
The recent African origin of modern humans or the "Out of Africa" theory (OOA) is the most widely accepted
paleo-anthropological model of the geographic origin and
early migration of
anatomically modern humans (''
Homo sapiens
Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
''). It follows the
early expansions of hominins out of Africa, accomplished by ''
Homo erectus
''Homo erectus'' ( ) is an extinction, extinct species of Homo, archaic human from the Pleistocene, spanning nearly 2 million years. It is the first human species to evolve a humanlike body plan and human gait, gait, to early expansions of h ...
'' and then ''
Homo neanderthalensis
Neanderthals ( ; ''Homo neanderthalensis'' or sometimes ''H. sapiens neanderthalensis'') are an extinct group of archaic humans who inhabited Europe and Western and Central Asia during the Middle to Late Pleistocene. Neanderthal extinctio ...
''.
The model proposes a "
single origin" of ''
Homo sapiens
Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
'' in the
taxonomic sense, precluding
parallel evolution in other regions of traits considered
anatomically modern, but not precluding multiple
admixture between ''H. sapiens'' and
archaic humans
''Homo'' () is a genus of great ape (family Hominidae) that emerged from the genus ''Australopithecus'' and encompasses only a single extant species, ''Homo sapiens'' (modern humans), along with a number of extinct species (collectively calle ...
in Europe and Asia. ''H. sapiens'' most likely developed in the
Horn of Africa
The Horn of Africa (HoA), also known as the Somali Peninsula, is a large peninsula and geopolitical region in East Africa.Robert Stock, ''Africa South of the Sahara, Second Edition: A Geographical Interpretation'', (The Guilford Press; 2004), ...
between 300,000 and 200,000 years ago,
although an alternative hypothesis argues that diverse morphological features of ''H. sapiens'' appeared locally in different parts of Africa and converged due to
gene flow
In population genetics, gene flow (also known as migration and allele flow) is the transfer of genetic variation, genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalent ...
between different populations within the same period. The "recent African origin" model proposes that all modern non-African populations are substantially descended from populations of ''H. sapiens'' that left Africa after that time.
There were at least several "out-of-Africa" dispersals of modern humans, possibly beginning as early as 270,000 years ago, including 215,000 years ago to at least Greece,
and certainly via northern Africa and the Arabian Peninsula about 130,000 to 115,000 years ago. There is evidence that modern humans had reached China around 80,000 years ago.
[.]
See als
''Modern humans in China ~80,000 years ago (?)''
Dieneks' Anthropology Blog. Practically all of these early waves seem to have gone extinct or retreated back, and present-day humans outside Africa descend mainly from a single expansion about 70,000–50,000 years ago,
via the so-called "
Southern Route". These humans spread rapidly along the coast of Asia and reached
Australia
Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
by around 65,000–50,000 years ago,
(though some researchers question the earlier Australian dates and place the arrival of humans there at 50,000 years ago at earliest,
while others have suggested that these first settlers of Australia may represent an older wave before the more significant out of Africa migration and thus not necessarily be ancestral to the region's later inhabitants
) while Europe was populated by an early offshoot which settled the Near East and Europe less than 55,000 years ago.
[.][.]
See also
In the 2010s, studies in
population genetics
Population genetics is a subfield of genetics that deals with genetic differences within and among populations, and is a part of evolutionary biology. Studies in this branch of biology examine such phenomena as Adaptation (biology), adaptation, s ...
uncovered evidence of
interbreeding that occurred between ''H. sapiens'' and
archaic humans
''Homo'' () is a genus of great ape (family Hominidae) that emerged from the genus ''Australopithecus'' and encompasses only a single extant species, ''Homo sapiens'' (modern humans), along with a number of extinct species (collectively calle ...
in Eurasia, Oceania and Africa,
indicating that modern population groups, while mostly derived from early ''H. sapiens'', are to a lesser extent also descended from regional variants of archaic humans.
Proposed waves

"Recent African origin", or ''Out of Africa II'', refers to the migration of
anatomically modern humans (''
Homo sapiens
Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
'') out of Africa after their emergence at c. 300,000 to 200,000 years ago, in contrast to "
Out of Africa I", which refers to the migration of archaic humans from Africa to Eurasia from before 1.8 and up to 0.5 million years ago.
Omo-Kibish I (Omo I) from southern Ethiopia is the oldest anatomically modern Homo sapiens skeleton currently known
(around 233,000 years old).
There are even older ''Homo sapiens'' fossils from
Jebel Irhoud
Jebel Irhoud or Adrar n Ighoud (; , Moroccan Arabic: ), is an archaeological site located just north of the town of Ighoud, Tlet Ighoud in Youssoufia Province, approximately south-east of the city of Safi, Morocco, Safi in Morocco. It is noted f ...
in Morocco which exhibit a mixture of modern and archaic features at around 315,000 years old.
Since the beginning of the 21st century, the picture of "recent single-origin" migrations has become significantly more complex, due to the discovery of modern-archaic admixture and the increasing evidence that the "recent out-of-Africa" migration took place in waves over a long time. As of 2010, there were two main accepted dispersal routes for the out-of-Africa migration of early anatomically modern humans, the "Northern Route" (via Nile Valley and Sinai) and the "Southern Route" via the
Bab-el-Mandeb strait.
* Posth et al. (2017) suggest that early ''Homo sapiens'', or "another species in Africa closely related to us", might have first migrated out of Africa around 270,000 years ago based on the closer affinity within Neanderthals'
mitochondrial genomes to ''Homo sapiens'' than
Denisovans
The Denisovans or Denisova hominins ( ) are an extinct species or subspecies of archaic human that ranged across Asia during the Lower Paleolithic, Lower and Middle Paleolithic, and lived, based on current evidence, from 285 thousand to 25 thou ...
.
[; see also .]
* Fossil evidence also points to an early ''Homo sapiens'' migration with the oldest known fossil coming from
Apidima Cave in Greece and dated at 210,000 years ago. Finds at
Misliya cave, which include a partial jawbone with eight teeth, have 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.
* An eastward dispersal from Northeast Africa to Arabia 150,000–130,000 years ago is based on the stone tools finds at
Jebel Faya dated to 127,000 years ago (discovered in 2011), although fossil evidence in the area is significantly later at 85,000 years ago.
Possibly related to this wave are the finds from
Zhirendong cave, Southern China, dated to more than 100,000 years ago. Other evidence of modern human presence in China has been dated to 80,000 years ago.
* The most significant out of Africa dispersal took place around 50,000–70,000 years ago via the so-called
Southern Route, either before or after the
Toba event, which happened between 69,000 and 77,000 years ago. This dispersal followed the southern coastline of Asia and reached Australia around 65,000–50,000 years ago or according to some research, by 50,000 years ago at earliest.
Western Asia was "re-occupied" by a different derivation from this wave around 50,000 years ago and Europe was populated from Western Asia beginning around 43,000 years ago.
* describes an additional wave of migration after the southern coastal route, a northern migration into Europe about 45,000 years ago. This possibility is ruled out by and , who argue for a single coastal dispersal, with an early offshoot into Europe.
Northern Route dispersal
Beginning 135,000 years ago, tropical Africa experienced
megadroughts which drove humans from the land and towards the sea shores, and forced them to cross over to other continents.
Fossils of early ''Homo sapiens'' were found in
Qafzeh and
Es-Skhul Caves in Israel and have been dated to 80,000 to 120,000 years ago. These humans seem to have either become extinct or retreated back to Africa 70,000 to 80,000 years ago, possibly replaced by southbound Neanderthals escaping the colder regions of ice-age Europe. Hua Liu ''et al.'' analyzed autosomal microsatellite markers dating to about 56,000 years ago. They interpret the
paleontological fossil as an isolated early offshoot that retracted back to Africa.
The discovery of stone tools in the
United Arab Emirates
The United Arab Emirates (UAE), or simply the Emirates, is a country in West Asia, in the Middle East, at the eastern end of the Arabian Peninsula. It is a Federal monarchy, federal elective monarchy made up of Emirates of the United Arab E ...
in 2011 at the
Faya-1 site in
Mleiha,
Sharjah
Sharjah (; ', Gulf Arabic: ''aš-Šārja'') is the List of cities in the United Arab Emirates, third-most populous city in the United Arab Emirates, after Dubai and Abu Dhabi. It is the capital of the Emirate of Sharjah and forms part of the D ...
, indicated the presence of modern humans at least 125,000 years ago, leading to a resurgence of the "long-neglected" North African route. This new understanding of the role of the Arabian dispersal began to change following results from archaeological and genetic studies stressing the importance of southern Arabia as a corridor for human expansions out of Africa.
In
Oman
Oman, officially the Sultanate of Oman, is a country located on the southeastern coast of the Arabian Peninsula in West Asia and the Middle East. It shares land borders with Saudi Arabia, the United Arab Emirates, and Yemen. Oman’s coastline ...
, a site was discovered by Bien Joven in 2011 containing more than 100 surface scatters of stone tools belonging to the late Nubian Complex, known previously only from
archaeological excavations in the
Sudan
Sudan, officially the Republic of the Sudan, is a country in Northeast Africa. It borders the Central African Republic to the southwest, Chad to the west, Libya to the northwest, Egypt to the north, the Red Sea to the east, Eritrea and Ethiopi ...
. Two optically stimulated luminescence age estimates placed the Arabian Nubian Complex at approximately 106,000 years old. This provides evidence for a distinct
Stone Age
The Stone Age was a broad prehistory, prehistoric period during which Rock (geology), stone was widely used to make stone tools with an edge, a point, or a percussion surface. The period lasted for roughly 3.4 million years and ended b ...
technocomplex in southern Arabia, around the earlier part of the
Marine Isotope Stage 5.
According to Kuhlwilm and his co-authors,
Neanderthal
Neanderthals ( ; ''Homo neanderthalensis'' or sometimes ''H. sapiens neanderthalensis'') are an extinction, extinct group of archaic humans who inhabited Europe and Western and Central Asia during the Middle Pleistocene, Middle to Late Plei ...
s
contributed genetically to modern humans then living outside of Africa around 100,000 years ago: humans which had already split off from other modern humans around 200,000 years ago, and this early wave of modern humans outside Africa also contributed genetically to the Altai Neanderthals.
[.]
See als
''Ancestors of Eastern Neandertals admixed with modern humans 100 thousand years ago''
Dienekes'Anthropology Blog. They found that "the ancestors of Neanderthals from the
Altai Mountains
The Altai Mountains (), also spelled Altay Mountains, are a mountain range in Central Asia, Central and East Asia, where Russia, China, Mongolia, and Kazakhstan converge, and where the rivers Irtysh and Ob River, Ob have their headwaters. The ...
and early modern humans met and interbred, possibly in the Near East, many thousands of years earlier than previously thought". According to co-author Ilan Gronau, "This actually complements archaeological evidence of the presence of early modern humans out of Africa around and before 100,000 years ago by providing the first genetic evidence of such populations." Similar
genetic admixture events have been
noted in other regions as well.
Southern Route dispersal
Coastal route

By some 50–70,000 years ago, a subset of the bearers of mitochondrial haplogroup
L3 migrated from
East Africa
East Africa, also known as Eastern Africa or the East of Africa, is a region at the eastern edge of the Africa, African continent, distinguished by its unique geographical, historical, and cultural landscape. Defined in varying scopes, the regi ...
into the
Near East
The Near East () is a transcontinental region around the Eastern Mediterranean encompassing the historical Fertile Crescent, the Levant, Anatolia, Egypt, Mesopotamia, and coastal areas of the Arabian Peninsula. The term was invented in the 20th ...
. It has been estimated that from a population of 2,000 to 5,000 individuals in Africa, only a small group, possibly as few as 150 to 1,000 people, crossed the Red Sea. The group that crossed the Red Sea travelled along the coastal route around
Arabia
The Arabian Peninsula (, , or , , ) or Arabia, is a peninsula in West Asia, situated north-east of Africa on the Arabian plate. At , comparable in size to India, the Arabian Peninsula is the largest peninsula in the world.
Geographically, the ...
and the
Persian Plateau to India, which appears to have been the first major settling point. argued for the route along the southern coastline of Asia, across about , reaching Australia by around 50,000 years ago.
Today at the
Bab-el-Mandeb straits, the
Red Sea
The Red Sea is a sea inlet of the Indian Ocean, lying between Africa and Asia. Its connection to the ocean is in the south, through the Bab-el-Mandeb Strait and the Gulf of Aden. To its north lie the Sinai Peninsula, the Gulf of Aqaba, and th ...
is about wide, but 50,000 years ago sea levels were lower (owing to glaciation) and the water channel was much narrower. Though the straits were never completely closed, they were narrow enough to have enabled crossing using simple rafts, and there may have been islands in between. Shell
middens 125,000 years old have been found in
Eritrea
Eritrea, officially the State of Eritrea, is a country in the Horn of Africa region of East Africa, with its capital and largest city being Asmara. It is bordered by Ethiopia in the Eritrea–Ethiopia border, south, Sudan in the west, and Dj ...
,
indicating that the diet of early humans included seafood obtained by
beachcombing
Beachcombing is an activity that consists of an individual "combing" (or searching) the beach and the intertidal zone, looking for things of value, interest or utility. A beachcomber is a person who participates in the activity of beachcombing ...
.
Toba eruption
The dating of the Southern Dispersal is a matter of dispute. It may have happened either pre- or post-Toba, a catastrophic volcanic eruption that took place between 69,000 and 77,000 years ago at the site of present-day
Lake Toba in Sumatra, Indonesia. Stone tools discovered below the layers of ash deposited in India may point to a pre-Toba dispersal but the source of the tools is disputed. An indication for post-Toba is haplo-group L3, that originated before the dispersal of humans out of Africa and can be dated to 60,000–70,000 years ago, "suggesting that humanity left Africa a few thousand years after Toba". Some research showing slower than expected genetic mutations in human DNA was published in 2012, indicating a revised dating for the migration to between 90,000 and 130,000 years ago.
Some more recent research suggests a migration out-of-Africa of around 50,000-65,000 years ago of the ancestors of modern non-African populations, similar to most previous estimates.
West Asia
Following the fossils dating 80,000 to 120,000 years ago from
Qafzeh and
Es-Skhul Caves in Israel there are no ''H. sapiens'' fossils in the
Levant
The Levant ( ) is the subregion that borders the Eastern Mediterranean, Eastern Mediterranean sea to the west, and forms the core of West Asia and the political term, Middle East, ''Middle East''. In its narrowest sense, which is in use toda ...
until the
Manot 1 fossil from
Manot Cave in Israel, dated to 54,700 years ago, though the dating was questioned by . The lack of fossils and stone tool industries that can be safely associated with modern humans in the Levant has been taken to suggest that modern humans were outcompeted by Neanderthals until around 55,000 years ago, who would have placed a barrier on modern human dispersal out of Africa through the Northern Route. Climate reconstructions also support a Southern Route dispersal of modern humans as the
Bab-el-Mandeb strait experienced a climate more conductive to human migration than the northern landbridge to the Levant during the major human dispersal out of Africa.
A 2023 study proposed that Eurasians and Africans genetically diverged ~100,000 years ago. Main Eurasians then lived in the Saudi Peninsula, genetically isolated from at least 85 kya, before expanding north 54 kya. For reference, Homo sapiens and Neanderthals diverged ~500 kya.
Oceania
Fossils from
Lake Mungo, Australia, have been dated to about 42,000 years ago.
Other fossils from a site called
Madjedbebe have been dated to at least 65,000 years ago, though some researchers doubt this early estimate and date the Madjedbebe fossils at about 50,000 years ago at the oldest.
Phylogenetic data suggests that an early Eastern Eurasian (Eastern non-African) meta-population trifurcated somewhere in
eastern South Asia, and gave rise to the Australo-Papuans, the Ancient Ancestral South Indians (AASI), as well as East/Southeast Asians, although Papuans may have also received some gene flow from an earlier group (xOoA), around 2%, next to additional archaic admixture in the
Sahul region.
[Genetics and material culture support repeated expansions into Paleolithic Eurasia from a population hub out of Africa, Vallini et al. 2022 (April 4, 2022) Quote: "''Taken together with a lower bound of the final settlement of Sahul at 37 kya it is reasonable to describe Papuans as either an almost even mixture between East-Eurasians and a lineage basal to West and East-Eurasians which occurred sometimes between 45 and 38kya, or as a sister lineage of East-Eurasians with or without a minor basal OoA or xOoA contribution. We here chose to parsimoniously describe Papuans as a simple sister group of Tianyuan, cautioning that this may be just one out of six equifinal possibilities.''"]
According to one study, Papuans could have either formed from a mixture between an East Eurasian lineage and lineage basal to West and East Asians, or as a sister lineage of East Asians with or without a minor basal OoA or xOoA contribution.
A Holocene hunter-gatherer sample (Leang_Panninge) from
South Sulawesi
South Sulawesi () is a Provinces of Indonesia, province in the South Peninsula, Sulawesi, southern peninsula of Sulawesi, Indonesia. The Selayar Islands archipelago to the south of Sulawesi is also part of the province. The capital and largest ci ...
was found to be genetically in between East-Eurasians and Australo-Papuans. The sample could be modeled as ~50% Papuan-related and ~50% Basal-East Asian-related (Andamanese Onge or Tianyuan). The authors concluded that Basal-East Asian ancestry was far more widespread and the peopling of Insular Southeast Asia and Oceania was more complex than previously anticipated.
East and Southeast Asia
In China, the
Liujiang man
The Liujiang man () is among the earliest modern humans (''Homo sapiens'') found in East Asia.
The remains were discovered in the Tongtianyan Cave (通天岩) in Liujiang, Guangxi, China.
The remains were excavated in 1958 and consist of a w ...
( zh, c=柳江人) is among the earliest modern humans found in
East Asia
East Asia is a geocultural region of Asia. It includes China, Japan, Mongolia, North Korea, South Korea, and Taiwan, plus two special administrative regions of China, Hong Kong and Macau. The economies of Economy of China, China, Economy of Ja ...
.
The date most commonly attributed to the remains is 67,000 years ago.
High rates of variability yielded by various dating techniques carried out by different researchers place the most widely accepted range of dates with 67,000 BP as a minimum, but do not rule out dates as old as 159,000 BP.
claim that modern human teeth have been found in China dating to at least 80,000 years ago.
Tianyuan man from China has a probable date range between 38,000 and 42,000 years ago, while
Liujiang man
The Liujiang man () is among the earliest modern humans (''Homo sapiens'') found in East Asia.
The remains were discovered in the Tongtianyan Cave (通天岩) in Liujiang, Guangxi, China.
The remains were excavated in 1958 and consist of a w ...
from the same region has a probable date range between 67,000 and 159,000 years ago. According to 2013 DNA tests, Tianyuan man is related "to many present-day
Asians and
Native Americans".
Tianyuan is similar in
morphology to Liujiang man, and some
Jōmon period
In Japanese history, the is the time between , during which Japan was inhabited by the Jōmon people, a diverse hunter-gatherer and early agriculturalist population united by a common culture, which reached a considerable degree of sedentism an ...
modern humans found in Japan, as well as modern East and Southeast Asians.
A 2021 study about the population history of Eastern Eurasia, concluded that distinctive
Basal-East Asian (East-Eurasian) ancestry originated in
Mainland Southeast Asia
Mainland Southeast Asia (historically known as Indochina and the Indochinese Peninsula) is the continental portion of Southeast Asia. It lies east of the Indian subcontinent and south of Mainland China and is bordered by the Indian Ocean to th ...
at ~50,000BC from a distinct southern Himalayan route, and expanded through multiple migration waves southwards and northwards respectively.
Americas
Genetic studies concluded that
Native Americans descended from a single founding population that initially split from a Basal-East Asian source population in Mainland Southeast Asia around 36,000 years ago, at the same time at which the proper
Jōmon people split from Basal-East Asians, either together with Ancestral Native Americans or during a separate expansion wave. They also show that the basal northern and southern Native American branches, to which all other Indigenous peoples belong, diverged around 16,000 years ago. An indigenous American sample from 16,000BC in
Idaho
Idaho ( ) is a landlocked U.S. state, state in the Pacific Northwest and Mountain states, Mountain West subregions of the Western United States. It borders Montana and Wyoming to the east, Nevada and Utah to the south, and Washington (state), ...
, which is craniometrically similar to modern Native Americans as well as
Paleosiberians, was found to have largely East-Eurasian ancestry and showed high affinity with contemporary East Asians, as well as Jōmon period samples of Japan, confirming that Ancestral Native Americans split from an East-Eurasian source population in Eastern Siberia.
Europe
According to , an early offshoot from the southern dispersal with haplogroup N followed the Nile from East Africa, heading northwards and crossing into Asia through the
Sinai. This group then branched, some moving into Europe and others heading east into Asia. This hypothesis is supported by the relatively late date of the arrival of modern humans in Europe as well as by archaeological and DNA evidence. Based on an analysis of 55 human mitochondrial genomes (mtDNAs) of hunter-gatherers, argue for a "rapid single dispersal of all non-Africans less than 55,000 years ago". By 45,000 years ago, modern humans are known to have reached northwestern Europe.
Genetic reconstruction
Mitochondrial haplogroups
Within Africa

The first lineage to branch off from
Mitochondrial Eve was
L0. This haplogroup is found in high proportions among the
San of Southern Africa and the
Sandawe of East Africa. It is also found among the
Mbuti people. These groups branched off early in human history and have remained relatively genetically isolated since then.
Haplogroups L1,
L2, and
L3 are descendants of L1–L6, and are largely confined to Africa. The macro haplogroups
M and
N, which are the lineages of the rest of the world outside Africa, descend from L3. L3 is about 70,000 years old, while haplogroups M and N are about 65–55,000 years old.
The relationship between such gene trees and demographic history is still debated when applied to dispersals.
Of all the lineages present in Africa, the female descendants of only one lineage,
mtDNA haplogroup L3, are found outside Africa. If there had been several migrations, one would expect descendants of more than one lineage to be found. L3's female descendants, the
M and
N haplogroup lineages, are found in very low frequencies in Africa (although
haplogroup M1 populations are very ancient and diversified in
North
North is one of the four compass points or cardinal directions. It is the opposite of south and is perpendicular to east and west. ''North'' is a noun, adjective, or adverb indicating Direction (geometry), direction or geography.
Etymology
T ...
and
North-east Africa) and appear to be more recent arrivals. A possible explanation is that these mutations occurred in East Africa shortly before the exodus and became the dominant haplogroups thereafter by means of the
founder effect
In population genetics, the founder effect is the loss of genetic variation that occurs when a new population is established by a very small number of individuals from a larger population. It was first fully outlined by Ernst Mayr in 1942, us ...
. Alternatively, the mutations may have arisen shortly afterwards.
Southern Route and haplogroups M and N
Results from mtDNA collected from aboriginal Malaysians called
Orang Asli
The Orang Asli are a Homogeneity and heterogeneity, heterogeneous Indigenous peoples, indigenous population forming a national minority in Malaysia. They are the oldest inhabitants of Peninsular Malaysia.
As of 2017, the Orang Asli accounted f ...
indicate that the haplogroups M and N share characteristics with original African groups from approximately 85,000 years ago, and share characteristics with sub-haplogroups found in coastal south-east Asian regions, such as Australasia, the Indian subcontinent and throughout continental Asia, which had dispersed and separated from their African progenitor approximately 65,000 years ago. This southern coastal dispersal would have occurred before the dispersal through the Levant approximately 45,000 years ago. This hypothesis attempts to explain why haplogroup N is predominant in Europe and why haplogroup M is absent in Europe. Evidence of the coastal migration is thought to have been destroyed by the rise in sea levels during the
Holocene
The Holocene () is the current geologic time scale, geological epoch, beginning approximately 11,700 years ago. It follows the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene to ...
epoch. Alternatively, a small European founder population that had expressed haplogroup M and N at first, could have lost haplogroup M through random
genetic drift
Genetic drift, also known as random genetic drift, allelic drift or the Wright effect, is the change in the Allele frequency, frequency of an existing gene variant (allele) in a population due to random chance.
Genetic drift may cause gene va ...
resulting from a
bottleneck
Bottleneck may refer to:
* the narrowed portion (neck) of a bottle
Science and technology
* Bottleneck (engineering), where the performance of an entire system is limited by a single component
* Bottleneck (network), in a communication network
* ...
(i.e. a
founder effect
In population genetics, the founder effect is the loss of genetic variation that occurs when a new population is established by a very small number of individuals from a larger population. It was first fully outlined by Ernst Mayr in 1942, us ...
).
The group that crossed the Red Sea travelled along the coastal route around
Arabia
The Arabian Peninsula (, , or , , ) or Arabia, is a peninsula in West Asia, situated north-east of Africa on the Arabian plate. At , comparable in size to India, the Arabian Peninsula is the largest peninsula in the world.
Geographically, the ...
and
Persia
Iran, officially the Islamic Republic of Iran (IRI) and also known as Persia, is a country in West Asia. It borders Iraq to the west, Turkey, Azerbaijan, and Armenia to the northwest, the Caspian Sea to the north, Turkmenistan to the nort ...
until reaching India.
Haplogroup M is found in high frequencies along the southern coastal regions of Pakistan and India and it has the greatest diversity in India, indicating that it is here where the mutation may have occurred. Sixty percent of the Indian population belong to
Haplogroup M. The indigenous people of the
Andaman Islands also belong to the M lineage. The Andamanese are thought to be offshoots of some of the earliest inhabitants in Asia because of their long isolation from the mainland. They are evidence of the coastal route of early settlers that extends from India to
Thailand
Thailand, officially the Kingdom of Thailand and historically known as Siam (the official name until 1939), is a country in Southeast Asia on the Mainland Southeast Asia, Indochinese Peninsula. With a population of almost 66 million, it spa ...
and Indonesia all the way to eastern
New Guinea
New Guinea (; Hiri Motu: ''Niu Gini''; , fossilized , also known as Papua or historically ) is the List of islands by area, world's second-largest island, with an area of . Located in Melanesia in the southwestern Pacific Ocean, the island is ...
. Since M is found in high frequencies in highlanders from New Guinea and the Andamanese and New Guineans have dark skin and
Afro-textured hair, some scientists think they are all part of the same wave of migrants who departed across the Red Sea ~60,000 years ago in the
Great Coastal Migration. The proportion of haplogroup M increases eastwards from
Arabia
The Arabian Peninsula (, , or , , ) or Arabia, is a peninsula in West Asia, situated north-east of Africa on the Arabian plate. At , comparable in size to India, the Arabian Peninsula is the largest peninsula in the world.
Geographically, the ...
to India; in eastern India, M outnumbers N by a ratio of 3:1. Crossing into Southeast Asia, haplogroup N (mostly in the form of derivatives of its R subclade) reappears as the predominant lineage. M is predominant in East Asia, but amongst
Indigenous Australians
Indigenous Australians are people with familial heritage from, or recognised membership of, the various ethnic groups living within the territory of contemporary Australia prior to History of Australia (1788–1850), British colonisation. The ...
, N is the more common lineage. This haphazard distribution of Haplogroup N from Europe to Australia can be explained by
founder effect
In population genetics, the founder effect is the loss of genetic variation that occurs when a new population is established by a very small number of individuals from a larger population. It was first fully outlined by Ernst Mayr in 1942, us ...
s and
population bottlenecks.
Autosomal DNA

A 2002 study of African, European, and Asian populations, found greater genetic diversity among Africans than among Eurasians, and that genetic diversity among Eurasians is largely a subset of that among Africans, supporting the out of Africa model. A large study by Coop ''et al''. (2009) found evidence for
natural selection
Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the Heredity, heritable traits characteristic of a population over generation ...
in
autosomal
An autosome is any chromosome that is not a sex chromosome. The members of an autosome pair in a diploid cell have the same morphology, unlike those in allosomal (sex chromosome) pairs, which may have different structures. The DNA in autosome ...
DNA outside of Africa. The study distinguishes non-African sweeps (notably
KITLG variants associated with
skin color), West-Eurasian sweeps (
SLC24A5) and East-Asian sweeps (
MC1R, relevant to skin color). Based on this evidence, the study concluded that human populations encountered novel selective pressures as they expanded out of Africa.
[;
summary in ] MC1R and its relation to skin color had already been discussed by . According to this study, Papua New Guineans continued to be exposed to selection for dark skin color so that, although these groups are distinct from Africans in other places, the allele for dark skin color shared by contemporary Africans, Andamanese and New Guineans is an archaism. suggest
convergent evolution
Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last comm ...
. A 2014 study by Gurdasani et al. indicates that the higher genetic diversity in Africa was further increased in some regions by relatively recent Eurasian migrations affecting parts of Africa.
Pathogen DNA
Another promising route towards reconstructing human genetic genealogy is via the
JC virus (JCV), a type of human
polyomavirus which is carried by 70–90 percent of humans and which is usually transmitted vertically, from parents to offspring, suggesting codivergence with human populations. For this reason, JCV has been used as a genetic marker for human evolution and migration. This method does not appear to be reliable for the migration out of Africa; in contrast to human genetics, JCV strains associated with African populations are not basal. From this conclude that either a basal African strain of JCV has become extinct or that the original infection with JCV post-dates the migration from Africa.
Admixture of archaic and modern humans
Evidence for
archaic human species (descended from ''
Homo heidelbergensis
''Homo heidelbergensis'' is a species of archaic human from the Middle Pleistocene of Europe and Africa, as well as potentially Asia depending on the taxonomic convention used. The species-level classification of ''Homo'' during the Middle Pleis ...
'') having
interbred with modern humans outside of Africa, was discovered in the 2010s. This concerns primarily
Neanderthal
Neanderthals ( ; ''Homo neanderthalensis'' or sometimes ''H. sapiens neanderthalensis'') are an extinction, extinct group of archaic humans who inhabited Europe and Western and Central Asia during the Middle Pleistocene, Middle to Late Plei ...
admixture in all modern populations except for
Sub-Saharan Africans but evidence has also been presented for
Denisova hominin admixture in
Australasia
Australasia is a subregion of Oceania, comprising Australia, New Zealand (overlapping with Polynesia), and sometimes including New Guinea and surrounding islands (overlapping with Melanesia). The term is used in a number of different context ...
(i.e. in
Melanesians
Melanesians are the predominant and Indigenous peoples of Oceania, indigenous inhabitants of Melanesia, in an area stretching from New Guinea to the Fiji Islands. Most speak one of the many languages of the Austronesian languages, Austronesian l ...
,
Aboriginal Australians
Aboriginal Australians are the various indigenous peoples of the Mainland Australia, Australian mainland and many of its islands, excluding the ethnically distinct people of the Torres Strait Islands.
Humans first migrated to Australia (co ...
and some
Negritos). The rate of Neanderthal admixture to European and Asian populations as of 2017 has been estimated at between about 2–3%.
Archaic admixture in some Sub-Saharan African populations hunter-gatherer groups (
Biaka Pygmies and
San), derived from archaic hominins that broke away from the modern human lineage around 700,000 years ago, was discovered in 2011. The rate of admixture was estimated at 2%.
Admixture from archaic hominins of still earlier divergence times, estimated at 1.2 to 1.3 million years ago, was found in
Pygmies
In anthropology, pygmy peoples are ethnic groups whose average height is unusually short. The term pygmyism is used to describe the phenotype of endemic short stature (as opposed to disproportionate dwarfism occurring in isolated cases in a po ...
,
Hadza and five
Sandawe in 2012.
From an analysis of
Mucin 7, a highly divergent haplotype that has an estimated coalescence time with other variants around 4.5 million years BP and is specific to African populations, it is inferred to have been derived from interbreeding between African modern and archaic humans.
A study published in 2020 found that the
Yoruba and
Mende populations of West Africa derive between 2% and 19% of their genome from an as-yet unidentified archaic hominin population that likely diverged before the split of modern humans and the ancestors of Neanderthals and Denisovans.
Stone tools
In addition to genetic analysis, Petraglia ''et al.'' also examines the small stone tools (
microlith
A microlith is a small Rock (geology), stone tool usually made of flint or chert and typically a centimetre or so in length and half a centimetre wide. They were made by humans from around 60,000 years ago, across Europe, Africa, Asia and Austral ...
ic materials) from the Indian subcontinent and explains the expansion of population based on the reconstruction of paleoenvironment. He proposed that the stone tools could be dated to 35 ka in South Asia, and the new technology might be influenced by environmental change and population pressure.
History of the theory
Classical paleoanthropology

The cladistic relationship of humans with the African
apes was suggested by
Charles Darwin
Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English Natural history#Before 1900, naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all speci ...
after studying the behaviour of African
apes, one of which was displayed at the
London Zoo.
The anatomist
Thomas Huxley had also supported the hypothesis and suggested that African apes have a close evolutionary relationship with humans.
These views were opposed by the German biologist
Ernst Haeckel
Ernst Heinrich Philipp August Haeckel (; ; 16 February 1834 – 9 August 1919) was a German zoologist, natural history, naturalist, eugenics, eugenicist, Philosophy, philosopher, physician, professor, marine biology, marine biologist and artist ...
, who was a proponent of the
Out of Asia theory. Haeckel argued that humans were more closely related to the primates of South-east Asia and rejected Darwin's African hypothesis.
In ''
The Descent of Man
''The Descent of Man, and Selection in Relation to Sex'' is a book by English natural history, naturalist Charles Darwin, first published in 1871, which applies evolutionary theory to human evolution, and details his theory of sexual selection, ...
'', Darwin speculated that humans had descended from apes, which still had small brains but walked upright, freeing their hands for uses which favoured intelligence; he thought such apes were African:
In 1871, there were hardly any human fossils of ancient hominins available. Almost fifty years later, Darwin's speculation was supported when anthropologists began finding fossils of ancient small-brained hominins in several areas of Africa (
list of hominina fossils). The hypothesis of ''recent'' (as opposed to
archaic) African origin developed in the 20th century. The "Recent African origin" of modern humans means "single origin" (monogenism) and has been used in various contexts as an antonym to polygenism. The debate in anthropology had swung in favour of monogenism by the mid-20th century. Isolated proponents of polygenism held forth in the mid-20th century, such as
Carleton Coon, who thought as late as 1962 that ''H. sapiens'' arose five times from ''H. erectus'' in five places.
Multiregional origin hypothesis
The historical alternative to the recent origin model is the
multiregional origin of modern humans, initially proposed by
Milford Wolpoff in the 1980s. This view proposes that the derivation of anatomically modern human populations from ''H. erectus'' at the beginning of the
Pleistocene
The Pleistocene ( ; referred to colloquially as the ''ice age, Ice Age'') is the geological epoch (geology), epoch that lasted from to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fin ...
1.8 million years BP, has taken place within a continuous world population. The hypothesis necessarily rejects the assumption of an
infertility barrier between ancient Eurasian and African populations of ''Homo''. The hypothesis was controversially debated during the late 1980s and the 1990s. The now-current terminology of "recent-origin" and "Out of Africa" became current in the context of this debate in the 1990s. Originally seen as an antithetical alternative to the recent origin model, the multiregional hypothesis in its original "strong" form is obsolete, while its various modified weaker variants have become variants of a view of "recent origin" combined with
archaic admixture. Stringer (2014) distinguishes the original or "classic" Multiregional model as having existed from 1984 (its formulation) until 2003, to a "weak" post-2003 variant that has "shifted close to that of the Assimilation Model".
Mitochondrial analyses
In the 1980s,
Allan Wilson together with
Rebecca L. Cann and
Mark Stoneking worked on genetic dating of the matrilineal most recent common ancestor of modern human populations (dubbed "
Mitochondrial Eve"). To identify informative
genetic marker
A genetic marker is a gene or DNA sequence with a known location on a chromosome that can be used to identify individuals or species. It can be described as a variation (which may arise due to mutation or alteration in the genomic loci) that can ...
s for tracking human evolutionary history, Wilson concentrated on
mitochondrial DNA
Mitochondrial DNA (mtDNA and mDNA) is the DNA located in the mitochondrion, mitochondria organelles in a eukaryotic cell that converts chemical energy from food into adenosine triphosphate (ATP). Mitochondrial DNA is a small portion of the D ...
(mtDNA), which is maternally inherited. This DNA material mutates quickly, making it easy to plot changes over relatively short times. With his discovery that human mtDNA is genetically much less diverse than chimpanzee mtDNA, Wilson concluded that modern human populations had diverged recently from a single population while older human species such as
Neanderthal
Neanderthals ( ; ''Homo neanderthalensis'' or sometimes ''H. sapiens neanderthalensis'') are an extinction, extinct group of archaic humans who inhabited Europe and Western and Central Asia during the Middle Pleistocene, Middle to Late Plei ...
s and ''
Homo erectus
''Homo erectus'' ( ) is an extinction, extinct species of Homo, archaic human from the Pleistocene, spanning nearly 2 million years. It is the first human species to evolve a humanlike body plan and human gait, gait, to early expansions of h ...
'' had become extinct.
With the advent of
archaeogenetics
Archaeogenetics is the study of ancient DNA using various molecular genetic methods and DNA resources. This form of genetic analysis can be applied to human, animal, and plant specimens. Ancient DNA can be extracted from various fossilized spec ...
in the 1990s, the dating of mitochondrial and
Y-chromosomal haplogroups became possible with some confidence. By 1999, estimates ranged around 150,000 years for the
mt-MRCA and 60,000 to 70,000 years for the migration out of Africa.
From 2000 to 2003, there was controversy about the mitochondrial DNA of "
Mungo Man 3" (LM3) and its possible bearing on the
multiregional hypothesis. LM3 was found to have more than the expected number of sequence differences when compared to modern human DNA (
CRS).
Comparison of the mitochondrial DNA with that of ancient and modern
aborigines, led to the conclusion that Mungo Man fell outside the range of genetic variation seen in Aboriginal Australians and was used to support the multiregional origin hypothesis. A reanalysis of LM3 and other ancient specimens from the area published in 2016, showed it to be akin to modern Aboriginal Australian sequences, inconsistent with the results of the earlier study.
Y-chromosome analyses

The
Y chromosome
The Y chromosome is one of two sex chromosomes in therian mammals and other organisms. Along with the X chromosome, it is part of the XY sex-determination system, in which the Y is the sex-determining chromosome because the presence of the ...
, which is paternally inherited, does not go through much recombination and thus stays largely the same after inheritance. Similar to Mitochondrial Eve, this could be studied to track the male most recent common ancestor ("
Y-chromosomal Adam" or Y-MRCA).
The most basal lineages have been detected in
West
West is one of the four cardinal directions or points of the compass. It is the opposite direction from east and is the direction in which the Sun sets on the Earth.
Etymology
The word "west" is a Germanic word passed into some Romance langu ...
,
Northwest
The points of the compass are a set of horizontal, radially arrayed compass directions (or azimuths) used in navigation and cartography. A '' compass rose'' is primarily composed of four cardinal directions—north, east, south, and west— ...
and
Central Africa
Central Africa (French language, French: ''Afrique centrale''; Spanish language, Spanish: ''África central''; Portuguese language, Portuguese: ''África Central'') is a subregion of the African continent comprising various countries accordin ...
, suggesting plausibility for the Y-MRCA living in the general region of "Central-Northwest Africa".
A
Stanford University School of Medicine study was done by comparing Y-chromosome sequences and mtDNA in 69 men from different geographic regions and constructing a family tree. It was found that the Y-MRCA lived between 120,000 and 156,000, and the Mitochondrial Eve lived between 99,000 and 148,000 years ago, which not only predates some proposed waves of migration, but also meant that both lived in the
African continent around the same time period.
Another study finds a plausible placement in "the north-western quadrant of the African continent" for the emergence of the A1b haplogroup. The 2013 report of
haplogroup A00 found among the
Mbo people of western present-day
Cameroon
Cameroon, officially the Republic of Cameroon, is a country in Central Africa. It shares boundaries with Nigeria to the west and north, Chad to the northeast, the Central African Republic to the east, and Equatorial Guinea, Gabon, and the R ...
is also compatible with this picture.
The revision of Y-chromosomal phylogeny since 2011 has affected estimates for the likely geographical origin of Y-MRCA as well as estimates on time depth. By the same reasoning, future discovery of presently-unknown archaic haplogroups in living people would again lead to such revisions. In particular, the possible presence of between 1% and 4%
Neanderthal-derived DNA in Eurasian genomes implies that the (unlikely) event of a discovery of a single living Eurasian male exhibiting a Neanderthal patrilineal line would immediately push back T-MRCA ("time to MRCA") to at least twice its current estimate. However, the discovery of a Neanderthal Y-chromosome by Mendez ''et al''. was tempered by a 2016 study that suggests the extinction of Neanderthal patrilineages, as the lineage inferred from the Neanderthal sequence is outside of the range of
contemporary human genetic variation.
Questions of geographical origin would become part of the debate on Neanderthal evolution from ''
Homo erectus
''Homo erectus'' ( ) is an extinction, extinct species of Homo, archaic human from the Pleistocene, spanning nearly 2 million years. It is the first human species to evolve a humanlike body plan and human gait, gait, to early expansions of h ...
''.
See also
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Human timeline
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Notes
References
Sources
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Further reading
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External links
* Encyclopædia Britannica,
Human Evolution'
Human Timeline (Interactive)–
Smithsonian,
National Museum of Natural History
The National Museum of Natural History (NMNH) is a natural history museum administered by the Smithsonian Institution, located on the National Mall in Washington, D.C., United States. It has free admission and is open 364 days a year. With 4.4 ...
(August 2016).
Age constraints for the Trachilos footprints from CreteNature
Nature is an inherent character or constitution, particularly of the Ecosphere (planetary), ecosphere or the universe as a whole. In this general sense nature refers to the Scientific law, laws, elements and phenomenon, phenomena of the physic ...
(October 2021).
{{Authority control
Prehistoric migrations
Human evolution
Human mitochondrial genetics
Human genetic history