Multiregional origin of modern humans

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A graph detailing the evolution to modern humans using the Multiregional theory of human evolution. The horizontal lines represent 'multiregional evolution' gene flow between regional lineages.

Multiregional theories may differ regarding the amount of mixing argued to have occurred between the different regional groups. Carleton Coon‎ argued for limited mixing, while others have argued for extensive. Recent genetic evidence have disproven the most strict form of the recent African origin theory by finding the occurrence of limited interbreeding with, and not the total replacement of, groups such as Neanderthals and Denisovans.

The multiregional hypothesis is a scientific model that provides an explanation for the pattern of human evolution proposed by Milford H. Wolpoff[1] in 1988.[2] Multiregional origin holds that the evolution of humanity from the beginning of the Pleistocene 2.5 million years BP to the present day has been within a single, continuous human species, evolving worldwide to modern Homo sapiens.

A competing theory, the recent African origin of modern humans (also known as "Out of Africa theory"), has emerged as the near consensus view since the 1990s,[3][4] proposing that modern humans arose in Africa around 100-200,000 years ago, moving out of Africa around 50-60,000 years ago to replace existing human species such as Homo erectus and the Neanderthals.[5]

Proponents of multiregional origin point to fossil and genomic data[6] as support for their hypothesis. The gene flow and sexual reproduction between modern and ancestral human population has not been ruled out.[7][8] There is no evidence for a contribution of Neanderthal mitochondrial DNA (37 genes) [9] or Y-DNA (78 genes) to modern humans; the studies claiming to find evidence of older diversity in ancestry are on the autosomal DNA (20-25k genes) making up the vast majority of the human genome.

Regional continuity

Our upright posture may have originated in a common ancestor of humans and great apes who lived in Europe - and not in Africa, as previously thought. That’s the conclusion reached by an international research team headed by Professor Madelaine Böhme from the Senckenberg Center for Human Evolution and Palaeoenvironment at the University of Tübingen in a study published Wednesday in the journal Nature. Böhme has discovered fossils of a previously unknown primate in southern Germany. The fossils of Danuvius guggenmosi, which lived 11.62 million years ago, suggest that it was well adapted to both walking upright on two legs as well as using all four limbs while climbing. The ability to walk upright is considered a key characteristic of humans. The researchers say their analysis of the fossils show that Danuvius were able to walk on two legs nearly twelve million years ago. Up to now, the oldest evidence of an upright gait is a mere six million years old, and was found on the Mediterranean island of Crete as well as in Kenya. “The finds in southern Germany are a milestone in palaeoanthropology, because they raise fundamental questions about our previous understanding of the evolution of the great apes and humans,” says Böhme. Working with the Professor from Tübingen were researchers from Bulgaria, Germany, Canada and the United States. Ever since Darwin, the early evolution of humans and our cousins, the great apes, has been intensely debated. At the center of these debates is the question of how humans came to walk on two legs. Did bipedal humans evolve from tree-dwelling, monkey-like apes which moved on all fours? From brachiating apes similar to orangutans? Or from knuckle-walking apes like chimpanzees and gorillas? Over the last 150 years many hypotheses have been advocated, but supporting fossil evidence has so far mostly been lacking. The Danuvius guggenmosi fossils were discovered between 2015 and 2018. Working in the Hammerschmiede clay pit in the Allgäu region of Bavaria, Böhme and her team excavated more than 15,000 fossil vertebrate bones from the ancient humid and forested ecosystems that were abundant in southern Germany at that time. The new primate fossils include the remains of at least four individuals. The most complete skeleton, of a male Danuvius, has body proportions similar to modern-day bonobos. Thanks to completely preserved limb bones, vertebra, finger and toe bones, the researchers were able to reconstruct the way Danuvius moved about in its environment. “For the first time, we were able to investigate several functionally important joints, including the elbow, hip, knee and ankle, in a single fossil skeleton of this age,” Böhme says. “It was astonishing for us to realize how similar certain bones are to humans, as opposed to great apes.”[10]

The term "multiregional hypothesis" was first coined in the early 1980s by Milford H. Wolpoff and colleagues as an explanation for the apparent similarities seen in Homo erectus and Homo sapiens fossils from the same region, what they called regional continuity.[2]

Wolpoff rejected the earlier proposal by Coon of parallel evolution,[2] and proposed a theory based on clinal variation that would allow for the necessary balance between local selection and a global species. He proposed that Homo erectus, Neanderthals, Homo sapiens and other humans were a single species. This species arose in Africa two million years ago as H. erectus and then spread out over the world, developing adaptations to regional conditions.

It was proposed that for periods of time some populations became isolated, developing in a different direction, but through continuous interbreeding, replacement, genetic drift and selection, adaptations that were an advantage anywhere on earth would spread, keeping the development of the species in the same overall direction, while maintaining adaptations to regional factors. Eventually, the more unusual local varieties of the species would have disappeared in favor of modern humans, retaining some regional adaptations, but with many common features.[2]

Fossil evidence

Some proponents of the multiregional hypothesis, including Wolpoff, argue that fossil evidence is more reliable than estimates based on genetic evidence and molecular clocks, which they contend are subject to genetic drift, bottlenecks and other complicating factors.


Multiregionalists claimed that the discovery of a possible hybrid Homo sapiens X neanderthalensis fossil child at the Abrigo do Lagar Velho rock-shelter site in Portugal in 1999 further supports the multiregional hypothesis, by reflecting the admixture of diverse human populations.[11] Two other archaeologists dispute this: "the analysis by Duarte et al. of the Lagar Velho child's skeleton is a brave and imaginative interpretation, of which it is unlikely that a majority of paleoanthropologists will consider proven."[12]

In an article appearing in the Proceedings of the National Academy of Sciences[13] in 2007, Erik Trinkaus has brought together the available data, which shows that early modern humans did exhibit evidence of Neanderthal traits, saying, "When you look at all of the well dated and diagnostic early modern European fossils, there is a persistent presence of anatomical features that were present among the Neandertals but absent from the earlier African modern humans...Early modern Europeans reflect both their predominant African early modern human ancestry and a substantial degree of admixture between those early modern humans and the indigenous Neandertals."[14]

Peking man

Shang et al. see continuity in skeletal remains of archaic people from east Asia.[15]

Early modern humans

Wolpoff and colleagues published an analysis in 2001 of character traits of the skulls of early modern human fossils, which failed to reject a theory of dual ancestry from Javan Homo erectus for Australian early modern humans and Neanderthals for Central European modern humans, and which they said ruled out a replacement model.[16] A subsequent analysis comparing differences of Neanderthal skulls to those of modern humans using 3D morphometric techniques showed a large difference between the two populations, such that Harvati & al concluded that "we interpret the evidence presented here as supporting the view that Neanderthals represent an extinct human species and therefore refute the regional continuity model for Europe."[17] It has been argued that these differences are consistent with an evolving lineage, as ancestors are never identical to their descendants.[18]

New early modern human remains were unearthed in 2003 in Tianyuan Cave, Zhoukoudian. 14C dated 42-39 ky Tianyuan 1 holotype is the oldest, directly dated EMH in eastern Eurasia. Tianyuan 1 exhibits a series of typical modern, derived modern human features and few archaic traits. Some late archaic human traits include a large hamulus length, anterior to posterior dental proportions and a broad and rounded distal phalangeal tuberosityhis.[19]

The oldest European EMH remains were discovered in 2002 in cave named Peştera cu Oase near the Iron Gates in the Danubian corridor. Oase 1 holotype revealed specific traits combining a variety of archaic Homo traits, derived early modern humans, and possibly Neanderthal features. Modern human attributes place it close to European early modern humans among Late Pleistocene samples. The fossil belongs to the few findings in Europe which could be directly dated and is considered the oldest known early modern human fossil from Europe. Two laboratories independently yielded microfiltrated collagen with 14C averaging to 34,950 B.P.[20] In Europe around 40-30 ka evolved latitudinal cline between intermixed Neanderthal traits on west and EEMH on east. The human population phenotypic continuity exist in subsequent generations.[11][20]

Genetic evidence

By analysing haplotype data, Alan Templeton found support for three waves of human migration out of Africa, the first being 1.9 million years ago, and concluded that it was impossible that existing Eurasian populations had not interbred with African migrants.[21]

Studies on past population bottlenecks that can be inferred from molecular data have led multiregionalists to conclude that the recent single-origin hypothesis is untenable because there are no population size bottlenecks affecting all genes that are more recent than 2 million years ago.

  • Microcephalin D allele introgressed into the modern human gene pool points to the Neanderthal lineage as possible source and compelling evidence of admixture among the human loci.[22][23][24]
  • RRM2P4[25] (ribonucleotide reductase M2 subunit pseudogene 4) [6]. Present day human population clades A and B have 2.3 Ma time to most recent common ancestor (TMRCA).[26] The gene tree branches of RRM2P4 point to eastern Asian ancestry.[27]
  • PDHA1 (pyruvate dehydrogenase) locus on X chromosome has estimated coalescent-time depth of 1.86 Ma.[27][28][29]
  • MAPT locus 17q21.3 split into deep genetic lineages H1 and H2 . H2 lineage in European population suggest inheritance from Neanderthals [30][31][32][33][34].
  • ASAH1. Related to mental activity N-Acylsphingosine Amidohydrolase gene two V and M deep genetic lineages[35] have TMRCA 2.4±.4 Ma.[36] Linkage disequilibrium 62% and small nucleotide diversity 0.05% indicate a signature of positive Darwinian selection for the V lineage. The M lineage is attributed to ancient population structure of humans in Africa.[37]
  • X-chromosome genes DMD44, APXL, AMELX, TNFSF5 show S-N heterogeneous patterns of variation and may play role in diversity-reducing selection in non-Africans[38].
  • CMAH CMP-N-acetylneuraminic acid hydroxylase pseudogene show 2.9 Ma genetic history [39].
  • NAT2 [40] SNPs cluster predictably with linages originating in sub-Saharan Africa, Europe, and East Asia.[41] NAT1*11A 0.29 Ma suggest ancient structured population with gene tree rooted in Eurasia.[42][43].
  • ALMS1 suggest ancient and complex evolutionary history.[44]
  • Genome polymorphism: Inversion polymorphism: known 5-million-base pair (Mbp) 8p23.1, 1-Mbp on 17q21.3 and novel 1.2-Mbp on 15q24, 2.1-Mbp 15q13, 1.7-Mbp 17q12 [10] . In the sample of 8 gnomes from worldwide sample including Yuruba Kidd&al group found 4 million SNPs and 796,273 small indels (1−100 bp in size); 15 large regions of excess nucleotide variation 500 kbp to 3 Mbp. Two of variable sites are described detailed above.[45]

Proponents of the multiregional hypothesis show genetic sequences of several loci in the human genome with million year old genealogy.[46][47][48][49][50][51] Those data of deep genetic lineages are explained in the multiregional theory framework as a result of heredity from structured ancestral population.[52] The data are not interpreted in light of the RAO hypothesis postulating recent replacement where separated million years ago genetic lineages are at best unpredicted.[53][54]

Criticism of the multiregional hypothesis

A competing, rather phony theory, the recent African origin of modern humans (also known as "Out of Africa"), has emerged as the near consensus view since the 1990s,[3] proposing that modern humans arose in Africa around 100-200,000 years ago, moving out of Africa around 50-60,000 years ago to replace existing human species such as Homo erectus and the Neanderthals.

Ancient mitochondrial DNA sequence, extracted from 37,000 years old Neanderthal specimens, and mitochondrial DNA from present day humans have different sequences. However, the largest example of sequenced Neanderthal nuclear DNA comprised 1 million base pairs compared to a human nuclear genome size of roughly 3 billion base pairs. This amounts to a comparison of only 0.033% of the genomes.

Not Out Of Africa (Lefkowitz).jpg

Not Out Of Africa

The US-American classicist Mary Lefkowitz, author of Not Out Of Africa (New York 1996, paperback edition 1997), is the Andrew W. Mellon Professor in the Humanities at Wellesley College. She is the author of many books on ancient Greece and Rome, including Lives of the Greek Poets and Women in Greek Myth, as well as articles for the "Wall Street Journal" and the "New Republic". She is the coeditor of Women's Life in Greece and Rome and Black Athena Revisited.

Not Out of Africa has sparked widespread debate over the teaching of revisionist history in schools and colleges. Was Socrates black? Did Aristotle steal his ideas from the library in Alexandria? Do we owe the underlying tenets of our democratic civilizaiton to the Africans? Mary Lefkowitz explains why politically motivated histories of the ancient world are being written and shows how Afrocentrist claims blatantly contradict the historical evidence. Not Out of Africa is an important book that protects and argues for the necessity of historical truths and standards in cultural education. For this new paperback edition, Mary Lefkowitz has written an epilogue in which she responds to her critics and offers topics for further discussion. She has also added supplementary notes, a bibliography with suggestions for further reading, and a glossary of names.

When Lefkowitz first encountered Afrocentrism in 1991, she was appalled. She discovered that there were people writing books and teaching that Greek civilization had derived from, or had even been “stolen” from Egypt. They were making claims that the Ancient Egyptians were black, as were Socrates, Cleopatra, and other important cultural figures in the Ancient World. They maintained that Greece had been invaded from Africa in the Middle of the 2nd millennium, that Greek religion and mystery systems were based on Egyptian prototypes and that what was called “Greek” philosophy was in fact the secret wisdom of Egyptian lodges of a Masonic type. She also discovered that these arguments were being supported by gross errors of fact, such as the idea that Aristotle had plundered the Egyptian library at Alexandria as a basis for his own massive philosophical and scientific writings. In fact, of course, the library at Alexandria was founded by Macedonian Greeks at least 30 years after Aristotle’s death. She explained that the spreading of the myths occurred because Afrocentric literature was widely read and that it was being undiscerningly taught, not merely in a number of school districts but also in some universities. Furthermore, when she had attempted to question Afrocentric speakers on her own campus (Wellesley) she had been rudely rebuffed. Even worse, when she appealed to colleagues for help they often failed to support her. Their ostensible grounds for this reluctance was the relativist position that as all history is fiction, there was room for many different stories. Thus, for them Afrocentrist history was no less true than the classicists’ version of the roots of Greek civilization. However, Mary Lefkowitz believes that another and more significant reason why her colleagues let her down, was the fear of being labeled as racist. She comprehends the Afrocentrists as living in a sealed off intellectual ghetto, impervious to outside information, where they pay no attention to the truth of their propositions but are purely concerned with the “feel good” factor and boosting the low self-esteem of African-Americans. While she has some respect for this motive, she denies that it has any place in the writing and teaching of history which must always remain objective. Thus, she has felt obliged to stand up and be counted against what she sees as the Afrocentrist assault on the basic principles of education, respect for the facts, logical argument and open debate. Mary Lefkowitz reiterates Arthur M. Schlesinger Jr.’s charge that Afrocentrist history is purely an attempt to promote group self-esteem, whereas history should consist of: “dispassionate analysis, judgment and perspective [...]” Schlesinger (1917–2007), historian, social critic, and public intellectual, charged that ‘Afrocentrism’ isolated both African scholarship and people of African ancestry from the rest of humanity. He expressed the views of many when he stated:

“What this fellow Asante has been saying, essentially, is that Africa is the source of all good and Europe the source of all evil."

In his book The Disuniting of America: Reflections on a Multicultural Society (1991), Schlesinger Jr., a former advisor to the John F. Kennedy and other US administrations as well as winner of the Pulitzer Prize, states that a new attitude, one that celebrates difference and abandons assimilation, may replace the classic image of the melting pot in which differences are submerged in democracy. He argues that ethnic awareness has had many positive consequences to unite a nation with a "history of prejudice". However, the "cult of ethnicity," if pushed too far, may endanger the unity of society. According to Schlesinger, multiculturalists are "very often ethnocentric separatists who see little in the Western heritage other than Western crimes." Their "mood is one of divesting Americans of their sinful European inheritance and seeking redemptive infusions from non-Western cultures."

New fossils suggest human ancestors evolved in Europe, not Africa

Homo sapiens have been on earth for 200,000 years — give or take a few ten-thousand-year stretches. Much of that time is shrouded in the fog of prehistory. What we do know has been pieced together by deciphering the fossil record through the principles of evolutionary theory. Yet new discoveries contain the potential to refashion that knowledge and lead scientists to new, previously unconsidered conclusions. A set of 8-million-year-old teeth may have done just that. Researchers recently inspected the upper and lower jaw of an ancient European ape. Their conclusions suggest that humanity’s forebearers may have arisen in Europe before migrating to Africa, potentially upending a scientific consensus that has stood since Darwin’s day. As reported in New Scientist, the 8- to 9-million-year-old hominin jaw bones were found at Nikiti, northern Greece, in the ’90s. Scientists originally pegged the chompers as belonging to a member of Ouranopithecus, an genus of extinct Eurasian ape. David Begun, an anthropologist at the University of Toronto, and his team recently reexamined the jaw bones. They argue that the original identification was incorrect. Based on the fossil’s hominin-like canines and premolar roots, they identify that the ape belongs to a previously unknown proto-hominin. The researchers hypothesize that these proto-hominins were the evolutionary ancestors of another European great ape Graecopithecus, which the same team tentatively identified as an early hominin in 2017. Graecopithecus lived in south-east Europe 7.2 million years ago. If the premise is correct, these hominins would have migrated to Africa 7 million years ago, after undergoing much of their evolutionary development in Europe. Begun points out that south-east Europe was once occupied by the ancestors of animals like the giraffe and rhino, too. “It’s widely agreed that this was the found fauna of most of what we see in Africa today,” he told New Scientists. “If the antelopes and giraffes could get into Africa 7 million years ago, why not the apes?” It’s worth noting that Begun has made similar hypotheses before. Writing for the Journal of Human Evolution in 2002, Begun and Elmar Heizmann of the Natural history Museum of Stuttgart discussed a great ape fossil found in Germany that they argued could be the ancestor (broadly speaking) of all living great apes and humans. “Found in Germany 20 years ago, this specimen is about 16.5 million years old, some 1.5 million years older than similar species from East Africa,” Begun said in a statement then. “It suggests that the great ape and human lineage first appeared in Eurasia and not Africa.”[55]

See also

Further reading


  1. Wolpoff, MH; Hawks J, Caspari R (2000). "Multiregional, not multiple origins". Am J Phys Anthropol 112 (1): 129–36. doi:<129::AID-AJPA11>3.0.CO;2-K 10.1002/(SICI)1096-8644(200005)112:1<129::AID-AJPA11>3.0.CO;2-K. PMID 10766948.
  2. 2.0 2.1 2.2 2.3 Wolpoff, MH; JN Spuhler, FH Smith, J Radovcic, G Pope, DW Frayer, R Eckhardt, and G Clark (1988). "Modern human origins". Science 241 (4867): 772–4. doi:10.1126/science.3136545. PMID 3136545.
  3. 3.0 3.1 Hua Liu, et al. (2006). "A geographically explicit genetic model of worldwide human-settlement history". American Journal of Human Genetics 79 (2): 230–237. doi:10.1086/505436. PMID 16826514. “Currently available genetic and archaeological evidence is generally interpreted as supportive of a recent single origin of modern humans in East Africa. However, this is where the near consensus on human settlement history ends, and considerable uncertainty clouds any more detailed aspect of human colonization history.”
  4. Weaver, Timothy D; Charles C. Roseman (2008). "New developments in the genetic evidence for modern human origins". Evolutionary Anthropology: Issues, News, and Reviews 17 (1): 69–80. Wiley-Liss. doi:10.1002/evan.20161.
  5. Fagundes, NJ; Ray N, Beaumont M, Neuenschwander S, Salzano FM, Bonatto SL, Excoffier L. (2007). "Statistical evaluation of alternative models of human evolution". Proc Natl Acad Sci USA 104 (45): 17614–9. doi:10.1073/pnas.0708280104. PMID 17978179.
  6. 6.0 6.1 Cox, Mp; Mendez, Fl; Karafet, Tm; Pilkington, Mm; Kingan, Sb; Destro-Bisol, G; Strassmann, Bi; Hammer, Mf (Jan 2008). "Testing for archaic hominin admixture on the X chromosome: model likelihoods for the modern human RRM2P4 region from summaries of genealogical topology under the structured coalescent" (Free full text). Genetics 178 (1): 427–37. doi:10.1534/genetics.107.080432. ISSN 0016-6731. PMID 18202385.
  7. Relethford, JH (2008). "Genetic evidence and the modern human origins debate". Heredity 100 (6): 555–63. Macmillan. doi:10.1038/hdy.2008.14. PMID 18322457.
  8. Wall, JD; Hammer MF (2006). "Archaic admixture in the human genome". Curr Opin Genet Dev 16 (6): 606–10. doi:10.1016/j.gde.2006.09.006. PMID 17027252.
  9. Hodgson, JA; Disotell TR (2008). "No evidence of a Neanderthal contribution to modern human diversity.". Genome Biology 9 (2): 206. BioMed Central. doi:10.1186/gb-2008-9-2-206. PMID 18304371.
  10. New human ancestor discovered in Europe, 2019 (Archive)
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  12. Chunky Gravettian child; Ian Tattersall and Jeffrey H. Schwartz
  13. Trinkaus, E (May 2007). "European early modern humans and the fate of the Neandertals" (Free full text). Proceedings of the National Academy of Sciences of the United States of America 104 (18): 7367–72. doi:10.1073/pnas.0702214104. ISSN 0027-8424. PMID 17452632.
  14. The Emerging Fate Of The Neandertals
  15. Shang et al. (1999). "An early modern human from Tianyuan Cave, Zhoukoudian, China". Proceedings of the National Academy of Sciences 104 (16): 6573. doi:10.1073/pnas.0702169104. PMID 17416672.
  16. Wolpoff, Milford H; John Hawks, David W Frayer, Keith Hunley (2001). "Modern Human Ancestry at the Peripheries: A Test of the Replacement Theory". Science 291 (5502): 293–297. AAAS. doi:10.1126/science.291.5502.293. PMID 11209077.
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  18. Wolpoff, Milford; Bruce Mannheim, Alan Mann, John Hawks, Rachel Caspari, Karen R. Rosenberg, David W. Frayer, George W. Gill and Geoffrey Clark (2004). "Why not the Neandertals?". World Archaeology 36: 527. doi:10.1080/0043824042000303700.
  19. Shang, H; Tong, H; Zhang, S; Chen, F; Trinkaus, E (Apr 2007). "An early modern human from Tianyuan Cave, Zhoukoudian, China" (Free full text). Proceedings of the National Academy of Sciences of the United States of America 104 (16): 6573–8. doi:10.1073/pnas.0702169104. ISSN 0027-8424. PMID 17416672.
  20. 20.0 20.1 Trinkaus, E; Moldovan, O; Milota, S; Bîlgăr, A; Sarcina, L; Athreya, S; Bailey, Se; Rodrigo, R; Mircea, G; Higham, T; Ramsey, Cb; Van, Der, Plicht, J (Sep 2003). "An early modern human from the Peştera cu Oase, Romania" (Free full text). Proceedings of the National Academy of Sciences of the United States of America 100 (20): 11231–6. doi:10.1073/pnas.2035108100. ISSN 0027-8424. PMID 14504393. “"When multiple measurements are undertaken, the mean result can be determined through averaging the activity ratios. For Oase 1, this provides a weighted average activity ratio of 〈14a〉 = 1.29 ± 0.15%, resulting in a combined OxA-GrA 14C age of 34,950, +990, and –890 B.P."”
  21. Williams, Robyn (2004). "Are We Neanderthals?". The Science Show (ABC Radio). Retrieved 2009-05-30. 
  22. Evans, Pd; Mekel-Bobrov, N; Vallender, Ej; Hudson, Rr; Lahn, Bt (Nov 2006). "Evidence that the adaptive allele of the brain size gene microcephalin introgressed into Homo sapiens from an archaic Homo lineage" (Free full text). Proceedings of the National Academy of Sciences of the United States of America 103 (48): 18178–83. doi:10.1073/pnas.0606966103. ISSN 0027-8424. PMID 17090677. “"... As such, microcephalin shows by far the most compelling evidence of admixture among the human loci examined thus far. Speculation about the identity of the archaic Homo population from which the microcephalin D allele introgressed into the modern human gene pool points to the Neanderthal lineage as a potential (although by no means only) candidate. Anatomically modern humans and Neanderthals shared a long period of coexistence, from as early as 130,000 years ago in the Middle East (39) to as late as 35,000 years ago in Europe (40), consistent with the estimated introgression time of the microcephalin D allele at or sometime before (approx)37,000 years ago. Furthermore, the worldwide frequency distribution of the D allele, exceptionally high outside of Africa but low in sub-Saharan Africa (29), suggests, but does not necessitate, admixture with an archaic Eurasian population. ..."”
  23. Trinkaus, E (May 2007). "European early modern humans and the fate of the Neandertals" (Free full text). Proceedings of the National Academy of Sciences of the United States of America 104 (18): 7367–72. doi:10.1073/pnas.0702214104. ISSN 0027-8424. PMID 17452632.
  24. Evans, Pd; Gilbert, Sl; Mekel-Bobrov, N; Vallender, Ej; Anderson, Jr; Vaez-Azizi, Lm; Tishkoff, Sa; Hudson, Rr; Lahn, Bt (Sep 2005). "Microcephalin, a gene regulating brain size, continues to evolve adaptively in humans". Science (New York, N.Y.) 309 (5741): 1717–20. doi:10.1126/science.1113722. ISSN 0036-8075. PMID 16151009.
  25. sequence and gene tree for RRM2P4 haplotypes
  26. Time-scaled gene tree of the RRM2P4. TMRCA between A, B clades 2.3 Mya jpeg
  27. 27.0 27.1 Garrigan, D; Mobasher, Z; Severson, T; Wilder, Ja; Hammer, Mf (Feb 2005). "Evidence for archaic Asian ancestry on the human X chromosome" (Free full text). Molecular biology and evolution 22 (2): 189–92. doi:10.1093/molbev/msi013. ISSN 0737-4038. PMID 15483323.
  28. Rosalind M. Harding (March 16, 1999). "More on the X files". Proceedings of the National Academy of Sciences 96 (6): 2582–2584. doi:10.1073/pnas.96.6.2582. “"the pattern of diversity at the PDHA1 locus unexpected is that this extreme structure is observed in a polymorphism with an estimated total coalescent-time depth of 1.86 million years"”
  29. Harris, E. E. ;Jody Hey (1999). "[pdf X chromosome evidence for ancient human histories]". Proceedings of the National Academy of Sciences 96: 3320. doi:10.1073/pnas.96.6.3320.
  30. J. Hardy, A. Pittman, A. Myers, K. Gwinn-Hardy, H.C. Fung, R. de Silva, M. Hutton and J. Duckworth (2005). "Evidence suggesting that Homo neanderthalensis contributed the H2 MAPT haplotype to Homo sapiens". Biochemical Society Transactions 33, part 4;. “"We suggest that the H2 haplotype is derived from Homo neanderthalensis and entered H. sapiens populations during the coexistence of these species in Europe from approx. 45 000 to 18 000 years ago and that the H2 haplotype has been under selection pressure since that time, possibly because of the role of this H1 haplotype in neurodegenerative disease."..."The tau (MAPT ) locus is very unusual. Over a region of approx. 1.8 Mb, there are two haplotype clades in European populations, H1 and H2 [6,7]. In other populations, only the H1 occurs and shows a normal pattern of recombination"”
  31. Shaw-Smith, C; Pittman, Am; Willatt, L; Martin, H; Rickman, L; Gribble, S; Curley, R; Cumming, S; Dunn, C; Kalaitzopoulos, D; Porter, K; Prigmore, E; Krepischi-Santos, Ac; Varela, Mc; Koiffmann, Cp; Lees, Aj; Rosenberg, C; Firth, Hv; De, Silva, R; Carter, Np (Sep 2006). "Microdeletion encompassing MAPT at chromosome 17q21.3 is associated with developmental delay and learning disability". Nature genetics 38 (9): 1032–7. doi:10.1038/ng1858. ISSN 1061-4036. PMID 16906163.
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