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[[File:Kowswamp1 cropped.png|thumb|180px|right|Cast of anatomically modern human Kow Swamp 1 skull from Australia with a face to vault angle matching that of Sangiran 17.]]
[[File:Kowswamp1 cropped.png|thumb|180px|right|Cast of anatomically modern human Kow Swamp 1 skull from Australia with a face to vault angle matching that of Sangiran 17.]]


Proponents of the multiregional hypothesis see regional continuity of certain morphological traits spanning the [[Pleistocene]] in different regions across the globe as evidence against a [[Out of Africa|single replacement model]] from Africa. In general, three major regions are recognized: [[Europe]], [[China]], and [[Indonesia]] (often including [[Australia]]).<ref>Wolpoff, M. H. (1985). Human evolution at the peripheries: the pattern at the eastern edge. Hominid Evolution: past, present and future, 355-365.</ref><ref>Frayer, D. W., Wolpoff, M. H., Thorne, A. G., Smith, F. H., & Pope, G. G. (1993). "Theories of modern human origins: the paleontological test". ''American Anthropologist''. 95(1): 14-50.</ref><ref>Wolpoff, M.H., A.G. Thorne, F.H. Smith, D.W. Frayer, and G.G. Pope: Multiregional Evolution: A World-Wide Source for Modern Human Populations. In: ''Origins of Anatomically Modern Humans'', edited by M.H. Nitecki and D.V. Nitecki. Plenum Press, New York. pp. 175-199.</ref> Wolpoff cautions that the continuity in certain skeletal features in these regions should not be seen in a racial context, instead calling them "morphological clades"; defined as sets of traits that "uniquely characterise a geographic region".<ref>Wolpoff, M. H. (1989). Multiregional evolution: the fossil alternative to Eden. In: ''The human revolution: behavioural and biological perspectives on the origins of modern humans''. 1: 62-108.</ref> According to Wolpoff and Thorne (1981): "We do not regard a morphological clade as a unique lineage, nor do we believe it necessary to imply a particular
⚫
Proponents of the multiregional hypothesis see regional continuity of certain morphological traits from archaic humans to modern humans, demonstrating regional genetic continuity, even as changes in other traits occur in parallel over time across all regions, demonstrating lateral genetic exchange.<ref name=isbn1416577963.25-26>{{cite book | title=Race and Human Evolution | author = Wolpoff, Milford, and Caspari, Rachel | year=1997 | publisher=Simon & Schuster | pages = 25–26 }}</ref> For example, in 2001 Wolpoff and colleagues published an analysis of character traits of the skulls of early modern human fossils in Australia and central Europe. They concluded that the diversity of these recent humans could not "result exclusively from a single late Pleistocene dispersal", and implied dual ancestry from Javan ''Homo erectus'' for Australia and from Neanderthals for Central Europe.<ref name="Wolpoff 2001">{{cite journal | last=Wolpoff | first=Milford H. | coauthors=John Hawks, David W. Frayer, Keith Hunley | year=2001 | title=Modern Human Ancestry at the Peripheries: A Test of the Replacement Theory | journal=Science | publisher=AAAS | volume=291 | pages=293–297 | url=http://www.sciencemag.org/cgi/content/abstract/291/5502/293 | doi=10.1126/science.291.5502.293 | pmid=11209077 | issue=5502 |bibcode = 2001Sci...291..293W }}</ref><ref name=10.1073/pnas.0808160106>Analysis of the full brain case shape confirms the idea that dispersal from a single population could not explain the early modern human variability, but does not confirm ties to regional archaic humans. {{cite journal | doi=10.1073/pnas.0808160106 | author=Gunz, P., ''et. al.'' | title=Early modern human diversity suggests subdivided population structure and a complex out-of-Africa scenario | url=http://www.pnas.org/content/106/15/6094.abstract | journal=Proceedings of the National Academy of Sciences of the United States of America | pmc=2669363 | volume=106 | issue=15 | pages=6094–6098 | date=2009-04-14 | pmid=19307568 | bibcode=2009PNAS..106.6094G}}</ref>
taxonomic status for it".<ref>Thorne, A. G., & Wolpoff, M. H. (1981). Regional continuity in Australasian Pleistocene hominid evolution. American Journal of Physical Anthropology, 55(3), 337-349.</ref> Critics of Multiregionalism have pointed out that no single human trait is unique to a geographical region (i.e. confined to one population and not found in any other) but Wolpoff ''et al''. (2000) note that regional continuity only recognizes combinations of features, not traits if individually accessed, a point they elsewhere compare to the forensic identification of a human skeleton:

{{Quote|"Regional continuity... is not the claim that such features do not appear elsewhere; the genetic structure of the human species makes such a possibility unlikely to the extreme. There may be uniqueness in ''combinations'' of traits, but no single trait is likely to have been unique in a particular part of the world although it might appear to be so because of the incomplete sampling provided by the spotty human fossil record."}}

⚫
Wolpoff also stresses that regional continuity works in conjunction with genetic exchanges between populations. The long-term or regional continuity in certain morphological traits is explained by [[Alan Thorne]]'s "Centre and Edge"<ref>Thorne, A.G. 1981. The Centre and the Edge: The signifi-cance of Australian hominids to African Palaeoan-thropology. Proceedings of the 8th Pan-African Congressof Prehistory (Nairobi), pp. 180–181. Nairobi: National Museums of Kenya.</ref> population genetics model which resolves Weidenreich's paradox of "how did populations retain geographical distinctions and yet evolve together?". For example, in 2001 Wolpoff and colleagues published an analysis of character traits of the skulls of early modern human fossils in Australia and central Europe. They concluded that the diversity of these recent humans could not "result exclusively from a single late Pleistocene dispersal", and implied dual ancestry from Javan ''Homo erectus'' for Australia and from Neanderthals for Central Europe, involving interbreeding with Africans.<ref name="Wolpoff 2001">{{cite journal | last=Wolpoff | first=Milford H. | coauthors=John Hawks, David W. Frayer, Keith Hunley | year=2001 | title=Modern Human Ancestry at the Peripheries: A Test of the Replacement Theory | journal=Science | publisher=AAAS | volume=291 | pages=293–297 | url=http://www.sciencemag.org/cgi/content/abstract/291/5502/293 | doi=10.1126/science.291.5502.293 | pmid=11209077 | issue=5502 |bibcode = 2001Sci...291..293W }}</ref><ref name=10.1073/pnas.0808160106>Analysis of the full brain case shape confirms the idea that dispersal from a single population could not explain the early modern human variability, but does not confirm ties to regional archaic humans. {{cite journal | doi=10.1073/pnas.0808160106 | author=Gunz, P., ''et. al.'' | title=Early modern human diversity suggests subdivided population structure and a complex out-of-Africa scenario | url=http://www.pnas.org/content/106/15/6094.abstract | journal=Proceedings of the National Academy of Sciences of the United States of America | pmc=2669363 | volume=106 | issue=15 | pages=6094–6098 | date=2009-04-14 | pmid=19307568 | bibcode=2009PNAS..106.6094G}}</ref>


=== Southeast Asia ===
=== Southeast Asia ===

Revision as of 20:15, 7 May 2014

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. In Weidenreich's original graphic[which?][citation needed] (which is more accurate than this one), there were also diagonal lines between the populations, e.g. between African H. erectus and Archaic Asians and between Asian H. erectus and Archaic Africans. This created a "trellis" (as Wolpoff called it) or a "network" that emphasized gene flow between geographic regions and within time. It is important to remember that the populations on the chart are not discrete – i.e., they do not represent different species, but are samples within a long lineage experiencing extensive gene flow.

The multiregional hypothesis is a scientific model that provides an alternate explanation to the widely accepted, mainstream "Out of Africa" model for the pattern of human evolution. The hypothesis holds that humans first arose near the beginning of the Pleistocene two million years ago and subsequent human evolution has been within a single, continuous human species. This species encompasses archaic human forms such as Homo erectus and Neanderthals as well as modern forms, and evolved worldwide to the diverse populations of modern Homo sapiens sapiens. The theory contends that humans evolve through a combination of adaptation within various regions of the world and gene flow between those regions. Proponents of multiregional origin point to fossil and genomic data and continuity of archaeological cultures as support for their hypothesis.[1]

The term "multiregional hypothesis" was coined in the early 1980s by Milford H. Wolpoff and colleagues, who used the theory to explain regional similarities between archaic humans and modern humans in various regions, in what they called regional continuity.[2][3] Wolpoff proposed that the mechanism of clinal variation allowed for the necessary balance between both local selection and overall evolution as a global species, with Homo erectus, Neanderthals, Homo sapiens and other human forms as subspecies. This species arose in Africa two million years ago as H. erectus and then spread out over the world, developing adaptations to regional conditions. Some populations became isolated for periods of time, developing in different directions, 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. By these mechanisms, surviving local varieties of the species evolved into modern humans, retaining some regional adaptations but with many features common to all regions.[3]

History

The Multiregional hypothesis was proposed in 1984 by Milford H. Wolpoff, Alan Thorne and Xinzhi Wu.[4] Wolpoff credits Franz Weidenreich's "Polycentric" theory of human origins as a major influence, but cautions that this should not be confused with polygenism, or Carleton Coon's model that minimized gene flow.[5][6][7] According to Wolpoff, Multiregionalism was misinterpreted by William W. Howells, who confused Weidenreich's theory with a polygenic "candelabra model" in his publications spanning five decades:

"How did Multiregional evolution get stigmatized as polygeny? We believe it comes from the confusion of Weidenreich's ideas, and ultimately of our own, with Coon's. The historic reason for linking Coon's and Weidenreich's ideas came from the mischaracterizations of Weidenreich's Polycentric model as a candelabra (Howells, 1942, 1944, 1959, 1993), that made his Polycentric model appear much more similar to Coon's than it actually was."[8]

Through the influence of Howells, many other anthropologists and biologists have confused Multiregionalism with polygenism i.e. seperate or multiple origins for different populations. Alan Templeton for example notes that this confusion has led to the error that gene flow between different populations was added to the Multiregional hypothesis as a "special pleading in response to recent difficulties", despite the fact: "parallel evolution was never part of the multiregional model, much less its core, whereas gene flow was not a recent addition, but rather was present in the model from the very beginning"[9] (emphasis in original). Despite this, Multiregionalism is still confused with polygenism, or Coon's model of racial origins, which Wolpoff and his colleagues have distanced themselves.[10]

Fossil evidence

Replica of Sangiran 17 Homo erectus skull from Indonesia showing obtuse face to vault angle determined by fitting of bones at brow.
Cast of anatomically modern human Kow Swamp 1 skull from Australia with a face to vault angle matching that of Sangiran 17.

Proponents of the multiregional hypothesis see regional continuity of certain morphological traits spanning the Pleistocene in different regions across the globe as evidence against a single replacement model from Africa. In general, three major regions are recognized: Europe, China, and Indonesia (often including Australia).[11][12][13] Wolpoff cautions that the continuity in certain skeletal features in these regions should not be seen in a racial context, instead calling them "morphological clades"; defined as sets of traits that "uniquely characterise a geographic region".[14] According to Wolpoff and Thorne (1981): "We do not regard a morphological clade as a unique lineage, nor do we believe it necessary to imply a particular taxonomic status for it".[15] Critics of Multiregionalism have pointed out that no single human trait is unique to a geographical region (i.e. confined to one population and not found in any other) but Wolpoff et al. (2000) note that regional continuity only recognizes combinations of features, not traits if individually accessed, a point they elsewhere compare to the forensic identification of a human skeleton:

"Regional continuity... is not the claim that such features do not appear elsewhere; the genetic structure of the human species makes such a possibility unlikely to the extreme. There may be uniqueness in combinations of traits, but no single trait is likely to have been unique in a particular part of the world although it might appear to be so because of the incomplete sampling provided by the spotty human fossil record."

Wolpoff also stresses that regional continuity works in conjunction with genetic exchanges between populations. The long-term or regional continuity in certain morphological traits is explained by Alan Thorne's "Centre and Edge"[16] population genetics model which resolves Weidenreich's paradox of "how did populations retain geographical distinctions and yet evolve together?". For example, in 2001 Wolpoff and colleagues published an analysis of character traits of the skulls of early modern human fossils in Australia and central Europe. They concluded that the diversity of these recent humans could not "result exclusively from a single late Pleistocene dispersal", and implied dual ancestry from Javan Homo erectus for Australia and from Neanderthals for Central Europe, involving interbreeding with Africans.[17][18]

Southeast Asia

Alan Thorne held that there was regional continuity in the human fossils in southeast Asia. Wolpoff, initially skeptical, became convinced when reconstructing the Sangiran 17 Homo erectus skull from Indonesia, when he was surprised that the skull's face to vault angle matched that of the Australian modern human Kow Swamp 1 skull. Wolpoff had expected the skull to match that of the Homo erectus specimens from China like the Dali skull, but instead, the face to vault angle seemed to be retained regionally over time, even while the fossils in the two regions showed parallel increases in brain case size and parallel reductions in masticatory structures over the intervening approximately 750,000 years.[19]

Laos

Findings in Tam Pa Ling, Laos suggest that anatomically modern humans (AMH) have lived in the region roughly 46 kya. These results were found using an AMH cranium fragment (named TPL1) found in sediment in the Tam Pa Ling cave (also known as "The Cave of The Monkeys"). There is a direct U-dating of the fossil (63.3 kya)[20] and surrounding stratigraphy dating (>46kya), found using Optically stimulated luminescence and Accelerator mass spectrometry.[21]

China

Replica of Homo erectus ("Peking man") skull from China.

Franz Weidenreich, who oversaw the excavations of numerous "Peking man" Homo erectus fossils at Zhoukoudian in the early 20th century, believed the fossil record demonstrated certain unique features linking prehistoric and modern human populations in China. Many subsequent Chinese paleoanthropologists, such as Wu Xinzhi, were also disposed to favor the multiregional hypothesis for the same reason.[22]

Proponents claim that recent finds support regional human continuity in China. The Tianyuan 1 specimen unearthed in 2003 in Tianyuan Cave, Zhoukoudian, and Carbon 14 dated to 42–39 kya exhibits a series of typical modern human features such as a distinct chin. However, the skeleton also has archaic traits such as low anterior to posterior dental proportions indicating relatively large molars and certain leg bone proportions typical of archaic forms such as Neanderthals. Shang et al. (1999) conclude that this combination of modern and archaic traits "implies that a simple spread of modern humans from Africa is unlikely."[23] A jaw bone found in 2008 and dated to 110 kya may also exhibit a mixture of archaic and modern human traits.[24]

Europe

Comparison of modern human and Neanderthal skull.

Proponents of the multiregional hypothesis argue for regional continuity in Europe on the basis of skeletal anatomy, morphology and genetics of speech, and the archaeology of the middle to upper paleolithic transition, which they believe to be inconsistent with the possibility of complete replacement of the Neanderthals in Europe without interbreeding.[25]

Some detractors of the theory have argued, in contrast, that the morphological differences between Neanderthals and early and modern humans indicate that they are different species, based on skull differences more distinct than between any subspecies pairs examined except for the two subspecies of gorilla, implying limited or no interbreeding.[26][27]

Many of the multiregional claims regarding skeletal morphology in Europe center on forms with both archaic Neanderthal traits and modern traits, to provide evidence of interbreeding rather than replacement. Examples include the Lapedo child found in Portugal[28] and the Oase 1 mandible from Peştera cu Oase, Romania,[29] though the Lapedo child example is disputed by some.[30] In a 2007 paper examining numerous samples from European early modern humans, later European humans from the Gravettian period, and the earlier Neanderthal and east African populations from whom the first two populations could have descended, Erik Trinkaus identified numerous features in the later European samples which were absent from the African sample, but present in the Neanderthal sample. These features included various aspects of skull and mandible shape, tooth shape and size, and shapes and proportions of other bones. Trinkaus concluded that early modern Europeans had predominant African ancestry with a substantial degree of admixture from the Neanderthals then indigenous to Europe.[31][32]

Genetic evidence

Genetic evidence from the late 1980s on the mitochondrial genome indicated that all living people can trace their maternal line of descent to a single female living in Africa about 200,000 years ago, the so-called Mitochondrial Eve. This originally led to a hypothesis that Homo sapiens evolved in Africa, with a founder population of humans leaving Africa and eventually replacing all archaic humans then living elsewhere (without interbreeding), known as the "Out of Africa" or "Replacement Hypothesis".[33][34][35] Recent analyses of DNA taken directly from Neanderthal specimens indicates that they or their ancestors contributed to the genome of all humans outside of Africa, indicating there was some degree of interbreeding with Neanderthals before their replacement.[36] It has also been shown that Denisova hominins contributed to the DNA of Melanesians and Australians through interbreeding.[37]

Mitochondrial DNA

Human mitochondrial DNA tree. "Mitochondrial Eve" is near the top of the diagram, next to the jagged arrow pointing to "Outgroup", and her distance from any nonafrican groups indicates that living human mitochondrial lineages coalesce in Africa.

A 1987 analysis of mitochondrial DNA from 147 people from around the world indicated that their mitochondrial lineages all coalesced in a common ancestor in Africa about 200,000 years ago. The analysis suggested that this reflected the worldwide expansion of modern humans as a new species, replacing rather than mixing with local archaic humans.[38] Later analysis of mitochondrial DNA from Neanderthals and from the denisova hominin indicated that those mitochondrial strains had diverged from the living human mitochondrial line long before 200,000 years ago, consistent with lack of interbreeding between early modern and archaic humans.[39][40]

The original mitochondrial DNA results and the resulting recent African replacement theory posed a serious challenge to the multiregional hypothesis.[41] Mitochondrial DNA alone, however, could not entirely rule out interbreeding between early modern and archaic humans, since archaic human mitochondrial strains from such interbreeding could have been lost due to genetic drift or a selective sweep.[42][43]

Nuclear DNA

Initial analysis of Y chromosome DNA, which like mitochondrial DNA, is inherited from only one parent, was consistent with a recent African replacement model. However, the mitochondrial and Y chromosome data could not be explained by the same modern human expansion out of Africa; the Y chromosome expansion would have involved genetic mixing that retained regionally local mitochondrial lines. In addition, the Y chromosome data indicated a later expansion back into Africa from Asia, demonstrating that gene flow between regions was not unidirectional.[44]

An early analysis of 15 noncoding sites on the X chromosome found additional inconsistencies with the recent African replacement hypothesis. The analysis found a multimodal distribution of coalescence times to the most recent common ancestor for those sites, contrary to the predictions for recent African replacement; in particular, there were more coalescence times near 2 million years ago (mya) than expected, suggesting an ancient population split around the time humans first emerged from Africa as Homo erectus, rather than more recently as suggested by the mitochondrial data. While most of these X chromosome sites showed greater diversity in Africa, consistent with African origins, a few of the sites showed greater diversity in Asia rather than Africa. For four of the 15 gene sites that did show greater diversity in Africa, the sites' varying diversity by region could not be explained by simple expansion from Africa, as would be required by the recent African replacement hypothesis.[45]

Later analyses of X chromosome and autosomal DNA continued to find sites with deep coalescence times inconsistent with a single origin of modern humans,[46][47][48][49][50] diversity patterns inconsistent with a recent expansion from Africa,[51] or both.[52][53] For example, analyses of a region of RRM2P4 (ribonucleotide reductase M2 subunit pseudogene 4) showed a coalescence time of about 2 Mya, with a clear root in Asia,[54][55] while the MAPT locus at 17q21.31 is split into two deep genetic lineages, one of which is common in and largely confined to the present European population, suggesting inheritance from Neanderthals.[56][57][58][59] In the case of the Microcephalin D allele, evidence for rapid recent expansion indicated introgression from an archaic population.[60][61][62][63]

In a 2005 review and analysis of the genetic lineages of 25 chromosomal regions, Alan Templeton found evidence of more than 34 occurrences of gene flow between Africa and Eurasia. Of these occurrences, 19 were associated with continuous restricted gene exchange through at least 1.46 million years ago; only 5 were associated with a recent expansion from Africa to Eurasia. Three were associated with the original expansion of Homo erectus out of Africa around 2 million years ago, 7 with an intermediate expansion out of Africa at a date consistent with the expansion of Acheulean tool technology, and a few others with other gene flows such as an expansion out of Eurasia and back into Africa subsequent to the most recent expansion out of Africa. Templeton rejected a hypothesis of recent African replacement with greater than 99% certainty (p < 10−17).[64]

Ancient DNA

Main article: Archaic human admixture with modern humans

By 2006, extraction of DNA directly from some archaic human samples was becoming possible. The earliest analyses were of Neanderthal DNA, and indicated that the Neanderthal contribution to modern human genetic diversity was no more than 20%, with a most likely value of 0%.[65] By 2010, however, detailed DNA sequencing of the Neanderthal specimens from Europe indicated that the contribution was nonzero, with Neanderthals sharing 1-4% more genetic variants with living non-Africans than with living humans in sub-Saharan Africa.[66][67] In late 2010, a recently discovered non-Neanderthal archaic human, the Denisova hominin from southern Siberia, was found to share 4-6% more of its genome with living Melanesian humans than with any other living group, supporting lateral gene transfer between two regions outside of Africa.[68][69] In August 2011, human leukocyte antigen (HLA) alleles from the archaic Denisovan and Neanderthal genomes were found to show patterns in the modern human population demonstrating origins from these non-African populations; the ancestry from these archaic alleles at the HLA-A site was more than 50% for modern Europeans, 70% for Asians, and 95% for Papua New Guineans.[70] Proponents of the multiregional hypothesis believe the combination of regional continuity inside and outside of Africa and lateral gene transfer between various regions around the world supports the multiregional hypothesis. However, "Out of Africa" Theory proponents also explain this with the fact that genetic changes occur on a regional basis rather than a continental basis, and populations close to each other are likely to share certain specific regional SNPs while sharing most other genes in common.[71][72] Migration Matrix theory (A=Mt) indicates that dependent upon the potential contribution of Neanderthal ancestry, we would be able to calculate the percentage of Neanderthal mtDNA contribution to the human species. As we do not know the specific migration matrix, we are unable to input the exact data, which would answer these questions irrefutably.[73]

Recent African Origin

Main article: Recent African origin of modern humans

The primary competing scientific hypothesis is currently recent African origin of modern humans, which proposes that modern humans arose as a new species 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 with limited interbreeding: at least once with Neanderthals and once with Denisovans.[74][75][76][77][78] This differs from the multiregional hypothesis in that the multiregional model predicts interbreeding with local human populations in any such migration.[76]

See also

References

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  13. ^ Wolpoff, M.H., A.G. Thorne, F.H. Smith, D.W. Frayer, and G.G. Pope: Multiregional Evolution: A World-Wide Source for Modern Human Populations. In: Origins of Anatomically Modern Humans, edited by M.H. Nitecki and D.V. Nitecki. Plenum Press, New York. pp. 175-199.
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  15. ^ Thorne, A. G., & Wolpoff, M. H. (1981). Regional continuity in Australasian Pleistocene hominid evolution. American Journal of Physical Anthropology, 55(3), 337-349.
  16. ^ Thorne, A.G. 1981. The Centre and the Edge: The signifi-cance of Australian hominids to African Palaeoan-thropology. Proceedings of the 8th Pan-African Congressof Prehistory (Nairobi), pp. 180–181. Nairobi: National Museums of Kenya.
  17. ^ Wolpoff, Milford H. (2001). "Modern Human Ancestry at the Peripheries: A Test of the Replacement Theory". Science. 291 (5502). AAAS: 293–297. Bibcode:2001Sci...291..293W. doi:10.1126/science.291.5502.293. PMID 11209077. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  18. ^ Analysis of the full brain case shape confirms the idea that dispersal from a single population could not explain the early modern human variability, but does not confirm ties to regional archaic humans. Gunz, P.; et al. (2009-04-14). "Early modern human diversity suggests subdivided population structure and a complex out-of-Africa scenario". Proceedings of the National Academy of Sciences of the United States of America. 106 (15): 6094–6098. Bibcode:2009PNAS..106.6094G. doi:10.1073/pnas.0808160106. PMC 2669363. PMID 19307568. {{cite journal}}: Explicit use of et al. in: |author= (help)
  19. ^ Cite error: The named reference isbn1416577963.25-26 was invoked but never defined (see the help page).
  20. ^ Demeter F, Shackelford LL, Bacon A-M, Duringer P, Westaway K, Sayavongkhamdy T, Braga J, Sichanthongtip P, Khamdalavong P and Ponche J-L 2012. Anatomically modern human in Southeast Asia (Laos) by 46 ka. Proceedings of the National Academy of Sciences 109(36).
  21. ^ http://www.pnas.org/content/early/2012/12/07/1216774109.full.pdf
  22. ^ Cite error: The named reference isbn1416577963.28-29 was invoked but never defined (see the help page).
  23. ^ Shang; Tong, H.; Zhang, S.; Chen, F.; Trinkaus, E.; et al. (1999). "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. Bibcode:2007PNAS..104.6573S. doi:10.1073/pnas.0702169104. ISSN 0027-8424. PMC 1871827. PMID 17416672. {{cite journal}}: Explicit use of et al. in: |author= (help)
  24. ^ Phil McKenna, "Chinese challenge to 'out of Africa' theory", New Scientist, November 2009.
  25. ^ Wolpoff, Milford (2004). "Why not the Neandertals?". World Archaeology. 36 (4): 527. doi:10.1080/0043824042000303700. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
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Further reading

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