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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.

The multiregional hypothesis is a theory of evolution to anatomically modern humans, Homo sapiens sapiens. The multiregional hypothesis holds that the evolution of humanity from the beginning of the Pleistocene 1.8 million years BP to the present day has been within a single, continuous human species, evolving worldwide from Homo erectus to modern Homo sapiens.^[1]

A competing theory of the recent African origin of modern humans (also known as "Out of Africa") has emerged as the near consensus view since the 1990s,^[2]^[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.^[4]

Proponents of the multiregional hypothesis point to fossil and genomic data^^[5] as support for their hypothesis. The gene flow, interbreeding or "admixture" between modern and ancestral human population has not been ruled out,^[6]^[7] although there is not yet any paleogenetic evidence for a contribution from Neanderthals to modern Europeans.^[8] To differentiate the current theory from earlier hypothesis some researchers prefer the term multiregional evolution.

History

Weidenreich-Coon

The multiregional hypothesis has its origin in the work of Franz Weidenreich in the 1930s, based on his examination of Peking Man. Weidenreich was an anatomist and observed numerous anatomical characteristics that he thought Peking Man had in common with modern Asians. The Weidenreich Theory stated that human races have evolved independently in the Old World from Homo erectus to Homo sapiens sapiens, while at the same time there was gene flow between the various populations. According to the Weidenreich Theory, genes that were generally adaptive (such as those for intelligence and communication) would flow relatively rapidly from one part of the world to the other, while those that were locally adaptive, would not. A vocal proponent of the Weidenreich theory was Carleton Coon.^[9]

Regional continuity

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.^[10]

Wolpoff rejected the earlier proposal by Coon of parallel evolution,^[10] 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.^[1]

Fossil evidence

Some supporters 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.

Neanderthals

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 Neandertal traits. "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," Trinkaus said. "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 Harvatti & 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]

Genetic evidence

By analysing haplotype data, Alan Templeton found support for three waves of human migration out of Africa, the first 1.9 million years ago, and concluded that it was impossible that existing Eurasian populations had not interbred with African migrants.^[19]

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.^[20]^[21]^[22]
RRM2P4 (ribonucleotide reductase M2 subunit pseudogene 4)^[23] region on X chromosome.^[24]
PDHA1 (pyruvate dehydrogenase) locus on X chromosome.^[25]
DMD44
APXL, AMELX, TNFSF5 ^[26]
ASAH1 two lineages V and M deep genetic lineages^[27] have TMRCA 2-2.8 My ^[28]^[29]
MAPT (microtubule associated protein tau) 17q21.3 split into deep genetic lineages H1 and H2 . H2 lineage in European population sugest inheritance from Neanderthals ^[30]^[31]^[32]^[33]^[34].

Proponents of the multiregional hypothesis show genetic sequences of several loci in the human genome with million year old genealogy^[35]^[36]^[37]^[38]^[39]^[40]. Those data of deep genetic lineages are explained in the multiregional theory framework as a result of heredity from structured ancestral population^[41]. 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. ^[42]^[43]

Some genetic lineages has potential to bring genetic evidence to multiregional evolution but the complexity of problem need more research.

HLA polymorphism has possible genealogy from 0.5 to 10 My and may reflect gene history before Homo lineage diverged from Pan^[44]. Many MHC HLA alleles are quite ancient: often an allele from a particular human HLA gene is more closely related to an allele found in chimpanzees than it is to another human allele from the same gene. The coevolutionary complexity is at least cubical. Anthropogenesis may be traced to multiple pathogenesis and again to genetic history of multiple hosts.^[45]

Researchers

The most prominent current proponents of the multiregional hypothesis are Milford H. Wolpoff, Wu Xinzhi, Alan G. Thorne, James Calcagno^[46], John Hawks^[47], Alan Templeton, and Erik Trinkaus.

Criticism of the multiregional hypothesis

Multiregional evolution contrasts with the "Recent African Origin" (RAO) theory. According to the latter theory, human evolution was a consequence of many cases of species replacement, as newer species replaced older ones across the human range^{[citation needed]}. Modern human origins, according to the RAO, is the most recent example of species replacement.

In 1997, testing performed on mitochondrial DNA extracted from a Neanderthal skeleton showed modern humans and Neanderthals last shared a common ancestor between 500,000 and 800,000 years ago, and furthermore that all modern humans mtDNA are more closely related to each other than to the Neanderthals mtDNA.^{[citation needed]}

Political implications

Leonard Lieberman and Fatimah Jackson have suggested that any new support for a biological concept of race will likely come from another source, namely, the study of human evolution. They therefore ask what, if any, implications current models of human evolution may have for any biological conception of race.^[48]

The major implication for race in the multiregional evolution continuity model involves the time depth of a million or more years in which race differentiation might evolve in diverse ecological regions [...]. This must be balanced against the degree of gene flow and the transregional operation of natural selection on encephalization due to development of tools and, more broadly, culture.^[49]

References

^ ^a ^b Wolpoff, MH (2000). "Multiregional, not multiple origins". Am J Phys Anthropol. 112 (1): 129–36. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ Hua Liu, et al. A Geographically Explicit Genetic Model of Worldwide Human-Settlement History. American Journal of Human Genetics, volume 79 (2006), pages 230–237, quote: 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.
^ Weaver, Timothy D (2008). "New developments in the genetic evidence for modern human origins". Evolutionary Anthropology: Issues, News, and Reviews. 17 (1). Wiley-Liss: 69–80. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ Fagundes, NJ (2007). "Statistical evaluation of alternative models of human evolution". Proc Natl Acad Sci U S A. 104 (45): 17614–9. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ Cox, MP (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". Genetics. 178 (1): 427–37. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ Relethford, JH (2008). "Genetic evidence and the modern human origins debate". Heredity. 100 (6). Macmillan: 555–63.
^ Wall, JD (2006). "Archaic admixture in the human genome". Curr Opin Genet Dev. 16 (6): 606–10. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ Hodgson, JA (2008). "No evidence of a Neanderthal contribution to modern human diversity". Genome Biology. 9 (2). BioMed Central: 206. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ The Races of Europe by Carleton Coon 1939 (Hosted by the Society for Nordish Physical Anthropology)
^ ^a ^b Wolpoff, MH (1988). "Modern Human Origins". Science. 241 (4867): 772–4. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ The early Upper Paleolithic human skeleton from the Abrigo do Lagar Velho (Portugal) and modern human emergence in Iberia ;Duarte C, 2. Maurício J, Pettitt P, Souto P, Trinkaus E, van der Plicht H, Zilhão J (1999) Proc Natl Acad Sci USA 96:7604–7609,[1]
^ Chunky Gravettian child; Ian Tattersall and Jeffrey H. Schwartz .[2]
^ http://www.pnas.org/cgi/content/abstract/104/18/7367 European early modern humans and the fate of the Neandertals
^ http://www.sciencedaily.com/releases/2007/04/070423185434.htm The Emerging Fate Of The Neandertals
^ 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. {{cite journal}}: Explicit use of et al. in: |author= (help)
^ Wolpoff, Milford H (2001). "Modern Human Ancestry at the Peripheries: A Test of the Replacement Theory". Science. 291. AAAS: 293–297. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ Harvati, Katerina (2004). "Neanderthal taxonomy reconsidered: Implications of 3D primate models of intra- and interspecific differences". PNAS. 101 (5): 1147–1152. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ Why not the Neandertals? Milford H. Wolpoff, Bruce Mannheim, Alan Mann, John Hawks, Rachel Caspari, Karen R. Rosenberg, David W. Frayer, George W. Gill and Geoffrey Clark; DOI: 10.1080/0043824042000303700; url: [3]
^ Williams, Robyn (2004). "Are We Neanderthals?". The Science Show. ABC Radio. Retrieved 2009-05-30.
^ Evidence that the adaptive allele of the brain size gene microcephalin introgressed into Homo sapiens from an archaic Homo lineage; PNAS | November 28, 2006 | vol. 103 | no. 48 | 18178-18183 ; quote: "... 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. Finally, our estimate of the separation time between D and non-D alleles (i.e., {approx}1,100,000 years with a lower-bound confidence interval of {approx}530,000 years) is largely consistent with the divergence time between modern humans and Neanderthals based on mitochondrial DNA (mtDNA) sequence difference (320,000–740,000 years; refs. 41 and 42) and with the earliest appearance of Neanderthals in the fossil record {approx}500,000 years ago (43). It would be of great interest to sequence the microcephalin locus in Neanderthals or other archaic Homo lineages, should it become technically feasible to retrieve and analyze nuclear DNA from ancient hominid remains. Our results not only provide genetic evidence in support of the possibility of admixture between modern humans and an archaic Homo lineage but also support the notion that the biological evolution of modern humans might have benefited from the contribution of adaptive alleles from our archaic relatives. In the case of microcephalin, it is all the more intriguing given the fact that the adaptive allele is associated with an important brain development gene. ..." URL:http://www.pnas.org/cgi/content/full/103/48/18178
^ http://www.pnas.org/cgi/content/full/104/18/7367
^ Microcephalin, a Gene Regulating Brain Size, Continues to Evolve Adaptively in Humans; Science 9 September 2005: Vol. 309. no. 5741, pp. 1717 - 1720 DOI: 10.1126/science.1113722; Patrick D. Evans & all; URL:http://www.sciencemag.org/cgi/content/full/309/5741/1717
^ [4]
^ 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. Murray P. Cox Genetics, Vol. 178, 427-437, January 2008, doi:10.1534/genetics.107.080432 URL: http://www.genetics.org/cgi/content/full/178/1/427
^ More on the X files. Rosalind M. Harding. PNAS Vol. 96, Issue 6, 2582-2584, March 16, 1999 http://www.pnas.org/cgi/content/full/96/6/2582; quote: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
^ Hammer, M. F., D. Garrigan, E. Wood, J. A. Wilder, Z. Mobasher, A. Bigham, J. G. Krenz, and M. W. Nachman. 2004. Heterogeneous patterns of variation among multiple human X-linked loci: the possible role of diversity-reducing selection in non-Africans. Genetics 167:1841–1853. doi:10.1534/genetics.103.025361 [5]; qoute:results indicate that a simple out-of-Africa bottleneck model is not sufficient to explain the observed patterns of sequence variation and that diversity-reducing selection acting at a subset of loci and/or a more complex neutral model must be invoked.
^ ASAH1 SL and ML region SNP DNA seguences [jpeg
^ http://www.genetics.org/cgi/content-nw/full/178/3/1505/FIG4
^ Population Genetic Analysis of the N-Acylsphingosine Amidohydrolase Gene Associated With Mental Activity in Humans Hie Lim Kim and Yoko Satta; doi:10.1534/genetics.107.083691 ; url: [6]
^ Evidence suggesting that Homo neanderthalensis contributed the H2 MAPT haplotype to Homo sapiens; J. Hardy, A. Pittman, A. Myers, K. Gwinn-Hardy, H.C. Fung, R. de Silva, M. Hutton and J. Duckworth; Biochemical Society Transactions (2005) Volume 33, part 4; qoute: 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
^ Microdeletion encompassing MAPT at chromosome 17q21.3 is associated with developmental delay and learning disability
^ Evolutionary toggling of the MAPT 17q21.31 inversion region; Michael C Zody, Zhaoshi Jiang, Hon-Chung Fung, Francesca Antonacci, LaDeana W Hillier, Maria Francesca Cardone, Tina A Graves, Jeffrey M Kidd, Ze Cheng, Amr Abouelleil, Lin Chen, John Wallis, Jarret Glasscock, Richard K Wilson, Amy Denise Reily, Jaime Duckworth, Mario Ventura, John Hardy, Wesley C Warren & Evan E Eichler; Nature Genetics 40, 1076 - 1083 (2008); doi:10.1038/ng.193 [7]
^ Introgression and microcephalin FAQ John Hawks
^ H1 tau haplotype-related genomic variation at 17q21.3 as an Asian heritage of the European Gypsy population; Heredity (2008) 101, 416–419; doi:10.1038/hdy.2008.70; published online 23 July 2008; qoute:In this study, we examine the frequency of a 900 kb inversion at 17q21.3 in the Gypsy and Caucasian populations of Hungary, which may reflect the Asian origin of Gypsy populations. Of the two haplotypes (H1 and H2), H2 is thought to be exclusively of Caucasian origin, and its occurrence in other racial groups is likely to reflect admixture. In our sample, the H1 haplotype was significantly more frequent in the Gypsy population (89.8 vs 75.5%, P<0.001) and was in Hardy–Weinberg disequilibrium (P=0.017). The 17q21.3 region includes the gene of microtubule-associated protein tau, and this result might imply higher sensitivity to H1 haplotype-related multifactorial tauopathies among Gypsies.
^ Evidence for Archaic Asian Ancestry on the Human X Chromosome; Daniel Garrigan, Zahra Mobasher, Tesa Severson, Jason A. Wilder and Michael F. Hammer; Molecular Biology and Evolution 2005 22(2):189-192; doi:10.1093/molbev/msi013 [8]
^ Deep Haplotype Divergence and Long-Range Linkage Disequilibrium at Xp21.1 Provide Evidence That Humans Descend From a Structured Ancestral Population; Daniel Garrigan, Zahra Mobasher, Sarah B. Kingan, Jason A. Wilder and Michael F. Hammer; Genetics, Vol. 170, 1849-1856, August 2005, Copyright © 2005 doi:10.1534/genetics.105.041095 [9]
^ X chromosome evidence for ancient human histories; Eugene E. Harris and Jody Hey; PNAS March 16, 1999 vol. 96 no. 6 3320-3324 [10]
^ A common inversion under selection in Europeans; Stefansson H, Helgason A, Thorleifsson G, Steinthorsdottir V, Masson G, Barnard J, Baker A, Jonasdottir A, Ingason A, Gudnadottir VG, et al. Nature Genetics 37, 129 - 137 (2005) Published online: 16 January 2005; doi:10.1038/ng1508
^ Evidence that the adaptive allele of the brain size gene microcephalin introgressed into Homo sapiens from an archaic Homo lineage; Patrick D. Evans, Nitzan Mekel-Bobrov, Eric J. Vallender, Richard R. Hudson and Bruce T. Lahn; PNAS November 28, 2006 vol. 103 no. 48 18178-18183 [11]
^ Early modern human diversity suggests subdivided population structure and a complex out-of-Africa scenario Philipp Gunza, Fred L. Booksteina, Philipp Mitteroeckera, Andrea Stadlmayra, Horst Seidlera and Gerhard W. Webera; 10.1073/pnas.0808160106 [12]
^ [13]
^ Ancient lineages in the genome: A response to Fagundes et al; Daniel Garrigan and Michael F. Hammer; doi:10.1534/genetics.105.041095 [14]
^ Reply to Garrigan and Hammer: Ancient lineages and assimilation; Nelson J. R. Fagundes, Nicolas Ray, Mark Beaumont, Samuel Neuenschwande, Francisco M. Salzano†, Sandro L. Bonatto and Laurent Excoffier ;10.1073/pnas.0711261105 [15] qoute:We must repeat that our results do not exclude the occurrence of some admixture events between modern and archaic humans,
^ Genes, fossils, and behaviour; Robert Foley, North Atlantic Treaty Organization google books preview page 101
^ Infectious Disease and Host-Pathogen Evolution; Edited by Krishna R. Dronamraju ; ISBN-13: 9780521820660 | ISBN-10: 0521820669
^ Calcagno homepage link
^ John Hawks homepage link
^ Leonard Lieberman and Fatimah Linda C. Jackson (1995) "Race and Three Models of Human Origin" in American Anthropologist Vol. 97, No. 2, pp. 232-234
^ Leonard Lieberman and Fatimah Linda C. Jackson (1995) "Race and Three Models of Human Origin" in American Anthropologist Vol. 97, No. 2, pp. 237

Reviews

Templeton, AR (2002). "Out of Africa again and again". Nature. 416: 45–51.
Pearson, Osbjorn M (2004). "Has the Combination of Genetic and Fossil Evidence Solved the Riddle of Modern Human Origins?". Evolutionary Anthropology. 13: 145–159.
Adams, J (2008). "Human Evolutionary Tree". Nature Education. 1 (1). Macmillian.
Johanson, Donald C (May 2001). "Origins of Modern Humans: Multiregional or Out of Africa?". ActionBioscience. Retrieved 2009-05-30.

External links

[16] - 'Genomics refutes an exclusively African origin of humans' (pdf) Vinayak Eswaran, Henry Harpending, Alan R. Rogers, Journal of Human Evolution (2005)
[17] - 'Templeton tree'
[18] - 'The Hybrid Child from Portugal'
Biochem. Soc. Trans (2005) 33, 582-585 - J. Hardy and others - Molecular Mechanisms of Neurodegeneration (Evidence suggesting that Homo neanderthalensis contributed the H2 MAPT haplotype to Homo sapiens)
Kent Holsinger's web site - 'Drift and migration' (only 1 migrant per generation between populations of reasonable big sizes can prevent divergence in allelic frequencies)
Genetics - 'Deep Haplotype Divergence and Long-Range Linkage Disequilibrium at Xp21.1 Provide Evidence That Humans Descend From a Structured Ancestral Population' (first genetic evidence that statistically rejects the null hypothesis that our species descends from a single, historically panmictic population), Daniel Garrigan, Zahra Mobasher, Sarah B. Kingan, Jason A. Wilder, and Michael F. Hammer, University of Arizona, Tucson, Genetics, Vol. 170, 1849-1856, August 2005
Linfield.edu - 'The Origin of Modern Humans: Multiregional and Replacement Theories', Michael Roberts, Linfield College
[19] - 'Evidence for Archaic Asian Ancestry on the Human X Chromosome' (suggests ancient RRM2P4 lineage is remnant of introgressive hybrid of anatomically modern humans from Africa and archaic populations in Eurasia), Daniel Garrigan, Zahra Mobasher, Tesa Severson, Jason A. Wilder, Michael F. Hammer, University of Arizona, Tucson, Molecular Biology and Evolution, vol 22, no 2, p 189-192 (2005)
PNAS.org - 'Mitochondrial DNA sequences in ancient Australians: Implications for modern human origins', Gregory J. Adcock, Elizabeth S. Dennis, Simon Easteal, Gavin A. Huttley, Lars S. Jermiin, W. James Peacock, Alan Thorne, Australian National University, Proceedings of the National Academy of Sciences, vol 98, no 2, p 537-542 (January 16, 2001)
StephenJayGould.org - 'Out of Africa vs. Multiregionalism', Tod Billings (December 7, 1999)
TalkOrigins.org - 'The evolution of modern humans: where are we now?' Christopher B. Stringer, General Anthropology, vol 7, no 2, p 1-5 (2001)
Selection, nuclear genetic variation, and mtDNA

[multiregional-1] Wolpoff, MH (2000). "Multiregional, not multiple origins". Am J Phys Anthropol. 112 (1): 129–36. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[2] Hua Liu, et al. A Geographically Explicit Genetic Model of Worldwide Human-Settlement History. American Journal of Human Genetics, volume 79 (2006), pages 230–237, quote: 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.

[3] Weaver, Timothy D (2008). "New developments in the genetic evidence for modern human origins". Evolutionary Anthropology: Issues, News, and Reviews. 17 (1). Wiley-Liss: 69–80. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[statistical-4] Fagundes, NJ (2007). "Statistical evaluation of alternative models of human evolution". Proc Natl Acad Sci U S A. 104 (45): 17614–9. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[admixture-5] Cox, MP (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". Genetics. 178 (1): 427–37. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[heredity-6] Relethford, JH (2008). "Genetic evidence and the modern human origins debate". Heredity. 100 (6). Macmillan: 555–63.

[http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6VS0-4M21SWT-1&_user=10&_rdoc=1&_fmt=&_orig=search&_sort=d&view=c&_acct=C000050221&_version=1&_urlVersion=0&_userid=10&md5=43bc2fdddd454c715ebfe27b4ad6a48a-7] Wall, JD (2006). "Archaic admixture in the human genome". Curr Opin Genet Dev. 16 (6): 606–10. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[noneanderthal-8] Hodgson, JA (2008). "No evidence of a Neanderthal contribution to modern human diversity". Genome Biology. 9 (2). BioMed Central: 206. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[racesofeurope-9] The Races of Europe by Carleton Coon 1939 (Hosted by the Society for Nordish Physical Anthropology)

[wolpoff-10] Wolpoff, MH (1988). "Modern Human Origins". Science. 241 (4867): 772–4. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[11] The early Upper Paleolithic human skeleton from the Abrigo do Lagar Velho (Portugal) and modern human emergence in Iberia ;Duarte C, 2. Maurício J, Pettitt P, Souto P, Trinkaus E, van der Plicht H, Zilhão J (1999) Proc Natl Acad Sci USA 96:7604–7609,[1]

[12] Chunky Gravettian child; Ian Tattersall and Jeffrey H. Schwartz .[2]

[13] ttp://www.pnas.org/cgi/content/abstract/104/18/7367 European early modern humans and the fate of the Neandertals

[14] ttp://www.sciencedaily.com/releases/2007/04/070423185434.htm 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. {{cite journal}}: Explicit use of et al. in: |author= (help)

[peripheries-16] Wolpoff, Milford H (2001). "Modern Human Ancestry at the Peripheries: A Test of the Replacement Theory". Science. 291. AAAS: 293–297. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[17] Harvati, Katerina (2004). "Neanderthal taxonomy reconsidered: Implications of 3D primate models of intra- and interspecific differences". PNAS. 101 (5): 1147–1152. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[18] Why not the Neandertals? Milford H. Wolpoff, Bruce Mannheim, Alan Mann, John Hawks, Rachel Caspari, Karen R. Rosenberg, David W. Frayer, George W. Gill and Geoffrey Clark; DOI: 10.1080/0043824042000303700; url: [3]

[19] Williams, Robyn (2004). "Are We Neanderthals?". The Science Show. ABC Radio. Retrieved 2009-05-30.

[20] Evidence that the adaptive allele of the brain size gene microcephalin introgressed into Homo sapiens from an archaic Homo lineage; PNAS | November 28, 2006 | vol. 103 | no. 48 | 18178-18183 ; quote: "... 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. Finally, our estimate of the separation time between D and non-D alleles (i.e., {approx}1,100,000 years with a lower-bound confidence interval of {approx}530,000 years) is largely consistent with the divergence time between modern humans and Neanderthals based on mitochondrial DNA (mtDNA) sequence difference (320,000–740,000 years; refs. 41 and 42) and with the earliest appearance of Neanderthals in the fossil record {approx}500,000 years ago (43). It would be of great interest to sequence the microcephalin locus in Neanderthals or other archaic Homo lineages, should it become technically feasible to retrieve and analyze nuclear DNA from ancient hominid remains. Our results not only provide genetic evidence in support of the possibility of admixture between modern humans and an archaic Homo lineage but also support the notion that the biological evolution of modern humans might have benefited from the contribution of adaptive alleles from our archaic relatives. In the case of microcephalin, it is all the more intriguing given the fact that the adaptive allele is associated with an important brain development gene. ..." URL:http://www.pnas.org/cgi/content/full/103/48/18178

[21] ttp://www.pnas.org/cgi/content/full/104/18/7367

[22] Microcephalin, a Gene Regulating Brain Size, Continues to Evolve Adaptively in Humans; Science 9 September 2005: Vol. 309. no. 5741, pp. 1717 - 1720 DOI: 10.1126/science.1113722; Patrick D. Evans & all; URL:http://www.sciencemag.org/cgi/content/full/309/5741/1717

[23] [4]

[24] 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. Murray P. Cox Genetics, Vol. 178, 427-437, January 2008, doi:10.1534/genetics.107.080432 URL: http://www.genetics.org/cgi/content/full/178/1/427

[25] More on the X files. Rosalind M. Harding. PNAS Vol. 96, Issue 6, 2582-2584, March 16, 1999 http://www.pnas.org/cgi/content/full/96/6/2582; quote: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

[26] Hammer, M. F., D. Garrigan, E. Wood, J. A. Wilder, Z. Mobasher, A. Bigham, J. G. Krenz, and M. W. Nachman. 2004. Heterogeneous patterns of variation among multiple human X-linked loci: the possible role of diversity-reducing selection in non-Africans. Genetics 167:1841–1853. doi:10.1534/genetics.103.025361 [5]; qoute:results indicate that a simple out-of-Africa bottleneck model is not sufficient to explain the observed patterns of sequence variation and that diversity-reducing selection acting at a subset of loci and/or a more complex neutral model must be invoked.

[27] ASAH1 SL and ML region SNP DNA seguences [jpeg

[28] ttp://www.genetics.org/cgi/content-nw/full/178/3/1505/FIG4

[29] Population Genetic Analysis of the N-Acylsphingosine Amidohydrolase Gene Associated With Mental Activity in Humans Hie Lim Kim and Yoko Satta; doi:10.1534/genetics.107.083691 ; url: [6]

[30] Evidence suggesting that Homo neanderthalensis contributed the H2 MAPT haplotype to Homo sapiens; J. Hardy, A. Pittman, A. Myers, K. Gwinn-Hardy, H.C. Fung, R. de Silva, M. Hutton and J. Duckworth; Biochemical Society Transactions (2005) Volume 33, part 4; qoute: 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] Microdeletion encompassing MAPT at chromosome 17q21.3 is associated with developmental delay and learning disability

[32] Evolutionary toggling of the MAPT 17q21.31 inversion region; Michael C Zody, Zhaoshi Jiang, Hon-Chung Fung, Francesca Antonacci, LaDeana W Hillier, Maria Francesca Cardone, Tina A Graves, Jeffrey M Kidd, Ze Cheng, Amr Abouelleil, Lin Chen, John Wallis, Jarret Glasscock, Richard K Wilson, Amy Denise Reily, Jaime Duckworth, Mario Ventura, John Hardy, Wesley C Warren & Evan E Eichler; Nature Genetics 40, 1076 - 1083 (2008); doi:10.1038/ng.193 [7]

[33] Introgression and microcephalin FAQ John Hawks

[34] H1 tau haplotype-related genomic variation at 17q21.3 as an Asian heritage of the European Gypsy population; Heredity (2008) 101, 416–419; doi:10.1038/hdy.2008.70; published online 23 July 2008; qoute:In this study, we examine the frequency of a 900 kb inversion at 17q21.3 in the Gypsy and Caucasian populations of Hungary, which may reflect the Asian origin of Gypsy populations. Of the two haplotypes (H1 and H2), H2 is thought to be exclusively of Caucasian origin, and its occurrence in other racial groups is likely to reflect admixture. In our sample, the H1 haplotype was significantly more frequent in the Gypsy population (89.8 vs 75.5%, P<0.001) and was in Hardy–Weinberg disequilibrium (P=0.017). The 17q21.3 region includes the gene of microtubule-associated protein tau, and this result might imply higher sensitivity to H1 haplotype-related multifactorial tauopathies among Gypsies.

[35] Evidence for Archaic Asian Ancestry on the Human X Chromosome; Daniel Garrigan, Zahra Mobasher, Tesa Severson, Jason A. Wilder and Michael F. Hammer; Molecular Biology and Evolution 2005 22(2):189-192; doi:10.1093/molbev/msi013 [8]

[36] Deep Haplotype Divergence and Long-Range Linkage Disequilibrium at Xp21.1 Provide Evidence That Humans Descend From a Structured Ancestral Population; Daniel Garrigan, Zahra Mobasher, Sarah B. Kingan, Jason A. Wilder and Michael F. Hammer; Genetics, Vol. 170, 1849-1856, August 2005, Copyright © 2005 doi:10.1534/genetics.105.041095 [9]

[37] X chromosome evidence for ancient human histories; Eugene E. Harris and Jody Hey; PNAS March 16, 1999 vol. 96 no. 6 3320-3324 [10]

[38] A common inversion under selection in Europeans; Stefansson H, Helgason A, Thorleifsson G, Steinthorsdottir V, Masson G, Barnard J, Baker A, Jonasdottir A, Ingason A, Gudnadottir VG, et al. Nature Genetics 37, 129 - 137 (2005) Published online: 16 January 2005; doi:10.1038/ng1508

[39] Evidence that the adaptive allele of the brain size gene microcephalin introgressed into Homo sapiens from an archaic Homo lineage; Patrick D. Evans, Nitzan Mekel-Bobrov, Eric J. Vallender, Richard R. Hudson and Bruce T. Lahn; PNAS November 28, 2006 vol. 103 no. 48 18178-18183 [11]

[40] Early modern human diversity suggests subdivided population structure and a complex out-of-Africa scenario Philipp Gunza, Fred L. Booksteina, Philipp Mitteroeckera, Andrea Stadlmayra, Horst Seidlera and Gerhard W. Webera; 10.1073/pnas.0808160106 [12]

[41] [13]

[42] Ancient lineages in the genome: A response to Fagundes et al; Daniel Garrigan and Michael F. Hammer; doi:10.1534/genetics.105.041095 [14]

[43] Reply to Garrigan and Hammer: Ancient lineages and assimilation; Nelson J. R. Fagundes, Nicolas Ray, Mark Beaumont, Samuel Neuenschwande, Francisco M. Salzano†, Sandro L. Bonatto and Laurent Excoffier ;10.1073/pnas.0711261105 [15] qoute:We must repeat that our results do not exclude the occurrence of some admixture events between modern and archaic humans,

[44] Genes, fossils, and behaviour; Robert Foley, North Atlantic Treaty Organization google books preview page 101

[45] Infectious Disease and Host-Pathogen Evolution; Edited by Krishna R. Dronamraju ; ISBN-13: 9780521820660 | ISBN-10: 0521820669

[46] Calcagno homepage link

[47] John Hawks homepage link

[48] Leonard Lieberman and Fatimah Linda C. Jackson (1995) "Race and Three Models of Human Origin" in American Anthropologist Vol. 97, No. 2, pp. 232-234

[49] Leonard Lieberman and Fatimah Linda C. Jackson (1995) "Race and Three Models of Human Origin" in American Anthropologist Vol. 97, No. 2, pp. 237

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@@ Line 1: / Line 1: @@
-{{Cleanup-rewrite|date=June 2009}}
+{{cleanup-rewrite|date=June 2009}}
 {{POV|date=June 2009}}
-{{Expert-subject-multiple|Evolutionary biology|Anthropology|Human Genetic History|date=June 2009}}
+{{Expert-subject-multiple|Evolutionary biology|Anthropology|Human Genetic History}}
 [[Image:Multiregionaltheory.gif|thumb|300px|right|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.]]
-The '''multiregional hypothesis''' is a theory of how [[anatomically modern humans]], ''Homo sapiens sapiens'', evolved with a worldwide distribution. The multiregional hypothesis holds that the [[Human evolution|evolution of humanity]] from the beginning of the [[Pleistocene]] 1.8 million years [[Before Present|BP]] to the present day has been within a single, continuous [[human species]], evolving worldwide from ''[[Homo erectus]]'' to modern ''Homo sapiens''.<ref name=multiregional>{{cite journal|last=Wolpoff|first=MH|coauthors=Hawks J, Caspari R|date=2000|title=Multiregional, not multiple origins|journal=Am J Phys Anthropol|volume=112|issue=1|pages=129-36|url=http://www3.interscience.wiley.com/journal/71008905/abstract}}</ref>
+The '''multiregional hypothesis''' is a theory of [[Human evolution|evolution]] to [[anatomically modern humans]], ''Homo sapiens sapiens''. The multiregional hypothesis holds that the [[Human evolution|evolution of humanity]] from the beginning of the [[Pleistocene]] 1.8 million years [[Before Present|BP]] to the present day has been within a single, continuous [[human species]], evolving worldwide from ''[[Homo erectus]]'' to modern ''Homo sapiens''.<ref name=multiregional>{{cite journal|last=Wolpoff|first=MH|coauthors=Hawks J, Caspari R|date=2000|title=Multiregional, not multiple origins|journal=Am J Phys Anthropol|volume=112|issue=1|pages=129-36|url=http://www3.interscience.wiley.com/journal/71008905/abstract}}</ref>
 A competing theory of the [[recent African origin of modern humans]] (also known as "Out of Africa") has emerged as the near consensus view since the 1990s,<ref>Hua Liu, et al. [http://dx.doi.org/10.1086/505436 A Geographically Explicit Genetic Model of Worldwide Human-Settlement History]. ''American Journal of Human Genetics'', volume 79 (2006), pages 230–237, quote: ''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.''</ref><ref>{{cite journal|last=Weaver|first=Timothy D|coauthors=Charles C. Roseman|date=2008|title=New developments in the genetic evidence for modern human origins|journal=Evolutionary Anthropology: Issues, News, and Reviews|publisher=Wiley-Liss|volume=17|issue=1|pages=69-80|url=http://www3.interscience.wiley.com/journal/117921411/abstract}}</ref> 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]].<ref name=statistical>{{cite journal|last=Fagundes|first=NJ|coauthors=Ray N, Beaumont M, Neuenschwander S, Salzano FM, Bonatto SL, Excoffier L.|date=2007|title=Statistical evaluation of alternative models of human evolution|journal=Proc Natl Acad Sci U S A|volume=104|issue=45|pages=17614-9|url=http://www.pnas.org/content/104/45/17614.long}}</ref>
@@ Line 39: / Line 39: @@
 ==Genetic evidence==
 {{Expand-section|date=May 2009}}
-{{Copyedit|section|date=June 2009}}
 By analysing haplotype data, Alan Templeton found support for three waves of human migration out of Africa, the first 1.9 million years ago, and concluded that it was impossible that existing Eurasian populations had not interbred with African migrants.<ref>{{cite news|url=http://www.abc.net.au/rn/scienceshow/stories/2004/1151858.htm|title=Are We Neanderthals?|last=Williams|first=Robyn|date=2004|work=The Science Show|publisher=ABC Radio|accessdate=2009-05-30}}</ref>
@@ Line 49: / Line 48: @@
 *PDHA1 ([[Pyruvate_dehydrogenase#Pathology|pyruvate dehydrogenase]]) locus on X chromosome.<ref>More on the X files. Rosalind M. Harding. PNAS Vol. 96, Issue 6, 2582-2584, March 16, 1999 http://www.pnas.org/cgi/content/full/96/6/2582; quote:''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''</ref>
 *DMD44
+*APXL, AMELX, TNFSF5 <ref>Hammer, M. F., D. Garrigan, E. Wood, J. A. Wilder, Z. Mobasher, A. Bigham, J. G. Krenz, and M. W. Nachman. 2004. Heterogeneous patterns of variation among multiple human X-linked loci: the possible role of diversity-reducing selection in non-Africans. Genetics 167:1841–1853. doi:10.1534/genetics.103.025361 [http://www.genetics.org/cgi/content/abstract/167/4/1841?ijkey=cb14a3724516d1a584feb8454d2c49cd72e003ee&keytype2=tf_ipsecsha]; qoute:''results indicate that a simple out-of-Africa bottleneck model is not sufficient to explain the observed patterns of sequence variation and that diversity-reducing selection acting at a subset of loci and/or a more complex neutral model must be invoked.''</ref>
+*[[ASAH1]] two lineages V and M deep genetic lineages<ref>[[ASAH1]] SL and ML region SNP DNA seguences [[http://www.genetics.org/content/vol178/issue3/images/large/1505fig3.jpeg jpeg]</ref> have [[Time to Most Recent Common Ancestor|TMRCA]] 2-2.8 My <ref>http://www.genetics.org/cgi/content-nw/full/178/3/1505/FIG4</ref><ref>Population Genetic Analysis of the N-Acylsphingosine Amidohydrolase Gene Associated With Mental Activity in Humans
+Hie Lim Kim and Yoko Satta; doi:10.1534/genetics.107.083691 ; url: [http://www.genetics.org/cgi/content/full/178/3/1505]</ref>
+*[[MAPT]] ([[Microtubule-associated protein|microtubule]] associated [[Tau protein|protein tau]]) [[chromosome 17|17]]q21.3 split into deep genetic lineages H1 and H2 . H2 lineage in European population sugest inheritance from Neanderthals <ref>Evidence suggesting that Homo neanderthalensis contributed the H2 MAPT haplotype to Homo sapiens;
+J. Hardy, A. Pittman, A. Myers, K. Gwinn-Hardy, H.C. Fung, R. de Silva, M. Hutton and J. Duckworth; Biochemical Society Transactions (2005) Volume 33, part 4; qoute: ''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</ref><ref>Microdeletion encompassing MAPT at chromosome 17q21.3 is associated with developmental delay and learning disability</ref><ref>Evolutionary toggling of the MAPT 17q21.31 inversion region; Michael C Zody, Zhaoshi Jiang, Hon-Chung Fung, Francesca Antonacci, LaDeana W Hillier, Maria Francesca Cardone, Tina A Graves, Jeffrey M Kidd, Ze Cheng, Amr Abouelleil, Lin Chen, John Wallis, Jarret Glasscock, Richard K Wilson, Amy Denise Reily, Jaime Duckworth, Mario Ventura, John Hardy, Wesley C Warren  &  Evan E Eichler; Nature Genetics 40, 1076 - 1083 (2008); doi:10.1038/ng.193 [http://www.nature.com/ng/journal/v40/n9/extref/ng.193-S1.pdf]</ref><ref>Introgression and microcephalin FAQ [http://johnhawks.net/weblog/reviews/neandertals/neandertal_dna/introgression_faq_2006.html John Hawks] </ref><ref>H1 tau haplotype-related genomic variation at 17q21.3 as an Asian heritage of the European Gypsy population; Heredity (2008) 101, 416–419; doi:10.1038/hdy.2008.70; published online 23 July 2008; qoute:''In this study, we examine the frequency of a 900 kb inversion at 17q21.3 in the Gypsy and Caucasian populations of Hungary, which may reflect the Asian origin of Gypsy populations. Of the two haplotypes (H1 and H2), H2 is thought to be exclusively of Caucasian origin, and its occurrence in other racial groups is likely to reflect admixture. In our sample, the H1 haplotype was significantly more frequent in the Gypsy population (89.8 vs 75.5%, P<0.001) and was in Hardy–Weinberg disequilibrium (P=0.017). The 17q21.3 region includes the gene of microtubule-associated protein tau, and this result might imply higher sensitivity to H1 haplotype-related multifactorial tauopathies among Gypsies.''</ref>.
 Proponents of the multiregional hypothesis show [[DNA sequencing|genetic sequences]] of several [[locus|loci]] in the [[human genome]] with million year old genealogy<ref>Evidence for Archaic Asian Ancestry on the Human X Chromosome; Daniel Garrigan, Zahra Mobasher, Tesa Severson, Jason A. Wilder and Michael F. Hammer;  Molecular Biology and Evolution 2005 22(2):189-192; doi:10.1093/molbev/msi013 [http://mbe.oxfordjournals.org/cgi/content/abstract/22/2/189?ijkey=239b0ce6c04434150a9be27400d3278dc06240ab&keytype2=tf_ipsecsha]</ref><ref>Deep Haplotype Divergence and Long-Range Linkage Disequilibrium at Xp21.1 Provide Evidence That Humans Descend From a Structured Ancestral Population; Daniel Garrigan, Zahra Mobasher, Sarah B. Kingan, Jason A. Wilder and Michael F. Hammer; Genetics, Vol. 170, 1849-1856, August 2005, Copyright © 2005
 doi:10.1534/genetics.105.041095 [http://www.genetics.org/cgi/content/abstract/170/4/1849?ijkey=d2b0021035eead07f6e001f7ebca97b8bfbf5f35&keytype2=tf_ipsecsha]</ref><ref>X chromosome evidence for ancient human histories; Eugene E. Harris and Jody Hey; PNAS  March 16, 1999   vol. 96  no. 6  3320-3324 [http://www.pnas.org/content/96/6/3320.abstract?ijkey=7004647b82dedfbf45fd48f027bf6fda72bd70b4&keytype2=tf_ipsecsha]</ref><ref>A common inversion under selection in Europeans; Stefansson H, Helgason A, Thorleifsson G, Steinthorsdottir V, Masson G, Barnard J, Baker A, Jonasdottir A, Ingason A, Gudnadottir VG, et al. Nature Genetics  37, 129 - 137 (2005) Published online: 16 January 2005; doi:10.1038/ng1508</ref><ref>Evidence that the adaptive allele of the brain size gene microcephalin introgressed into Homo sapiens from an archaic Homo lineage; Patrick D. Evans, Nitzan Mekel-Bobrov, Eric J. Vallender, Richard R. Hudson and   Bruce T. Lahn; PNAS  November 28, 2006   vol. 103  no. 48  18178-18183  [http://www.pnas.org/content/103/48/18178.full]</ref><ref>Early modern human diversity suggests subdivided population structure and a complex out-of-Africa scenario
@@ Line 55: / Line 60: @@
 Daniel Garrigan and Michael F. Hammer; doi:10.1534/genetics.105.041095 [http://www.pnas.org/content/105/2/E3.full]</ref><ref>Reply to Garrigan and Hammer: Ancient lineages and assimilation;
 Nelson J. R. Fagundes, Nicolas Ray, Mark Beaumont, Samuel Neuenschwande, Francisco M. Salzano†, Sandro L. Bonatto and Laurent Excoffier ;10.1073/pnas.0711261105 [http://www.pnas.org/content/105/2/E4.full] qoute:''We must repeat that our results do not exclude the occurrence of some admixture events between modern and archaic humans,''</ref>
+Some genetic lineages has potential to bring genetic evidence to multiregional evolution but the complexity of problem need more research.
+*[[Human leukocyte antigen|HLA]] polymorphism has possible genealogy from 0.5 to 10 My and may reflect gene history before ''Homo'' lineage diverged from ''Pan''<ref>Genes, fossils, and behaviour; Robert Foley, North Atlantic Treaty Organization [http://books.google.com/books?id=-wJxcwWQNGoC&pg=PA99&lpg=PA99&dq=HLA+polymorphisms+evolution&source=bl&ots=O4n1saqjfz&sig=595zKxgjnct8tJwHXWRjUjhxLnY&hl=en&ei=JSsySpeRHIG0NLT05ZAK&sa=X&oi=book_result&ct=result&resnum=6#PPA102,M1 google books preview page 101]</ref>. Many [[Major_histocompatibility_complex#MHC_evolution_and_allelic_diversity|MHC]] HLA alleles are quite ancient: often an allele from a particular human HLA gene is more closely related to an allele found in chimpanzees than it is to another human allele from the same gene. The [[coevolution|coevolutionary]] complexity is at least cubical. [[Anthropogenesis]] may be traced to multiple [[pathogenesis]] and again to genetic history of multiple [[Host (biology)|host]]s.<ref>Infectious Disease and Host-Pathogen Evolution; Edited by Krishna R. Dronamraju ; ISBN-13: 9780521820660 | ISBN-10: 0521820669</ref>
 ==Researchers==
@@ Line 63: / Line 71: @@
 Multiregional evolution contrasts with the "[[Recent African Origin]]" (RAO) theory. According to the latter theory, human evolution was a consequence of many cases of species replacement, as newer species replaced older ones across the human range{{Fact|really multiple Homo species replaced each others?|date=June 2009}}.  Modern [[human origins]], according to the RAO, is the most recent example of species replacement.
-In 1997, testing performed on mitochondrial DNA extracted from a Neanderthal skeleton showed modern humans and Neanderthals last shared a common ancestor between 500,000 and 800,000 years ago, and furthermore that all modern humans mtDNA are more closely related to each other than to the Neanderthals mtDNA.{{Fact|date=June 2009}}
+In 1997, testing performed on mitochondrial DNA extracted from a Neanderthal skeleton showed modern humans and Neanderthals last shared a common ancestor between 500,000 and 800,000 years ago, and furthermore that all modern humans mtDNA are more closely related to each other than to the Neanderthals mtDNA.{{fact}}
 ==Political implications==