao6266/F3.large.jpg) (H. sapiens divergence times) and Stringer

98. Caramelli, D; Lalueza-Fox, C; Vernesi, C; Lari, M; Casoli, A; Mallegni, F; Chiarelli, B; Dupanloup,

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