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Early human migrations

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Further information:History of human migrationandPre-modern human migration

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

Early human migrationsare the earliestmigrations and expansionsofarchaic and modern humansacross continents. They are believed to have begun approximately 2 million years ago with theearly expansions out of AfricabyHomo erectus.This initial migration was followed by otherarchaic humansincludingH. heidelbergensis,which lived around 500,000 years ago and was the likely ancestor ofDenisovansandNeanderthalsas well as modern humans. Early hominids had likely crossedland bridgesthat have now sunk.

Within Africa,Homo sapiensdispersed around the time of itsspeciation,roughly 300,000 years ago.[note 1]Therecent African originparadigm suggests that the anatomically modern humans outside of Africa descend from a population ofHomo sapiensmigrating fromEast Africaroughly 70–50,000 years ago and spreadingalong the southern coastof Asia and to Oceania by about 50,000 years ago. Modern humans spreadacross Europeabout 40,000 years ago.

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

The migrating modern human populations are known to haveinterbredwith 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 theLast Glacial Maximum,North Eurasianpopulations migratedto the Americasabout 20,000 years ago. Arctic Canada and Greenland were reached by thePaleo-Eskimoexpansion around 4,000 years ago. Finally,Polynesiawas populated within the past 2,000 years in the last wave of theAustronesian expansion.

Early humans (Homo erectusbeforeHomo sapiens)

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See also:Early expansions of hominins out of Africa

Theearliest humansdeveloped out ofaustralopithecineancestors about 3 million years ago, most likely in the area of theKenyan Rift Valley,where theoldest known stone toolshave been found. Stone tools recently discovered at theShangchensite in China and dated to 2.12 million years ago are claimed to be the earliest known evidence of hominins outside Africa, surpassingDmanisiin Georgia by 300,000 years.[6]

Homo erectus

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Main articles:Homo erectusandOldowan

Between 2 and less than a million years ago,Homospread throughout East Africa and toSouthern Africa(Homo ergaster), but not yet to West Africa. Around 1.8 million years ago,Homo erectusmigrated out of Africavia theLevantine corridorandHorn of AfricatoEurasia.This migration has been proposed as being related to the operation of theSaharan pump,around 1.9 million years ago.[citation needed]Homo erectusdispersed throughout most of theOld World,reaching as far asSoutheast Asia.Its distribution is traced by theOldowanlithic industry, by 1.3 million years ago extending as far north as the40th parallel(Xiaochangliang).

Key sites for this early migration out of Africa areRiwatin Pakistan (~2 Ma?[7]),Ubeidiyain the Levant (1.5 Ma) andDmanisiin the Caucasus (1.81 ± 0.03 Ma,p=0.05[8]).

Chinashows evidence ofHomo erectusfrom 2.12 mya in Gongwangling, in Lantian county.[9]TwoHomo erectusincisors have been found near Yuanmou, southern China, and are dated to 1.7 mya, and a cranium from Lantian has been dated to 1.63 mya. Artefacts from Majuangou III and Shangshazui in theNihewan basin,northern China, have been dated to 1.6–1.7 mya.[9][10]The archaeological site ofXihoudu(Tây hầu độ) inShanxiprovince is the earliest recordeduse of firebyHomo erectus,which is dated 1.27 million years ago.[11]

Southeast Asia(Java) was reached about 1.7 million years ago (Meganthropus). WesternEuropewas first populated around 1.2 million years ago (Atapuerca).[12]

Robert G. Bednarikhas suggested thatHomo erectusmay have built rafts and sailed oceans, a theory that has raised some controversy.[13]

AfterH. erectus

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

One million years after its dispersal,H. erectuswas diverging into new species.H. erectusis achronospeciesand was never extinct, so its "late survival" is a matter of taxonomic convention. Late forms ofH. erectusare 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 asH. antecessorin Europe around 800,000 years ago andH. heidelbergensisin Africa around 600,000 years ago.H. heidelbergensisin its turn spread across East Africa (H. rhodesiensis) and to Eurasia, where it gave rise toNeanderthalsandDenisovans.

H. heidelbergensis,Neanderthals and Denisovans expanded north beyond the50th parallel(Eartham Pit, Boxgrove500kya,Swanscombe Heritage Park400kya,Denisova Cave50 kya). It has been suggested that late Neanderthals may even have reached the boundary of theArctic,by c. 32,000 years ago, when they were being displaced from their earlier habitats byH. sapiens,based on 2011 excavations at the site of Byzovaya in theUrals(Komi Republic,65°01′N57°25′E/ 65.02°N 57.42°E/65.02; 57.42).[15]

Other archaic human species are assumed to have spread throughout Africa by this time, although the fossil record is sparse. Their presence is assumed based on traces ofadmixturewith modern humans found in the genome of African populations.[5][16][17][18]Homo naledi,discovered in South Africa in 2013 and tentatively dated to about 300,000 years ago, may represent fossil evidence of such an archaic human species.[19]

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

Homo sapiens

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Dispersal throughout Africa

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Further information:Macro-haplogroup L (mtDNA)andArchaeogenetics of sub-Saharan Africa

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

In September 2019, scientists reported the computerized determination, based on 260CT scans,of a virtualskull shapeof 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 inEastandSouth Africa.[28][29]

In July 2019, anthropologists reported the discovery of 210,000 year old remains of aH. sapiensand 170,000 year old remains of aH. neanderthalensisinApidima Cavein southernGreece,more than 150,000 years older than previousH. sapiensfinds in Europe.[30][31][32][33]

Early modern humans expanded to Western Eurasia and Central, Western and Southern Africa from the time of their emergence. Whileearly expansionsto Eurasia appear not to have persisted,[34][20]expansions toSouthernandCentral Africaresulted 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 lateAcheulean(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 modernKhoi-Sanexpanded 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 theMIS 5"megadrought",130,000 years ago, there were two ancestral population clusters in Africa, bearers ofmt-DNA haplogroup L0in southern Africa, ancestral to the Khoi-San, and bearers ofhaplogroup L1-6in 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 theCentral African foragerpopulations (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 inWest Africais difficult to interpret due to a sparsity of fossil evidence.Homo sapiensseems to have reached the westernSahelian zoneby 130,000 years ago, while tropical West African sites associated withH. sapiensare known only from after 130,000 years ago. Unlike elsewhere in Africa, archaicMiddle Stone Agesites appear to persist until very late, down to the Holocene boundary (12,000 years ago), pointing to the possibility of late survival ofarchaic humans,and latehybridizationwithH. sapiensin West Africa.[35]

Early northern Africa dispersal

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

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

A fragment of a jawbone with eight teeth found atMisliya Cavehas 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 theLevalloistype which could put the date of the first migration even earlier if the tools can be associated with the modern human jawbone finds.[42][43]

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

Fuller projectionmap showing early human migrations according tomitochondrialpopulation genetics(numbers aremillenniabefore present).

There is some evidence that modern humans left Africa at least 125,000 years ago using two different routes: through theNile Valley,theSinai Peninsulaand theLevant(Qafzeh Cave:120,000–100,000 years ago); and a second route through the present-dayBab-el-MandebStrait on the Red Sea (at that time, with a much lower sea level and narrower extension), crossing to theArabian Peninsula[45][46]and settling in places like the present-day United Arab Emirates (125,000 years ago)[47]and Oman (106,000 years ago),[48]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 atJebel Faya,those from Jwalapuram and some from Africa suggest that their creators were all modern humans.[49]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,ChongzuoCity: 100,000 years ago;[note 9]and theLiujiang hominid(Liujiang County): controversially dated at 139,000–111,000 years ago[54]). Dating results of theLunadong(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.[55][56]

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

Coastal migration

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

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

A small group from a population in East Africa, bearingmitochondrial haplogroup L3and numbering possibly fewer than 1,000 individuals,[61][62]crossed theRed Seastrait atBab-el-Mandeb,to what is nowYemen,after around 75,000 years ago.[63]A recent review has also shown support for the northern route through theSinai Peninsulaand theLevant.[20]Their descendants spread along thecoastal routearoundArabiaandPersiatoSouth Asiabefore 55,000 years ago. Other research supports a migration out of Africa between about 65,000 and 50,000 years ago.[58][64][60]The coastal migration between roughly 70,000 and 50,000 years ago is associated with mitochondrial haplogroupsMandN,both derivative of L3.

Along the wayH. sapiensinterbred with Neanderthals and Denisovans,[65]with Denisovan DNA making 0.2% of mainland Asian and Native American DNA.[66]

Nearby Oceania

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Main article:Peopling of Oceania

Migrations continued along the Asian coast to Southeast Asia and Oceania, colonisingAustraliaby around 65,000–50,000 years ago.[67][68][69]By reaching Australia,H. sapiensfor the first time expanded its habitat beyond that ofH. erectus.Denisovan ancestry is shared byMelanesians,Aboriginal Australians,and smaller scattered groups of people in Southeast Asia, such as theMamanwa,aNegritopeople in thePhilippines,suggesting the interbreeding took place in Eastern Asia where the Denisovans lived.[70][71][72]Denisovans may have crossed theWallace Line,withWallaceaserving as their lastrefugium.[73][74]Homo erectushad crossed the Lombok gap reaching as far as Flores, but never made it to Australia.[75]

The map shows the probable extent of land and water at the time of thelast glacial maximum,20,000 yrs ago and when the sea level was probably more than 110m lower than today.

During this time sea level was much lower and most ofMaritime Southeast Asiaformed one land mass known asSunda.Migration continued Southeast on thecoastal routeto thestraitsbetween Sunda andSahul,the continental land mass of present-day Australia andNew Guinea.The gaps on theWeber Lineare up to 90 km wide,[76]so the migration to Australia and New Guinea would have required seafaring skills. Migration also continued along the coast eventually turning northeast toChinaand finally reachingJapanbefore turning inland. This is evidenced by the pattern ofmitochondrial haplogroupsdescended fromhaplogroup M,and inY-chromosomehaplogroup C.

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

Dispersal throughout Eurasia

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Further information:Upper Paleolithic,Mammoth steppe,Interbreeding between archaic and modern humans,andMousterian
Successive dispersals of
Homo erectusgreatest extent (yellow),
Homo neanderthalensisgreatest extent (ochre) and
Homo sapiens(red).

The population brought toSouth Asiabycoastal migrationappears 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 theUpper Paleolithic,gave rise to the major population groups of theOld Worldand theAmericas.

Towards the West, Upper Paleolithic populations associated with mitochondrial haplogroupRand its derivatives, spread throughout Asia and Europe, with a back-migration ofM1to North Africa and the Horn of Africa several millennia ago.[dubious–discuss]

Presencein Europeis certain after 40,000 years ago, possibly as early as 43,000 years ago,[84]rapidly replacing the Neanderthal population. Contemporary Europeans haveNeanderthal ancestry,but it seems likely that substantial interbreeding with Neanderthals ceased before 47,000 years ago, i.e. took place before modern humans entered Europe.[85]

There is evidence frommitochondrial DNAthat modern humans have passed through at least onegenetic bottleneck,in which genome diversity was drastically reduced.Henry Harpendinghas 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.[86]Climatological and geological evidence suggests evidence for the bottleneck. The explosion ofToba,the largest volcanic eruption of theQuaternary,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 andfounder effectsmay have been maximised. The greater diversity amongst African genomes may reflect the extent of African refugia during the Toba incident.[87]However, a recent review highlights that the single-source hypothesis of non-African populations is less consistent with ancient DNA analysis than multiple sources with genetic mixing across Eurasia.[20]

Europe

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Main articles:Early European modern humans,Paleolithic Europe,Aurignacian,Gravettian,andArt of the Upper Paleolithic
Further information:Hominid dispersals in Europe
30,000-year-oldcave lionandbisonpainting found in theChauvet Cave,France.

The recent expansion ofanatomically modern humansreached Europe around 40,000 years ago from Central Asia and the Middle East, as a result of cultural adaption to big game hunting ofsub-glacial steppefauna.[88]Neanderthalswere present both in the Middle East and in Europe, and the arriving populations of anatomically modern humans (also known as "Cro-Magnon"orEuropean early modern humans)interbred with Neanderthal populationsto 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, reachedsites in Arctic Russiaby 40,000 years ago.[89]

Cro-Magnon are considered the first anatomically modern humans in Europe. They enteredEurasiaby theZagros Mountains(near present-dayIranand easternTurkey) around 50,000 years ago, with one group rapidly settling coastal areas around theIndian Oceanand another migrating north to the steppes ofCentral Asia.[90]Modern human remains dating to 45,000-47,000 have been found inGermany,[91]while finds of 43,000–45,000 years ago have been discovered in Italy[92]and Britain,[93]as well as in the European Russian Arctic from 40,000 years ago.[89][94]

Humans colonised the environment west of the Urals, hunting reindeer especially,[95]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.[95][96]

However, from recent research it is believed that the ecological crisis resulting from the eruption in c. 38,000 BCE of the super-volcano in thePhlegrean Fieldsnear Naples, which left much of eastern Europe covered in ash, wiped out both the last Neanderthal and the first Homo Sapiens populations of the early Upper Paleolithic.[97][98]Modern Europeans of today bear no trace of the genomes of the first Homo Sapiens Europeans, but only of those from after the ecological crisis of 38,000 BCE.[99]Modern humans then repopulated Europe from the east after the eruption and the ice age that took place from 38,000 to 36,000 BCE.[100]

Amitochondrial DNAsequence of two Cro-Magnons from thePaglicci Cavein Italy, dated to 23,000 and 24,000 years old (Paglicci 52 and 12), identified themtDNAasHaplogroup N,typical of the latter group.[101]

Migration of modern humans into Europe, based on simulation by Currat & Excoffier (2004)[102]
(YBP =Years before present)
Up to 37,500 YBP
Up to 35,000 YBP
Up to 32,500 YBP
Up to 30,000 YBP
Further information:Neanderthal extinctionandInterbreeding between archaic and modern humans

The expansion of modern human population is thought to have begun 45,000 years ago, and it may have taken 15,000–20,000 years for Europe to be colonized.[103][104]

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 modernHomo 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 theCro-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 theIberian Peninsula.Neanderthals disappeared about 40,000 years ago.[105]

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 yearsBP.In conjunction with archaeological and fossil evidence, interbreeding is thought to have occurred somewhere in Western Eurasia, possibly the Middle East.[85]Studies show a higher Neanderthal admixture in East Asians than in Europeans.[106][107]North African groups share a similar excess of derived alleles with Neanderthals as non-African populations, whereas Sub-Saharan African groups are the only modern human populations with no substantial Neanderthal admixture.[note 10]The Neanderthal-linked haplotype B006 of the dystrophin gene has also been found among nomadic pastoralist groups in the Sahel and Horn of Africa, who are associated with northern populations. Consequently, the presence of this B006 haplotype on the northern and northeastern perimeter of Sub-Saharan Africa is attributed to gene flow from a non-African point of origin.[note 11]

East, Southeast and North Asia

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Ancient North Eurasianpopulations from Siberia were an important genetic contributor toAncient Native AmericansandEastern European Hunter-Gatherers.Neolithic Iranian farmers andJōmon people(ancestors of theAinu people) also received geneflow from ANE-related populations.[110]

"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.[111]A 2013 study foundNeanderthal introgressionof 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.[112]

A 2016 study presented an analysis of the population genetics of theAinupeople 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.[113]A 2013 study associated severalphenotypicaltraits associated with Mongoloids with a single mutation of theEDARgene, dated to c. 35,000 years ago.[note 12][note 13]

Further information:Ancient North Eurasians

Mitochondrial haplogroupsA,BandGoriginated about 50,000 years ago, and bearers subsequently colonizedSiberia,KoreaandJapan,by about 35,000 years ago. Parts of these populations migrated to North America during theLast Glacial Maximum.

A review paper by Melinda A. Yang (in 2022) summarized and concluded that a distinctive "Basal-East Asian population" referred to as 'East- and Southeast Asian lineage' (ESEA); which is ancestral to modern East Asians,Southeast Asians,Polynesians,andSiberians,originated inMainland Southeast Asiaat ~50,000BC, and expanded through multiple migration waves southwards and northwards respectively. This ESEA lineage gave rise to various sublineages, and is also ancestral to theHoabinhian hunter-gatherersof Southeast Asia and the ~40,000 year oldTianyuanlineage found inNorthern China,but already differentiated and distinct fromEuropean-relatedandAustralasian-relatedlineages, found in other regions of prehistoric Eurasia. The ESEA lineage trifurcated from an earlier East-Eurasian or "eastern non-African" (ENA) meta-population, which also contributed to the formation of Ancient Ancestral South Indians (AASI) as well as to Australasians.[117]

Last Glacial Maximum

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Eurasia

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Further information:SolutreanandMagdalenian
Schematic illustration of theBeringiamigration based onmatrilineal genetics:Arrival of Central Asian populations to the BeringianMammoth steppec. 25,000 years ago, followed by a "swift peoplling of the Americas"[citation needed]c. 15,000 years ago.

Around 20,000 years ago, approximately 5,000 years after the Neanderthal extinction, theLast Glacial Maximumforced northern hemisphere inhabitants to migrate to severalshelters(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 theNeolithicexpansion from the Middle East, and still later theChalcolithicpopulation movements associated withIndo-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.[118]

Americas

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Main articles:Settlement of the AmericasandGenetic history of Indigenous peoples of the Americas

Paleo-Indiansoriginated fromCentral Asia,crossing theBeringia land bridgebetween eastern Siberia and present-day Alaska.[119]Humans lived throughout the Americas by the end of thelast glacial period,or more specifically what is known as thelate glacial maximum.[119][120][121][122]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.[123]

Conventional estimates have it that humans reached North America at some point between 15,000 and 20,000 years ago.[124][125][126][127]The traditional theory is that these early migrants moved when sea levels were significantly lowered due to theQuaternary glaciation,[120][123]following herds of now-extinctpleistocenemegafaunaalongice-free corridorsthat stretched between theLaurentideandCordilleranice sheets.[128]Another route proposed is that, either on foot or usingprimitive boats,they migrated down the Pacific coast toSouth Americaas far asChile.[129]Any archaeological evidence of coastal occupation during the last Ice Age would now have been covered by thesea level rise,up to a hundred metres since then.[130]The recent finding of indigenousAustralasiangenetic markers in Amazonia supports that a coastal route and subsequent isolation did occur with some migrants.[131]

Holocene migrations

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Further information:Pre-modern human migration,Mesolithic,andLinguistic homeland
Prehistoric migration routes for Y-chromosome Haplogroup N lineage following the retreat of ice sheets after the Last Glacial Maximum (22–18 kya).[132]

TheHoloceneis taken to begin 12,000 years ago, after the end of theLast Glacial Maximum.During theHolocene climatic optimum,beginning about 9,000 years ago, human populations which had been geographically confined torefugiabegan to migrate. By this time, most parts of the globe had been settled byH. sapiens;however, large areas that had been covered byglacierswere now re-populated.

This period sees the transition from theMesolithicto theNeolithicstage throughout thetemperate zone.The Neolithic subsequently gives way to theBronze AgeinOld Worldcultures and the gradual emergence of thehistorical recordin theNear EastandChinabeginning 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 majorlanguage familiessuch as theNiger-Congo,Nilo-Saharan,Afro-Asiatic,Uralic,Sino-TibetanorIndo-Europeanphyla. The speculativeNostratic theorypostulates the derivation of the major language families of Eurasia (excluding Sino-Tibetan) from a single proto-language spoken at the beginning of the Holocene period.

Eurasia

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

Evidence published in 2014 from genome analysis of ancient human remains suggests that the modern native populations of Europe largely descend from three distinct lineages: "Western Hunter-Gatherers",derivative of the Cro-Magnon population of Europe,Early European Farmersintroduced to Europe from the Near East during theNeolithic RevolutionandAncient North Eurasianswho expanded to Europe in the context of theIndo-European expansion.[133]The Ancient North Eurasian component was introduced to Western Europe by people related to theYamnaya culture.[134]Additional ANE ancestry is found in European populations through Paleolithic interactions withEastern Hunter-Gatherers.[135]

Sub-Saharan Africa

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Further information:Nilo-SaharanandNiger-Congo

West-Eurasian back-migrations started in the earlyHoloceneor already earlier in thePaleolithicperiod (30-15kya), followed by pre-Neolithic andNeolithicmigration events from theMiddle East,mostly affecting Northern Africa, the Horn of Africa, and wider regions of the Sahel zone and East Africa.[136]

Pre-Neolithic and Neolithic migration events in Africa.[136]

TheNilotic peoplesare thought to be derived from an earlier undifferentiatedEastern Sudanicunity by the 3rd millennium BCE. The development of the Proto-Nilotes as a group may have been connected with their domestication oflivestock.The Eastern Sudanic unity must have been considerably earlier still, perhaps around the 5th millennium BCE (while the proposedNilo-Saharanunity would date to theUpper Paleolithicabout 15kya). The original locus of the early Nilotic speakers was presumably east of the Nile in what is nowSouth Sudan.The Proto-Nilotes of the 3rd millennium BCE werepastoralists,while their neighbors, the Proto-Central Sudanicpeoples, were mostly agriculturalists.[137]

TheNiger-Congophylum 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 inc. 3900 BCE.[138]TheBantu expansionhas spread theBantu languagesto Central, Eastern and Southern Africa, partly replacing theindigenouspopulations of these regions, including theAfrican Pygmies,Hadza peopleandSan people.Beginning about 3,000 years ago, it reached South Africa about 1,700 years ago.[139]

Some evidence (including a 2016 study by Busby et al.) suggests admixture from ancient and recent migrations fromEurasiainto parts of Sub-Saharan Africa.[140]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 theSahelzone, and found to a lesser degree in certain parts of Western Africa, andSouthern Africa(excluding recent immigrants).[141]

Indo-Pacific

[edit]
Further information:Austronesian peoples § Austronesian expansion
Chronological map of theAustronesian expansion.

The first seaborne human migrations were by theAustronesian peoples[dubious–discuss]originating fromTaiwanknown as the "Austronesian expansion".[142]Using advanced sailing technologies likecatamarans,outrigger boats,andcrab claw sails,they built the first sea-going ships and rapidly colonizedIsland Southeast Asiaat around 3000 to 1500 BCE. From thePhilippinesandEastern Indonesiathey colonizedMicronesiaby 2200 to 1000 BCE.[142][143]

A branch of the Austronesians reachedIsland Melanesiabetween 1600 and 1000 BCE, establishing theLapita culture(named after the archaeological site in Lapita,New Caledonia,where their characteristic pottery was first discovered). They are the direct ancestors of the modernPolynesians.They ventured intoRemote OceaniareachingVanuatu,New Caledonia,andFijiby 1200 BCE, andSamoaandTongaby 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 theCook Islands,theSociety Islands,and theMarquesas.From there, they further colonizedHawaiiby 900 CE,Rapa Nuiby 1000 CE, andNew Zealandby 1200 CE.[143][144][145]

In theIndian Ocean,Austronesians fromBorneoalso colonizedMadagascarand theComoros Islandsby around 500 CE. Austronesians remain the dominant ethnolinguistic group of the islands of the Indo-Pacific, and were the first to establish amaritime trade networkreaching as far west asEast Africaand theArabian Peninsula.They assimilated earlierPleistoceneto earlyHolocenehuman overland migrations throughSundalandlike thePapuansand theNegritosin Island Southeast Asia.[142][143]The Austronesian expansion was the last and the most far-reachingNeolithichuman migration event.[146]

Caribbean

[edit]

TheCaribbeanwas 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.[147]The descendants of these migrants include the ancestors of theTaínoandKalinago(Island Carib) peoples.[148]

Arctic

[edit]
Main article:Circumpolar peoples
Map showing the decline of the Paleo-EskimoDorset cultureand expansion of theThule people(900 to 1500 CE).

The earliest inhabitants of North America's central and eastern Arctic are referred to as theArctic small tool tradition(AST) and existed c. 2500 BCE. AST consisted of severalPaleo-Eskimocultures, including theIndependence culturesandPre-Dorsetculture.[149][150]

TheInuitare the descendants of theThule culture,which emerged from western Alaska around CE 1000 andgradually displacedthe Dorset culture.[151][152]

See also

[edit]

Notes

[edit]
  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. sapiensdivergence times) and Stringer (2012),[2](archaic admixture).
  2. ^Archaic admixture from various sources is known from Europe and Asia (Neanderthals), Southeast Asia and Melanesia (Denisovans) as well as from Western and Southern Africa. The proportion of admixture varies by region, but in all cases has been reported below 10%: In Eurasian mostly estimated at 1–4% (with a high estimate of 3.4–7.3% by Lohse (2014)[3]) in Melanesians estimated at 4–6% (Reich et al. (2010)).[4]Admixture of an unknown archaic hominin in Sub-Saharan African hunter-gatherer populations was estimated at 2% (Hammer et al. (2011)).[5]
  3. ^Homo erectus soloensis,found inJava,is considered the latest known specimen ofH. erectus.Formerly dated to as late as 50,000 to 40,000 years ago, a 2011 study pushed back the date of the extinction ofH. e. soloensisto 143,000 years ago at the latest, more likely before 550,000 years ago.[14]
  4. ^"Here we report the ages, determined by thermoluminescence dating, of fire-heated flint artefacts obtained from new excavations at the Middle Stone Age site of Jebel Irhoud, Morocco, which are directly associated with newly discovered remains of H. sapiens. A weighted average age places these Middle Stone Age artefacts and fossils at 315±34 thousand years ago. Support is obtained through the recalculated uranium series with electron spin resonance date of 286±32 thousand years ago for a tooth from the Irhoud 3 hominin mandible."[21]
  5. ^Estimated split times given in the source cited (in kya): Human-Neanderthal: 530–690, Deep Human [H. sapiens]: 250–360, NKSP-SKSP: 150–190, Out of Africa (OOA): 70–120.[1]
  6. ^"By ~130 ka two distinct groups of anatomically modern humans co-existed in Africa: broadly, the ancestors of many modern-day Khoe and San populations in the south and a second central/eastern African group that includes the ancestors of most extant worldwide populations. Early modern human dispersals correlate with climate changes, particularly the tropical African" megadroughts "of MIS 5 (marine isotope stage 5, 135–75 ka) which paradoxically may have facilitated expansions in central and eastern Africa, ultimately triggering the dispersal out of Africa of people carrying haplogroup L3 ~60 ka. Two south to east migrations are discernible within haplogroup L0. One, between 120 and 75 ka, represents the first unambiguous long-range modern human dispersal detected by mtDNA and might have allowed the dispersal of several markers of modernity. A second one, within the last 20 ka signalled by L0d, may have been responsible for the spread of southern click-consonant languages to eastern Africa, contrary to the view that these eastern examples constitute relicts of an ancient, much wider distribution."[36]
  7. ^"We studied the branching history of Pygmy hunter–gatherers and agricultural populations from Africa and estimated separation times and gene flow between these populations. The model identified included the early divergence of the ancestors of Pygmy hunter–gatherers and farming populations ~60,000 years ago, followed by a split of the Pygmies' ancestors into the Western and Eastern Pygmy groups – 20,000 years ago."[41]
  8. ^Early modern human presence outside of Africa has been proposed to date back to as early as 177,000 years ago.[34]
  9. ^The authors of Liu (2010) seem to accept that the individual has African recent ascentry, but with Asian archaic human admixture.[50]See also Dennell (2010).[51]Brief comments at[52]and[53]
  10. ^"We found that North African populations have a significant excess of derived alleles shared with Neandertals, when compared to sub-Saharan Africans. This excess is similar to that found in non-African humans, a fact that can be interpreted as a sign of Neandertal admixture. Furthermore, the Neandertal's genetic signal is higher in populations with a local, pre-Neolithic North African ancestry. Therefore, the detected ancient admixture is not due to recent Near Eastern or European migrations. Sub-Saharan populations are the only ones not affected by the admixture event with Neandertals."[108]
  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."[109]
  12. ^Traits affected by the mutation are sweat glands, teeth, hair shaft thickness and breast tissue.[114][115]
  13. ^East Asian genetics shows a number of concentrated alleles suggestive of selection pressures. This concerns the genesEDAR,ADH1B,ABCC1,andALDH2in particular. The East Asian types of ADH1B are associated withrice domesticationand would thus have arisen after the c. 11,000 years ago.[116]

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Further reading

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