Recent African origin of modern humans
The recent African origin of modern humans, also called the "Out of Africa" theory (OOA), recent single-origin hypothesis (RSOH), replacement hypothesis or recent African origin model (RAO), is, in paleoanthropology, the dominant model of the geographic origin and early migration of anatomically modern humans (Homo sapiens), which proposes a single area of origin for modern humans. According to this model, modern humans evolved in East Africa, and started to disperse through the world roughly 50,000 to 100,000 years ago. The single-origin hypothesis is cited as having the scientific consensus as of the mid-2000s.
The major competing hypothesis of "recent single origin" has been the multiregional origin of modern humans, which envisions a wave of Homo sapiens migrating earlier from Africa and interbreeding with local Homo erectus populations in multiple regions of the globe.
In the 2010s, the discovery of evidence of archaic admixture of modern humans outside of Africa with Neanderthals and Denisovans has complicated the picture. As of 2011, it appears likely that there were two waves of migration out of Africa. The first took place between 130,000–115,000 years ago via northern Africa, and appears to have mostly died out or retreated (although there is some evidence of a presence of modern humans in China about 80,000 years ago). A second dispersal took place via the so-called Southern Route, following the southern coastline of Asia, which led to the lasting colonization of Eurasia and Australia by around 50,000 years ago. Europe was populated by an early offshoot which settled the Near East and Europe (post-Toba hypothesis).
- 1 African origins of Homo sapiens
- 2 Dispersals out of Africa
- 3 Genetic reconstruction
- 4 History of the theory
- 5 See also
- 6 References
- 7 External links
African origins of Homo sapiens
Anatomically modern humans originated in Africa by about 200,000 years ago. The trend in cranial expansion and the acheulean elaboration of stone tool technologies which occurred between 400,000 years ago and the second interglacial period in the Middle Pleistocene (around 250,000 years ago) provide evidence for a transition from Homo erectus to H. sapiens. In the Recent African Origin (RAO) scenario, migration within and out of Africa eventually replaced the earlier dispersed H. erectus.
Beginning about 100,000 years ago, evidence of more sophisticated technology and artwork begins to emerge and by 50,000 years ago, fully modern behaviour becomes more prominent. Stone tools show regular patterns that are reproduced or duplicated with more precision while tools made of bone and antler appear for the first time.
Fossil finds outside Africa
Fossils of early Homo sapiens were found in Qafzeh cave in Israel and have been dated to 80,000 to 100,000 years ago. A fossil of a modern human dated to 54,700 years ago was found in Manot Cave in Israel, named Manot 1, though the dating was questioned by Groucutt et al. (2015). Fossils from Lake Mungo, Australia, have been dated to about 42,000 years ago. The Tianyuan cave remains in China's Liujiang region have a probable date range between 38,000 and 42,000 years ago. The Tianyuan specimens are most similar in morphology to Minatogawa Man, modern humans dated between 17,000 and 19,000 years ago and found on Okinawa Island, Japan.
Dispersals out of Africa
Anatomically modern humans originated in Africa by about 200,000 years ago. The accepted theory is that there were two dispersals from African populations, an early one via northern Africa which left traces in some human populations, and a second one which populated the entire world.
The first dispersal took place between 130,000–115,000 years ago via northern Africa, but died out or retreated. Chinese researchers question this extinction, claiming that modern humans were present in China already 80,000 years ago.
A second dispersal took place 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, crossing about 250 kilometres (155 mi) of sea, and colonized Australia by around 50,000 years ago. According to this theory, Europe was populated either by a migration out of India, which was repopulated from southeast Asia after the Toba-event (pre-Toba hypothesis), or by an early offshoot which settled the Near East and Europe (post-Toba hypothesis).
Early northern Africa dispersal
The early northern Africa dispersal took place between 130,000–115,000 years ago. The discovery of stone tools in the United Arab Emirates in 2011 indicated the presence of modern humans between 100,000 and 125,000 years ago, leading to a resurgence of the "long-neglected" North African route.
Fossils of early Homo sapiens were found in Qafzeh cave in Israel and have been dated 80,000 to 100,000 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.
According to Kuhlwilm and his co-authors, Neanderthals received gene flow from modern humans around 100,000 years ago, from humans which split off from other modern humans around 200,000 years ago. They asserted that "the ancestors of Neanderthals from the Altai Mountains 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 ka by providing the first genetic evidence of such populations."
Chinese research questions the extinction of this early dispersal. Shen, Wang et al. (2002) have dated Liujang Man to 111,000 to 139,000 years before the present. Liu, Martinón-Torres et al. (2015) claim that modern human teeth have been found in China dating to at least 80,000 years ago.
Southern Route dispersal
By some 70,000 years ago, a part of the bearers of mitochondrial haplogroup L3 migrated from East Africa into the Near East. 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 and Persia to India, which appears to be the first major settling point. Wells (2003) argued for the route along the southern coastline of Asia, across about 250 kilometres (155 mi), reaching Australia by around 50,000 years ago.
Today at the Bab-el-Mandeb straits, the Red Sea is about 20 kilometres (12 mi) wide but 50,000 years ago sea levels were 70 m (230 ft) lower (owing to glaciation) and the water was much narrower. Though the straits were never completely closed, they were narrow enough and there may have been islands in between to have enabled crossing using simple rafts. Shell middens 125,000 years old have been found in Eritrea, indicating the diet of early humans included seafood obtained by beachcombing.
Dating: pre-or post-Toba
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. Stone tools discovered below the layers of ash disposed 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". New 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.
Peopling of Europe
According to Meredith (2011), members of one branch of Homo sapiens left Africa at some point between 125,000 and 60,000 years ago and replaced other populations of the genus Homo such as Neanderthals and Homo erectus.
According to Macaulay et al. (2005), an early offshoot from the southern disposal 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, Posth et al. (2016) argue for a "rapid single dispersal of all non-Africans less than 55,000 years ago."
Several authors have argued for multiple dispersals. Wells (2003) describes two waves of migration, first the southern coastal route, then a northern migration into Europe at circa 45,000 years ago.[note 1] This possibility is ruled out by Macaulay et al. (2005) and Posth et al. (2016), arguing for a single coastal dispersal.
The first lineage to branch off from Mitochondrial Eve is 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 L1-6 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 84,000 years old and haplogroup M and N are about 63,000 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, only the female descendants of 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 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 after the departure through the founder effect. 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 and the creation of a phylogentic tree indicate that the hapologroups M and N share characteristics with original African groups from approximately 85,000 years ago and share characteristics with sub-haplogroups among coastal south-east Asian regions, such as Australasia, the Indian subcontinent and throughout continental Asia, which had dispersed and separated from its African origins approximately 65,000 years ago. This southern coastal dispersion would have occurred before the dispersion 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 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 resulting from a bottleneck (i.e. a founder effect).
The group that crossed the Red Sea travelled along the coastal route around Arabia and Persia 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 and Indonesia all the way to Papua New Guinea. 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 to India; in eastern India, M outnumbers N by a ratio of 3:1. Crossing into East Asia, haplogroup N reappears as the dominant lineage. M is predominant in South East Asia but amongst Indigenous Australians N is the more common lineage. This haphazard distribution of Haplogroup N from Europe to Australia can be explained by founder effects and population bottlenecks.
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 in autosomal 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. MC1R and its relation to skin color had already been discussed by Liu, Harding et al. (2000), p. 135. 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. Endicott et al. (2003) suggest convergent evolution.
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 Shackelton et al. (2006) 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) having interbred with modern humans outside of Africa, was discovered in the 2010s. This concerns primarily Neanderthal admixture in all modern populations except for Sub-Saharan Africans but evidence has also been presented for Denisova hominin admixture in Australasia (i.e. in Melanesians, Aboriginal Australians and some Negritos). The rate of admixture has been found to be relatively low (1–10%), and the result is still open to revision, as the presence of archaic markers has also been suggested as being due to shared ancestral traits rather than interbreeding.
In 2011, evidence was presented that for archaic admixture in some Sub-Saharan African populations (Biaka Pygmies and San), derived from archaic hominins that broke away from the modern human lineage around 700,000 years. After a survey for the introgressive haplotypes across Sub-Saharan populations, it was suggested that the admixture event happened with archaic hominins that possibly once inhabited Central Africa.
In addition to genetic analysis, Petraglia et al. also examines the small stone tools (microlithic materials) from 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
The cladistic relationship of humans with the African apes was suggested by Charles Darwin 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, 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, 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 each great region of the world the living mammals are closely related to the extinct species of the same region. It is, therefore, probable that Africa was formerly inhabited by extinct apes closely allied to the gorilla and chimpanzee; and as these two species are now man's nearest allies, it is somewhat more probable that our early progenitors lived on the African continent than elsewhere. But it is useless to speculate on this subject, for an ape nearly as large as a man, namely the Dryopithecus of Lartet, which was closely allied to the anthropomorphous Hylobates, existed in Europe during the Upper Miocene period; and since so remote a period the earth has certainly undergone many great revolutions, and there has been ample time for migration on the largest scale.— Charles Darwin, Descent of Man
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 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".
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 markers for tracking human evolutionary history, Wilson concentrated on mitochondrial DNA (mtDNA), passed from mother to child. 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 Neanderthals and Homo erectus had become extinct. With the advent of archaeogenetics 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 2000–2003, 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 on 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.
- Archaeogenetics of the Near East
- Behavioral modernity
- Coastal migration
- Dawn of Humanity (2015 PBS film)
- Early human migrations
- Genetics and archaeogenetics of South Asia
- Genetic history of Europe
- Genetic history of indigenous peoples of the Americas
- Genetic history of Italy
- Genetic history of North Africa
- Genetic history of the British Isles
- Genetic history of the Iberian Peninsula
- Hofmeyr Skull
- Human timeline
- Identical ancestors point
- Indo-Aryan migration theory
- Sahara pump theory
- The Incredible Human Journey
- Timeline of human evolution
- Kay Young McChesney: "Wells (2003) divided the descendants of men who left Africa into a genealogical tree with 11 lineages. Each genetic marker represents a single-point mutation (SNP) at a specific place in the genome. First, genetic evidence suggests that a small band with the marker M168 migrated out of Africa along the coasts of the Arabian Peninsula and India, through Indonesia, and reached Australia very early, between 60,000 and 50,000 years ago. This very early migration into Australia is also supported by Rasmussen et al. (2011). Second, a group bearing the marker M89 moved out of northeastern Africa into the Middle East 45,000 years ago. From there, the M89 group split into two groups. One group that developed the marker M9 went into Asia about 40,000 years ago. The Asian (M9) group split three ways: into Central Asia (M45), 35,000 years ago; into India (M20), 30,000 years ago; and into China (M122), 10,000 years ago. The Central Asian (M45) group split into two groups: toward Europe (M173), 30,000 years ago and toward Siberia (M242), 20,000 years ago. Finally, the Siberian group (M242) went on to populate North and South America (M3), about 10,000 years ago.
- Liu, Prugnolle et al. (2006). "Currently available genetic and archaeological evidence is supportive of a recent single origin of modern humans in East Africa. However, this is where the consensus on human settlement history ends, and considerable uncertainty clouds any more detailed aspect of human colonization history."
- Stringer, Chris (June 2003). "Human evolution: Out of Ethiopia". Nature. 423 (6941): 692–3, 695. Bibcode:2003Natur.423..692S. doi:10.1038/423692a. PMID 12802315.
- Robert Jurmain; Lynn Kilgore; Wenda Trevathan (20 March 2008). Essentials of Physical Anthropology. Cengage Learning. pp. 266–. ISBN 978-0-495-50939-4. Retrieved 14 June 2011.
- Wolpoff, Milford H.; Hawks, John; Caspari, Rachel (May 2000). "Multiregional, not multiple origins". Am. J. Phys. Anthropol. 112 (1): 129–36. doi:10.1002/(SICI)1096-8644(200005)112:1<129::AID-AJPA11>3.0.CO;2-K. PMID 10766948.
- Prüfer, K.; Racimo, F.; Patterson, N.; Jay, F.; Sankararaman, S.; Sawyer, S.; et al. (2014) [Online 2013]. "The complete genome sequence of a Neanderthal from the Altai Mountains". Nature. 505 (7481): 43–49. Bibcode:2014Natur.505...43P. doi:10.1038/nature12886. PMC . PMID 24352235.
- Armitage, Simon J.; Jasim, Sabah A.; Marks, Anthony E.; Parker, Adrian G.; Usik, Vitaly I.; Uerpmann, Hans-Peter (January 2011). "The southern route "out of Africa": evidence for an early expansion of modern humans into Arabia". Science. 331 (6016): 453–6. Bibcode:2011Sci...331..453A. doi:10.1126/science.1199113. PMID 21273486.
- Balter Michael (January 2011). "Was North Africa the launch pad for modern human migrations?" (PDF). Science. 331 (6013): 20–3. Bibcode:2011Sci...331...20B. doi:10.1126/science.331.6013.20. PMID 21212332.
- Cruciani, Fulvio; Trombetta, Beniamino; Massaia, Andrea; Destro-Bisol, Giovanni; Sellitto, Daniele; Scozzari, Rosaria (June 2011). "A revised root for the human Y chromosomal phylogenetic tree: the origin of patrilineal diversity in Africa". AJHG. 88 (6): 814–8. doi:10.1016/j.ajhg.2011.05.002. PMC . PMID 21601174.
- Smith, Tanya M.; Tafforeau, Paul; Reid, Donald J.; Grün, Rainer; Eggins, Stephen; Boutakiout, Mohamed; Hublin, Jean-Jacques (April 2007). "Earliest evidence of modern human life history in North African early Homo sapiens". PNAS. 104 (15): 6128–33. Bibcode:2007PNAS..104.6128S. doi:10.1073/pnas.0700747104. PMC . PMID 17372199.
- Liu, Martinón-Torres et al. (2015).
See also Modern humans in China ~80,000 years ago (?), Dieneks' Anthropology Blog.
- Macaulay et al. (2005).
- Posth et al. (2016).
See also mtDNA from 55 hunter-gatherers across 35,000 years in Europe, Dienekes' Anthroplogy Bog.
- Eric Delson; Ian Tattersall; John A. Van Couvering (2000). Encyclopedia of human evolution and prehistory. Taylor & Francis. p. 677–. ISBN 978-0-8153-1696-1. Retrieved 14 June 2011.
- White, Tim D.; Asfaw, Berhane; Degusta, David; Gilbert, Henry; Richards, Gary D.; Suwa, Gen; Howell, Clark F. (June 2003). "Pleistocene Homo sapiens from Middle Awash, Ethiopia". Nature. 423 (6941): 742–7. Bibcode:2003Natur.423..742W. doi:10.1038/nature01669. PMID 12802332.
- Hoffecker, John (2009). "The spread of modern humans in Europe". PNAS. 106 (38): 16040–16045. doi:10.1073/pnas.0903446106. PMC . PMID 19571003.
- Tattersall, Ian (2009). "Human origins: Out of Africa". PNAS. 106 (38): 16018–16021. doi:10.1073/pnas.0903207106.
- Liu, Prugnolle et al. (2006).
- Hershkovitz et al. (2015)
See also 55,000-Year-Old Skull Fossil Sheds New Light on Human Migration out of Africa, Science News.
- Bowler, James M.; Johnston, Harvey; Olley, John M.; Prescott, John R.; Roberts, Richard G.; Shawcross, Wilfred; Spooner, Nigel A. (2003). "New ages for human occupation and climatic change at Lake Mungo, Australia". Nature. 421 (6925): 837–40. Bibcode:2003Natur.421..837B. doi:10.1038/nature01383. PMID 12594511.
- Olleya JM, Roberts RG, Yoshida H, Bowler JM (2006). "Single-grain optical dating of grave-infill associated with human burials at Lake Mungo, Australia". Quaternary Science Reviews. 25 (19–20): 2469–2474. Bibcode:2006QSRv...25.2469O. doi:10.1016/j.quascirev.2005.07.022.
- Hu, Yaowu; Shang, Hong; Tong, Haowen; Nehlich, Olaf; Liu, Wu; Zhao, Chaohong; Yu, Jincheng; Wang, Changsui; Trinkaus, Erik; Richards, Michael P. (July 2009). "Stable isotope dietary analysis of the Tianyuan 1 early modern human". Proc. Natl. Acad. Sci. U.S.A. 106 (27): 10971–4. Bibcode:2009PNAS..10610971H. doi:10.1073/pnas.0904826106. PMC . PMID 19581579.
- Brown, Peter (August 1992). "Recent human evolution in East Asia and Australasia". Philos. Trans. R. Soc. Lond., B, Biol. Sci. 337 (1280): 235–42. doi:10.1098/rstb.1992.0101. PMID 1357698.
- Literature: Göran Burenhult: Die ersten Menschen, Weltbild Verlag, 2000. ISBN 3-8289-0741-5
- Beyin (2011).
- Two pronged AMH colonization of Eurasia. Or not
- Appenzeller (2012).
- Scerri, Eleanor M. L.; Drake, Nick A.; Jennings, Richard; Groucutt, Huw S. (1 October 2014). "Earliest evidence for the structure of Homo sapiens populations in Africa". Quaternary Science Reviews. 101: 207–216. doi:10.1016/j.quascirev.2014.07.019.
- Finlayson (2009), p. 68.
- Kuhlwilm et al. (2016).
See also Ancestors of Eastern Neandertals admixed with modern humans 100 thousand years ago, Dienekes'Anthropology Blog.
- Zhivotovsky; Rosenberg, NA; Feldman, MW; et al. (2003). "Features of Evolution and Expansion of Modern Humans, Inferred from Genomewide Microsatellite Markers". American Journal of Human Genetics. 72 (5): 1171–86. doi:10.1086/375120. PMC . PMID 12690579.
- Stix, Gary (2008). "The Migration History of Humans: DNA Study Traces Human Origins Across the Continents". Retrieved 14 June 2011.
- Metspalu M, Kivisild T, Metspalu E, Parik J, Hudjashov G, Kaldma K, Serk P, Karmin M, Behar DM, Gilbert MT, Endicott P, Mastana S, Papiha SS, Skorecki K, Torroni A, Villems R (August 2004). "Most of the extant mtDNA boundaries in south and southwest Asia were likely shaped during the initial settlement of Eurasia by anatomically modern humans". BMC Genet. 5: 26. doi:10.1186/1471-2156-5-26. PMC . PMID 15339343.
- Fernandes et. al (June 2006). "Absence of post-Miocene Red Sea land bridges: biogeographic implications". Journal of Biogeography. 33 (6): 961–966. doi:10.1111/j.1365-2699.2006.01478.x.
- Walter RC, Buffler RT, Bruggemann JH, Guillaume MM, Berhe SM, Negassi B, Libsekal Y, Cheng H, Edwards RL, von Cosel R, Néraudeau D, Gagnon M (May 2000). "Early human occupation of the Red Sea coast of Eritrea during the last interglacial". Nature. 405 (6782): 65–9. doi:10.1038/35011048. PMID 10811218.
- Catherine Brahic (24 November 2012). "Our True Dawn". New Scientist. Reed Business Information (2892): 34–7. ISSN 0262-4079.
- Groucutt et al. (2015).
- Young McChesney 2015.
- Gonder, Mary K.; Mortensen, Holly M.; Reed, Floyd A.; De Sousa, Alexandra; Tishkoff, Sarah A. (March 2007). "Whole-mtDNA genome sequence analysis of ancient African lineages". Mol. Biol. Evol. 24 (3): 757–68. doi:10.1093/molbev/msl209. PMID 17194802.
- Chen, Yu-Sheng; Olckers, Antonel; Schurr, Theodore G.; Kogelnik, Andreas M.; Huoponen, Kirsi; Wallace, Douglas C. (April 2000). "mtDNA variation in the South African Kung and Khwe-and their genetic relationships to other African populations". Am. J. Hum. Genet. 66 (4): 1362–83. doi:10.1086/302848. PMC . PMID 10739760.
- Maca-Meyer, Nicole; González, Ana M.; Larruga, José M.; Flores, Carlos; Cabrera, Vicente M. (13 August 2001). "Major genomic mitochondrial lineages delineate early human expansions". BMC Genet. 2: 13. doi:10.1186/1471-2156-2-13. PMC . PMID 11553319.
- Ingman M, Gyllensten U (July 2003). "Mitochondrial genome variation and evolutionary history of Australian and New Guinean aborigines". Genome Res. 13 (7): 1600–6. doi:10.1101/gr.686603. PMC . PMID 12840039.
- Yu, Ning; et al. (May 2002). "Larger Genetic Differences Within Africans Than Between Africans and Eurasians". Genetics. Genetics Society of America. Retrieved 7 April 2013.
- Coop G, Pickrell, Novembre, Kudaravalli, Li, Absher, Myers, Cavalli-Sforza, Feldman, Pritchard (June 2009). Schierup MH, ed. "The role of geography in human adaptation". PLoS Genet. 5 (6): e1000500. doi:10.1371/journal.pgen.1000500. PMC . PMID 19503611.; summary in Kliman (ed.), Encyclopedia of Evolutionary Biology (2016), p. 451
- Elizabeth Matisoo-Smith, K. Ann Horsburgh, DNA for Archaeologists, Routledge (2016).
- Rasmussen, M.; Guo, X.; Wang, Y.; Lohmueller, K.E.; Rasmussen, S.; Albrechtsen, A.; et al. (2011). "An Aboriginal Australian Genome Reveals Separate Human Dispersals into Asia". Science. 334 (6052): 94–98. doi:10.1126/science.1211177. PMC . PMID 21940856.
- Sánchez-Quinto, F.; Botigué, L.R.; Civit, S.; Arenas, C.; Ávila-Arcos, M.C.; Bustamante, C.D.; et al. (2012). "North African Populations Carry the Signature of Admixture with Neandertals". PLoS ONE. 7 (10): e47765. doi:10.1371/journal.pone.0047765. PMC . PMID 23082212.
- Hammer, M.F.; Woerner, A.E.; Mendez, F.L.; Watkins, J.C.; Wall, J.D. (2011). "Genetic evidence for archaic admixture in Africa". Proceedings of the National Academy of Sciences. 108 (37): 15123–15128. doi:10.1073/pnas.1109300108. PMC . PMID 21896735.
- Petraglia, M.; Clarkson, C.; Boivin, N.; Haslam, M.; Korisettar, R.; Chaubey, G.; Arnold, L. (2009). "Population increase and environmental deterioration correspond with microlithic innovations in South Asia ca. 35,000 years ago". Proceedings of the National Academy of Sciences. 106 (30): 12261–12266. doi:10.1073/pnas.0810842106. PMC . PMID 19620737.
- Peter Lafreniere (22 September 2010). Adaptive Origins: Evolution and Human Development. Taylor & Francis. p. 90. ISBN 978-0-8058-6012-2. Retrieved 14 June 2011.
- Robinson D, Ash PM (2010). The Emergence of Humans: An Exploration of the Evolutionary Timeline. New York: Wiley. ISBN 0-470-01315-X.
- Palmer D (2006). Prehistoric Past Revealed: The Four Billion Year History of Life on Earth. Berkeley: University of California Press. p. 43. ISBN 0-520-24827-9.
- Regal B (2004). Human evolution: a guide to the debates. Santa Barbara, Calif: ABC-CLIO. pp. 73–75. ISBN 1-85109-418-0.
- "The descent of man Chapter 6 – On the Affinities and Genealogy of Man". Darwin-online.org.uk. Retrieved 11 January 2011.
- Jackson JP Jr (2001). "'In Ways Unacademical': The Reception of Carleton S. Coon's The Origin of Races" (pdf). Journal of the History of Biology. 34 (2): 247–285. doi:10.1023/A:1010366015968.
- Stringer, C. B.; Andrews, P. (1988). "Genetic and fossil evidence for the origin of modern humans". Science. 239 (4845): 1263–1268. doi:10.1126/science.3125610. PMID 3125610. Stringer, C.; Bräuer, G. (1994). "Methods, misreading, and bias". American Anthropologist. 96 (2): 416–424. doi:10.1525/aa.1994.96.2.02a00080.
Stringer, C. B. (1992). "Replacement, continuity and the origin of Homo sapiens". In: Continuity or replacement? Controversies in Homo sapiens evolution. F. H. Smith (ed). Rotterdam: Balkema. pp. 9–24.
Bräuer, G.; Stringer, C. (1997). "Models, polarization, and perspectives on modern human origins". In: Conceptual issues in modern human origins research. New York: Aldine de Gruyter. pp. 191–201.
- Liu Wu in Zhisheng, Weijian Zhou (eds.), Quaternary Geology VSP (1997), p. 24.
- Stringer, C. (2001). "Modern human origins—distinguishing the models". Afr. Archaeol. Rev. 18: 67–75.
- Stringer, C (2002). "Modern human origins: progress and prospects". Philosophical Transactions of the Royal Society of London. 357 (1420): 563–579. doi:10.1098/rstb.2001.1057.
- Stringer, C. (2014). "Why we are not all multiregionalists now". Trends in Ecology & Evolution. 29 (5): 248–251. doi:10.1016/j.tree.2014.03.001.
- "Allan Wilson: Revolutionary Evolutionist". New Zealanders Heroes.
- Wallace, D; Brown, MD; Lott, MT (1999). "Mitochondrial DNA variation in human evolution and disease". Gene. 238 (1): 211–30. doi:10.1016/S0378-1119(99)00295-4. PMID 10570998. "evidence that our species arose in Africa about 150 000 years before present (YBP), migrated out of Africa into Asia about 60 000 to 70 000 YBP and into Europe about 40 000 to 50 000 YBP, and migrated from Asia and possibly Europe to the Americas about 20 000 to 30 000 YBP."
- Adcock GJ, Dennis ES, Easteal S, Huttley GA, Jarmiin LS, Peacock WJ, Thorne A (2001). "Mitochondrial DNA sequences in ancient Australians: Implications for modern human origins". PNAS. 98 (2): 537–542. doi:10.1073/pnas.98.2.537. PMC . PMID 11209053.
- Heupink TH, Subramanian S, Wright JL, Endicott P, Carrington Westaway M, Huynen L, Parson W, Millar C, Willerslev E, Lambert DM (2016). "Ancient mtDNA sequences from the First Australians revisited". PNAS. 113: 6892–7. doi:10.1073/pnas.1521066113. PMC . PMID 27274055.
- Appenzeller, Tim (2012). "Human migrations: Eastern odyssey. Humans had spread across Asia by 50,000 years ago. Everything else about our original exodus from Africa is up for debate.". Nature. 485 (7396).
- Beyin, Amanuel (2011). "Upper Pleistocene Human Dispersals out of Africa: A Review of the Current State of the Debate". International Journal of Evolutionary Biology. 2011 (615094): 1–17. doi:10.4061/2011/615094.
- Endicott, Phillip; Gilbert, M. Thomas P.; Stringer, Chris; Lalueza-Fox, Carles; Willerslev, Eske; Hansen, Anders J.; Cooper, Alan (January 2003). "The genetic origins of the Andaman Islanders". AJHG. 72 (1): 178–84. doi:10.1086/345487. PMC . PMID 12478481.
- Finlayson, Clive (2009). The humans who went extinct: why Neanderthals died out and we survived. Oxford University Press US. ISBN 978-0-19-923918-4.
- Groucutt, Huw S.; et al. (2015). "Rethinking the dispersal of Homo sapiens out of Africa". Evolutionary Anthropology. 24 (4 - July/August): 149–164. doi:10.1002/evan.21455.
- Harding, Rosalind M.; Healy, Eugene; Ray, Amanda J.; Ellis, Nichola S.; Flanagan, Niamh; Todd, Carol; Dixon, Craig; Sajantila, Antti; Jackson, Ian J.; Birch-Machin, Mark A.; Rees, Jonathan L. (April 2000). "Evidence for variable selective pressures at MC1R". AJHG. 66 (4): 1351–61. doi:10.1086/302863. PMC . PMID 10733465.
- Hershkovitz, Israel; Marder, Ofer; Ayalon, Avner; Bar-Matthews, Miryam; Yasur, Gal; Boaretto, Elisabetta; Caracuta, Valentina; Alex, Bridget; et al. (2015). "Levantine cranium from Manot Cave (Israel) foreshadows the first European modern humans". Nature. 520: 216–9. doi:10.1038/nature14134. PMID 25629628.
- Kuhlwilm, Martin; et al. (2016). "Ancient gene flow from early modern humans into Eastern Neanderthals". Nature. 530: 429–33. doi:10.1038/nature16544. PMC . PMID 26886800.
- Liu, Hua; Prugnolle, Franck; Manica, Andrea; Balloux, François (August 2006). "A geographically explicit genetic model of worldwide human-settlement history". AJHG. 79 (2): 230–7. doi:10.1086/505436. PMC . PMID 16826514.
- Liu, Wu; Martinón-Torres, María; Cai, Yan-jun; Xing, Song; Tong, Hao-wen; Pei, Shu-wen; Sier, Mark Jan; Wu, Xiao-hong; Edwards, R. Lawrence (2015). "The earliest unequivocally modern humans in southern China". Nature. 526: 696–9. doi:10.1038/nature15696. PMID 26466566.
- Macaulay, Vincent; Hill, Catherine; Achilli, Alessandro; Rengo, Chiara; Clarke, Douglas; Meehan, William; Blackburn, James; Semino, Ornella; Scozzari, Rosaria; Cruciani, Fulvio; Taha, Adi; Shaari, Norazila Kassim; Raja, Joseph Maripa; Ismail, Patimah; Zainuddin, Zafarina; Goodwin, William; Bulbeck, David; Bandelt, Hans-Jürgen; Oppenheimer, Stephen; Torroni, Antonio; Richards, Martin (13 May 2005). "Single, Rapid Coastal Settlement of Asia Revealed by Analysis of Complete Mitochondrial Genomes". Science. 308 (5724): 1034–6. doi:10.1126/science.1109792. PMID 15890885.
- Meredith, Martin (2011). Born in Africa: The Quest for the Origins of Human Life. New York City: PublicAffairs. ISBN 1-58648-663-2.
- Posth, Cosimo; Renaud, Gabriel; Mittnik, Alissa; Drucker, Dorothée G.; et al. (2016). "Pleistocene Mitochondrial Genomes Suggest a Single Major Dispersal of Non-Africans and a Late Glacial Population Turnover in Europe". Current Biology. 26: 827–833. doi:10.1016/j.cub.2016.01.037. PMID 26853362.
- Shackelton, Laura A.; Rambaut, Andrew; Pybus, Oliver G.; Holmes, Edward C. (2006). "JC Virus Evolution and Its Association with Human Populations". Journal of Virology. 80: 9928–33. doi:10.1128/JVI.00441-06. PMC . PMID 17005670.
- Shen, Guanjun; Wang, Wei; Wang, Qian; Zhao, Jianxin; Collerson, Kenneth; Zhou, Chunlin; Tobias, Phillip V. (2002). "U-Series dating of Liujiang hominid site in Guangxi, Southern China". J. Hum. Evol. 43 (6): 817–29. doi:10.1006/jhev.2002.0601. PMID 12473485.
- Young McChesney, Kai (2015), "Teaching Diversity. The Science You Need to Know to Explain Why Race Is Not Biological", SAGE open, 5: 2158244015611712, doi:10.1177/2158244015611712
- Wells, Spencer (2003) . The Journey of Man: A Genetic Odyssey. New York: Random House Trade Paperbacks. ISBN 0-8129-7146-9.
- Stringer, Chris (2011). The Origin of Our Species. London: Allen Lane. ISBN 978-1-84614-140-9.
- Wells, Spencer (2006). Deep ancestry: inside the Genographic Project. Washington, D.C: National Geographic. ISBN 0-7922-6215-8.
- Wade, N. (2006). Before the Dawn : Recovering the Lost History of Our Ancestors. Penguin Press HC, The. ISBN 1-59420-079-3.
- Sykes, Bryan (2004). The Seven Daughters of Eve: The Science That Reveals Our Genetic Ancestry. Corgi Adult. ISBN 0-552-15218-8.