Misplaced Pages

Genetic history of Egypt

Article snapshot taken from Wikipedia with creative commons attribution-sharealike license. Give it a read and then ask your questions in the chat. We can research this topic together.

This is an old revision of this page, as edited by Dbachmann (talk | contribs) at 14:34, 1 April 2013 (um what, "reconstructed DNA profile of Tutankhamun, based on a film that was made for the Discovery Channel"?). The present address (URL) is a permanent link to this revision, which may differ significantly from the current revision.

Revision as of 14:34, 1 April 2013 by Dbachmann (talk | contribs) (um what, "reconstructed DNA profile of Tutankhamun, based on a film that was made for the Discovery Channel"?)(diff) ← Previous revision | Latest revision (diff) | Newer revision → (diff) Main article: Archaeogenetics of the Near East See also: Population history of Egypt

The genetic history of the demographics of Egypt reflects Egypt's geographical location at the crossroads of several major cultural areas: the Mediterranean, the Middle East, the Sahara and East Africa.

In general, various DNA studies have found that the gene frequencies of North African populations are intermediate between those of the Near East, the Horn of Africa, southern Europe and Sub Saharan Africa, though NRY frequency distributions of the modern Egyptian population appear to be much more similar to those of the Middle East than to any sub-Saharan African population, suggesting a much larger Eurasian genetic component.

Ancient DNA

Contamination from handling and intrusion from microbes create obstacles to the recovery of Ancient DNA. Consequently most DNA studies have been carried out on modern Egyptian populations with the intent of learning about the influences of historical migrations on the population of Egypt. One successful study was performed on ancient mummies of the 12th Dynasty, by Paabo and Di Rienzo, which identified multiple lines of descent, some of which originated in sub-Saharan Africa.

Blood typing and DNA sampling on ancient Egyptian mummies is scant; however, blood typing of dynastic mummies found ABO frequencies to be most similar to modern Egyptians and some also to Northern Haratin populations. ABO blood group distribution shows that the Egyptians form a sister group to North African populations, including Berbers, Nubians and Canary Islanders.

DNA studies on King Tut and His Relatives

Swiss scientists at Zurich-based DNA genealogy centre, iGENEA, reconstructed DNA profile of Tutankhamun, based on a film that was made for the Discovery Channel, which showed that Tutankhamun has Haplogroup R1b1a2, to which more than 50% of European men and less than 1% of modern-day Egyptians belong to. However the Y-chromosome of King Tut has never been published and DNA profile displayed in Discovery Channel documentary may not belong to the Pharaoh. According to Carsten Pusch, a geneticist at Germany's University of Tübingen who was part of the team that unraveled Tut's DNA from samples taken from his mummy and mummies of his family members, iGENEA's claims are "simply impossible." R1b also shows up in parts of northern Africa, particularly some regions in Algeria, where tests have found it in 11.8% of subjects. It is also found in central Africa around Chad and Cameroon, but the Chadic-speaking area in Africa is dominated by the branch known as R1b1c (R-V88).

DNA studies on Ramesses III

A 2012 study done on the mummified remains of Ramesses III determined that the pharaoh's y-chromosome belonged to Haplogroup E1b1a (Y-DNA). The pharaoh's y-chromosome belongs to the most frequent haplogroup among Sub-Saharan African y-chromosomes.

DNA studies on modern Egyptians

Egypt has experienced several invasions during its history. However, these do not seem to account for more than about 10% overall of current Egyptians ancestry when the DNA evidence of the ancient mitochondrial DNA and modern Y chromosomes is considered. While Ivan van Sertima argue that the Egyptians were primarily Africoid before the many conquests of Egypt diluted the Africanity of the Egyptian people, other scholars such as Frank Yurco believe that Modern Egyptians are largely representative of the ancient population, and the DNA evidence appears to support this view.

In general, various DNA studies have found that the gene frequencies of modern North African populations are intermediate between those of the Horn of Africa and Eurasia, though possessing a greater genetic affinity with the populations of Eurasia than they do with Africa. The present population of the Sahara is Caucasoid in the extreme north, with a fairly gradual increase of Negroid component as one goes south. The results of these genetic studies is consistent with the historical record, which records significant bidirectional contact between Egypt and Nubia, and the Levant/Near East within the last few thousand years, but with general population continuity from the Early Dynastic Period of Egypt up to the modern day era.

Genetic analysis of modern Egyptians reveals that they have paternal lineages common to indigenous North-East African populations primarily (Algeria, Tunisia, Morocco), and to Near Eastern peoples to a lesser extent—these lineages would have spread during the Neolithic and were maintained by the predynastic period.

A study by Krings et al. (1999) on mitochondrial DNA clines along the Nile Valley found that a Eurasian cline runs from Northern Egypt to Southern Sudan and a Sub-Saharan cline from Southern Sudan to Northern Egypt. Another mtDNA study of modern Egyptians from the Gurna region near Thebes in Southern Egypt by Stevanovicth et al. 2004 revealed that Eurasian Out of Africa haplogroups represented 61.8% of the population, with the remainder being of Sub-Saharan (20.6%) and with a high frequency (17.6%) of haplogroup M1. According to the authors "This sedentary population presented similarities to the Ethiopian population by the L1 and L2 macrohaplogroup frequency (20.6%), by the West Eurasian component (defined by haplogroups H to K and T to X) and particularly by a high frequency (17.6%) of haplogroup M1... Our results suggest that the Gurna population has conserved the trace of an ancestral genetic structure from an ancestral East African population, characterized by a high M1 haplogroup frequency". The oral tradition of the Gurna people indicates that they, like most modern day Egyptians, descend from the Ancient Egyptians

Luis et al. (2004) found that the male haplogroups in a sample of 147 Egyptians were E1b1b (36.1%, predominantly E-M78), J (32.0%), G (8.8%), T(8.2%), and R (7.5%). E1b1b and its subclades are characteristic of some Afro-Asiatic speakers and are believed to have originated in either the Near East, North Africa, or the Horn of Africa. Cruciani et al. (2007) suggests that E-M78, E1b1b predominant subclade in Egypt, originated in "Northeastern Africa", which in the study refers specifically to Egypt and Libya

Other studies have shown that modern Egyptians have genetic affinities primarily with populations of Asia, North and Northeast Africa, and to a lesser extent Middle Eastern and European populations.

Some genetic studies done on modern Egyptians suggest that most do not have close relations to most Sub Saharan Africans, and other studies show that they are mostly related to other North Africans, and to a lesser extent southern European/Mediterranean and Middle Eastern populations. A 2004 mtDNA study of upper Egyptians from Gurna found a genetic ancestral heritage to modern Northeast Africans, characterized by a high M1 haplotype frequency and an L1 and L2 macrohaplogroup frequency of 20.6%. Another study links Egyptians in general with people from modern Eritrea and Ethiopia. Though there has been much debate of the origins of haplogroup M1 a recent 2007 study had concluded that M1 has West Asia origins not a Sub Saharan African origin Origin A 2003 Y chromosome study was performed by Lucotte on modern Egyptians, with haplotypes V, XI, and IV being most common. Haplotype V is common in Berbers and has a low frequency outside North Africa. Haplotypes V, XI, and IV are all predominantly North African/Horn African haplotypes, and they are far more dominant in Egyptians than in Near Eastern or European groups.

Y-Dna haplogroups

A study using the Y-chromosome of modern Egyptian males found similar results, namely that North East African haplogroups are predominant in the South but the predominant haplogroups in the North are characteristic of North African and Eurasian populations.

Population Nb A/B E(xE1b1b) E1b1b1 (M35) E1b1b1a (M78) E1b1b1b (M81) E1b1b1c (M123) F K G I J1 J2 R1a R1b Other Study
1 Egyptians 147 2.7% 2.7% 0 18.4% 8.2% 9.5% 0 7.5% 9.5% 0 19.7% 12.2% 3.4% 4.1% 2.1% Luis et al. (2004)
2 Egyptians from El-Hayez Oasis (Western Desert) 35 0 5.70% 5.7% 28.6% 28.6% 0 0 0 0 0 31.4% 0 0 0 0 Kujanová et al. (2009)
3 Egyptians from Siwa Oasis (Western Desert) 93 28.0% 6.5% 2.2% 6.5% 1.1% 2.2% 0 0 3.2% 0 7.5% 6.5% 0 28.0% 8.3% Dugoujon et al. (2009)
4 Northern Egyptians 44 2.3% 0 4.5% 27.3% 11.4% 9.1% 6.8% 2.3% 0 0 9.1% 9.1% 2.3% 9.9% 6.8% Arredi et al. (2004)
5 Southern Egyptians 29 0.0% 0 0 17.2% 6.9% 6.9% 17.2% 10.3% 0 3.4% 20.7% 3.4% 0 13.8% 0 Arredi et al. (2004)
Distribution of E1b1b1a (E-M78) and its subclades
Population N E-M78 E-M78* E-V12* E-V13 E-V22 E-V32 E-V65 Study
Southern Egyptians 79 50.6% 44.3% 1.3% 3.8% 1.3% Cruciani et al. (2007)
Egyptians from Bahari 41 41.4% 14.6% 2.4% 21.9% 2.4% Cruciani et al. (2007)
Northern Egyptians (Delta) 72 23.6% 5.6% 1.4% 13.9% 2.8% Cruciani et al. (2007)
Egyptians from Gurna Oasis 34 17.6% 5.9% 8.8% 2.9% Cruciani et al. (2007)
Egyptian from Siwa Oasis 93 6.4% 2.1% 4.3% Cruciani et al. (2007)

Autosomal DNA

In 13 January 2012, an exhaustive genetic study of North Africa's human populations was published. The researchers analyzed around 800,000 genetic markers, distributed throughout the entire genome in 125 North African individuals belonging to seven representative populations in the whole region (Saharawi, South Moroccans, North Moroccans, Algerians, Tunisians Berbers, Libyans and Egyptians) and the information obtained was compared with the information from the neighbouring populations. The results of this study show that there is a native genetic component ("Maghrebi") which defines North Africans. The study identified mainly two distinct, opposite gradients of ancestry: an east-to-west increase of this native North African ancestry and an east-to-west decrease in likely Near Eastern Arabic ancestry.

The study also reveals that the genetic composition of North Africa's human populations is very complex and is the result of five distinct ancestries : a local component (Maghrebi) dating back thirteen thousand years and the varied genetic influence of neighbouring populations on North African groups during successive migrations (European, Near Eastern, eastern and western sub-Saharan Africa). According to the authors, the people inhabiting North Africa today are not descendants of either the earliest occupants of this region fifty thousand years ago, or descendants of the most recent Neolithic populations. The data shows that the ancestors of today's North Africans were a group of populations which already lived in the region around thirteen thousand years ago. Furthermore, this local North African genetic component is very different from the one found in the populations in the south of the Sahara, which shows that the ancestors of today's North Africans were members of a subgroup of humanity who left Africa to conquer the rest of the world and who subsequently returned to the north of the continent to settle in the region. As well as this local component, North African populations were also observed to share genetic markers with all the neighbouring regions, as a result of more recent migrations, although these appear in different proportions. There is an influence from the Middle East, which becomes less marked as the distance from the Arabian Peninsula increases, similar proportions of European influence in all North African populations, and, in some populations (South Moroccans, Saharawi...), there are even individuals who present a large proportion of recent influence from the South of the Sahara in their genome.

Admixture analysis

Recent genetic analysis of North African populations have found that, despite the complex admixture genetic background, there is an autochthonous genomic component which is likely derived from "back-to-Africa" gene flow older than 12,000 years ago (ya) (i.e., prior to the Neolithic migrations). This local population substratum seems to represent a genetic discontinuity with the earliest modern human settlers of North Africa (those with the Aterian industry) given the estimated ancestry is younger than 40,000 years ago. North Morocco, Libya and Egypt carry high proportions of European and Near Eastern ancestral components, whereas Tunisian Berbers and Saharawi are those populations with highest autochthonous North African component.

Average ancestry proportions in North African populations estimated by ADMIXTURE for k = 4 different ancestries (October 2012)
Population N Maghreb Europe Near East Sub-Saharan Africa
Tunisia (Berbers) 18 93% 4% 2% 1%
Saharawi 18 55% 17% 10% 18%
Morocco North 18 44% 31% 14% 11%
Morocco South 16 44% 13% 10% 33%
Algeria 19 39% 27% 16% 18%
Libya 17 31% 28% 25% 16%
Egypt 19 19% 37% 30% 14%

The Copts

A study of Copts group in Sudan found relatively high frequencies of Sub-Saharan Haplogroup B (Y-DNA). The Sudanese Copts are converts to Egyptian Christianity and not ethnically related to Egyptian Copts. According to the study, the presence of Sub-Saharan haplogroups may also consistent with the historical record in which southern Egypt was colonized by Nilotic populations during the early state formation.

However, it is not generally accepted that Sudanese Copts are ethnically related to those of Egypt, as conversion of ethnic Nubian kings to Christianity occurred in the 6th century AD. According to tradition, a missionary sent by Byzantine empress Theodora arrived in Nobatia and started preaching the gospel about 540 AD. It is possible that the conversion process began earlier, however, under the aegis of Coptic missionaries from Egypt. The Nubian kings accepted the Monophysite Christianity already practiced in Egypt and acknowledged the spiritual authority of the Egyptian Coptic patriarch of Alexandria over the Nubian church, which in turn adopted the Coptic name for their church.

See also

References

  1. Cavalli-Sforza, History and Geography of Human Genes, The intermediacy of North Africa and to a lesser extent Europe is apparent
  2. ^ The Levant versus the Horn of Africa: Evidence for Bidirectional Corridors of Human Migrations – Luis; Rowold; Regueiro; Caeiro; Cinnioğlu; Roseman; Underhill; Cavalli-Sforza; and Herrera. – see http://www.pubmedcentral.nih.gov/articlerender.fcgi?artid=1182266
  3. ^ Cavalli-Sforza, L.L., P. Menozzi, and A. Piazza. 1994. The History and Geography of Human Genes. Princeton: Princeton University Press.
  4. ^ Bosch, E.; et al. (1997). "Population history of north Africa: evidence from classical genetic markers". Human Biology. 69 (3): 295–311. {{cite journal}}: Explicit use of et al. in: |first1= (help)
  5. Arredi B, Poloni E, Paracchini S, Zerjal T, Fathallah D, Makrelouf M, Pascali V, Novelletto A, Tyler-Smith C (2004). "A predominantly Nilo Saharan origin for Y-chromosomal DNA variation in North Africa". Am J Hum Genet. 75 (2): 338–45. doi:10.1086/423147. PMC 1216069. PMID 15202071.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  6. ^ Manni F, Leonardi P, Barakat A, Rouba H, Heyer E, Klintschar M, McElreavey K, Quintana-Murci L (2002). "Y-chromosome analysis in Egypt suggests a genetic regional continuity in Northeastern Africa". Hum Biol. 74 (5): 645–58. doi:10.1353/hub.2002.0054. PMID 12495079.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  7. Cavalli-Sforza. "Synthetic maps of Africa". The History and Geography of Human Genes. ISBN 0691087504. {{cite book}}: External link in |chapterurl= (help); Unknown parameter |chapterurl= ignored (|chapter-url= suggested) (help)The present population of the Sahara is Sudan in the extreme north, with an increase of Negroid component as one goes south
  8. Encyclopedia of the Archaeology of Ancient Egypt By Kathryn A. Bard, Steven Blake Shubert pp 278-279
  9. S.O.Y. Keita & A. J. Boyce, S. O. Y.; Boyce, A. J. (Anthony J.) (June 2009). "Genetics, Egypt, and History: Interpreting Geographical Patterns of Y Chromosome Variation". History in Africa. 32 (1): 221. doi:10.1353/hia.2005.0013. {{cite journal}}: Check date values in: |year= / |date= mismatch (help)
  10. Shomarka Keita (2005), S. O. Y. (2005). "Y-Chromosome Variation in Egypt". African Archaeological Review. 22 (2): 61. doi:10.1007/s10437-005-4189-4. ISBN ]. {{cite journal}}: Check |isbn= value: invalid character (help)CS1 maint: numeric names: authors list (link)
  11. Keita (2005). "History in the Interpretation of the Pattern of p49a,f TaqI RFLP Y-Chromosome Variation in Egypt". American Journal of Human Biology. 17 (5): 559–67. doi:10.1002/ajhb.20428. PMID 16136533.
  12. Shomarka Keita: What genetics can tell us
  13. Paabo, S., and A. Di Rienzo, A molecular approach to the study of Egyptian history. In Biological Anthropology and the Study of Ancient Egypt. V. Davies and R. Walker, eds. pp. 86-90. London: British Museum Press. 1993
  14. Borgognini Tarli S.M., Paoli G. 1982. Survey on paleoserological studies. Homo, 33: 69-89
  15. Cavalli-Sforza, L.L., P. Menozzi, and A. Piazza. 1994. The History and Geography of Human Genes. Princeton: Princeton University Press, 169-174
  16. Half of European men share King Tut's DNA. Reuters, 1 August 2011. Retrieved on 6 August 2011
  17. The Tutankhamun DNA Project, Igenea,
  18. King Tut Related to Half of European Men? Maybe Not, LiveScience, 3 August 2011
  19. Robino; Crobu, F; Di Gaetano, C; Bekada, A; Benhamamouch, S; Cerutti, N; Piazza, A; Inturri, S; Torre, C; et al. (2008). "Analysis of Y-chromosomal SNP haplogroups and STR haplotypes in an Algerian population sample". Journal International Journal of Legal Medicine. 122 (3): 251–5. doi:10.1007/s00414-007-0203-5. PMID 17909833. {{cite journal}}: Explicit use of et al. in: |author= (help)
  20. Cruciani, F; Santolamazza, P; Shen, P; Macaulay, V; Moral, P; Olckers, A; Modiano, D; Holmes, S; Destro-Bisol, G (2002). "A back migration from Asia to sub-Saharan Africa is supported by high-resolution analysis of human Y-chromosome haplotypes". American Journal of Human Genetics. 70 (5): 1197–214. doi:10.1086/340257. PMC 447595. PMID 11910562., pp. 13–14
  21. Cruciani; Trombetta, B; Sellitto, D; Massaia, A; Destro-Bisol, G; Watson, E; Beraud Colomb, E; Dugoujon, JM; Moral, P; et al. (2010). "Human Y chromosome haplogroup R-V88: a paternal genetic record of early mid Holocene trans-Saharan connections and the spread of Chadic languages". European Journal of Human Genetics. 18 (7): 800–7. doi:10.1038/ejhg.2009.231. PMC 2987365. PMID 20051990. {{cite journal}}: Explicit use of et al. in: |author= (help)
  22. Hawass (2012). "Revisiting the harem conspiracy and death of Ramesses III: anthropological, forensic, radiological, and genetic study". BMJ. doi:10.1136/bmj.e8268. {{cite journal}}: Cite journal requires |journal= (help)
  23. Egypt, Child of Africa. 1994. ISBN 1-56000-792-3. {{cite book}}: External link in |chapterurl= (help); Unknown parameter |chapterurl= ignored (|chapter-url= suggested) (help)
  24. Frank Yurco, "An Egyptological Review" in Mary R. Lefkowitz and Guy MacLean Rogers, eds. Black Athena Revisited. Chapel Hill: University of North Carolina Press, 1996. p. 62-100
  25. Cavalli-Sforza, History and Geography of Human Genes, The intermediacy of North Africa and to lesser extent East Africa between Africa and Europe is apparent
  26. Arredi B, Poloni E, Paracchini S, Zerjal T, Fathallah D, Makrelouf M, Pascali V, Novelletto A, Tyler-Smith C (2004). "A predominantly neolithic origin for Y-chromosomal DNA variation in North Africa". Am J Hum Genet. 75 (2): 338–45. doi:10.1086/423147. PMC 1216069. PMID 15202071.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  27. Cavalli-Sforza. "Synthetic maps of Africa". The History and Geography of Human Genes. ISBN 0-691-08750-4. {{cite book}}: External link in |chapterurl= (help); Unknown parameter |chapterurl= ignored (|chapter-url= suggested) (help)The present population of the Sahara is Caucasoid in the extreme north, with a fairly gradual increase of Negroid component as one goes south
  28. Krings, T; Salem, AE; Bauer, K; Geisert, H; Malek, AK; Chaix, L; Simon, C; Welsby, D; Di Rienzo, A (1992). "mtDNA Analysis of Nile River Valley Populations: Genetic Corridor or a Barrier to Migration?" (PDF). Am J Hum Genet. 64 (5): 1116–76. doi:10.1086/302314. PMC 1377841. PMID 10090902.
  29. ^ Lucotte, G.; Mercier, G. (2001). "Brief communication: Y-chromosome haplotypes in Egypt". American Journal of Physical Anthropology. 121 (1): 63–6. doi:10.1002/ajpa.10190. PMID 12687584. Cite error: The named reference "lucotte" was defined multiple times with different content (see the help page).
  30. Arredi B, Poloni E, Paracchini S, Zerjal T, Fathallah D, Makrelouf M, Pascali V, Novelletto A, Tyler-Smith C (2004). "A Predominantly Neolithic Origin for Y-Chromosomal DNA Variation in North Africa". Am J Hum Genet. 75 (2): 338–45. doi:10.1086/423147. PMC 1216069. PMID 15202071.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  31. Manni F, Leonardi P, Barakat A, Rouba H, Heyer E, Klintschar M, McElreavey K, Quintana-Murci L (2002). "Y-chromosome analysis in Egypt suggests a genetic regional continuity in Northeastern Africa". Hum Biol. 74 (5): 645–58. doi:10.1353/hub.2002.0054. PMID 12495079.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  32. Kings, T; Salem, AE; Bauer, K; Geisert, H; Malek, AK; Chaix, L; Simon, C; Welsby, D; Di Rienzo, A (1992). "mtDNA Analysis of Nile River Valley Populations: Genetic Corridor or a Barrier to Migration?" (PDF). Am J Hum Genet. 64 (5): 1116–76. doi:10.1086/302314. PMC 1377841. PMID 10090902.
  33. Stevanovitch, A.; Gilles, A.; Bouzaid, E.; Kefi, R.; Paris, F.; Gayraud, R. P.; Spadoni, J. L.; El-Chenawi, F.; Beraud-Colomb, E. (2004). "Mitochondrial DNA Sequence Diversity in a Sedentary Population from Egypt". Annals of Human Genetics. 68 (Pt 1): 23–39. doi:10.1046/j.1529-8817.2003.00057.x. PMID 14748828.
  34. Underhill (2002), Bellwood and Renfrew, ed., Inference of Neolithic Population Histories using Y-chromosome Haplotypes, Cambridge: McDonald Institute for Archaeological Research, ISBN 1-902937-20-1
  35. Kivisild T, Reidla M, Metspalu E; et al. (2004). "Ethiopian mitochondrial DNA heritage: tracking gene flow across and around the gate of tears". Am. J. Hum. Genet. 75 (5): 752–70. doi:10.1086/425161. PMC 1182106. PMID 15457403. {{cite journal}}: Explicit use of et al. in: |author= (help)CS1 maint: multiple names: authors list (link)
  36. ^ Stevanovitch A, Gilles A, Bouzaid E; et al. (2004). "Mitochondrial DNA sequence diversity in a sedentary population from Egypt". Ann. Hum. Genet. 68 (Pt 1): 23–39. doi:10.1046/j.1529-8817.2003.00057.x. PMID 14748828. {{cite journal}}: Explicit use of et al. in: |author= (help)CS1 maint: multiple names: authors list (link) Cite error: The named reference "pmid14748828" was defined multiple times with different content (see the help page).
  37. ^ Arredi B, Poloni E, Paracchini S, Zerjal T, Fathallah D, Makrelouf M, Pascali V, Novelletto A, Tyler-Smith C (2004). "A predominantly neolithic origin for Y-chromosomal DNA variation in North Africa". Am J Hum Genet. 75 (2): 338–45. doi:10.1086/423147. PMC 1216069. PMID 15202071.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  38. ^ Manni F, Leonardi P, Barakat A, Rouba H, Heyer E, Klintschar M, McElreavey K, Quintana-Murci L (2002). "Y-chromosome analysis in Egypt suggests a genetic regional continuity in Northeastern Africa". Hum Biol. 74 (5): 645–58. doi:10.1353/hub.2002.0054. PMID 12495079.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  39. Luca Cavalli-Sforza, Luigi (1996-08-05). The History and Geography of Human Genes. Princeton University Press. ISBN 978-0691029054. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  40. Cavalli-Sforza, L.L., P. Menozzi, and A. Piazza. 1994, The History and Geography of Human Genes. Princeton:Princeton University Press.
  41. Kivisild T, Reidla M, Metspalu E, Rosa A, Brehm A, Pennarun E, Parik J, Geberhiwot T, Usanga E; et al. (2004). "Ethiopian mitochondrial DNA heritage: tracking gene flow across and around the gate of tears". Am J Hum Genet. 75 (5): 752–70. {{cite journal}}: Explicit use of et al. in: |author= (help)CS1 maint: multiple names: authors list (link)
  42. Mitochondrial lineage M1 traces an early human backflow to Africa
  43. Keita, S.O. (2005). "History in the interpretation of the pattern of p49a, f TaqI RFLP Y-chromosome variation in Egypt: a consideration of multiple lines of evidence". Am J Hum Biol. 17 (5): 559–67. doi:10.1002/ajhb.20428. PMID 16136533. {{cite journal}}: |access-date= requires |url= (help)
  44. Luis JR, Rowold DJ, Regueiro M, Caeiro B, Cinnioglu C, Roseman C, Underhill PA, Cavalli-Sforza LL, Herrera RJ (2004) The Levant versus the Horn of Africa: evidence for bidirectional corridors of human migrations. Am J Hum Genet 74:532–544
  45. Martina Kujanová, Luísa Pereira, Verónica Fernandes, Joana B. Pereira, Viktor Černý, Near Eastern Neolithic genetic input in a small oasis of the Egyptian Western Desert, American Journal of Physical Anthropology, Volume 140, Issue 2, pages 336–346, October 2009
  46. Dugoujon J.M., Coudray C., Torroni A., Cruciani F., Scozzari F., Moral P., Louali N., Kossmann M. The Berber and the Berbers: Genetic and linguistic diversities
  47. Cruciani, F.; La Fratta, R.; Trombetta, B.; Santolamazza, P.; Sellitto, D.; Colomb, E. B.; Dugoujon, J.-M.; Crivellaro, F. et al. (2007), "Tracing Past Human Male Movements in Northern/Eastern Africa and Western Eurasia: New Clues from Y-Chromosomal Haplogroups E-M78 and J-M12", Molecular Biology and Evolution 24 (6): 1300–1311, DOI:10.1093/molbev/msm049, PMID 17351267
  48. Henn BM, Botigué LR, Gravel S, Wang W, Brisbin A, et al. (2012) Genomic Ancestry of North Africans Supports Back-to-Africa Migrations. PLoS Genet 8(1): e1002397. doi:10.1371/journal.pgen.1002397
  49. North African Populations Carry the Signature of Admixture with Neandertals. Sánchez-Quinto F, Botigué LR, Civit S, Arenas C, Ávila-Arcos MC, et al. (2012) North African Populations Carry the Signature of Admixture with Neandertals. PLoS ONE 7(10): e47765. doi:10.1371/journal.pone.0047765
  50. Hassan, Hisham Y.; Underhill, Peter A.; Cavalli-Sforza, Luca L.; Ibrahim, Muntaser E. (2008). "Y-Chromosome Variation Among Sudanese:Restricted Gene Flow, Concordance With Language, Geography, and History" (PDF). American Journal of Physical Anthropology. 137 (3): 316–23. doi:10.1002/ajpa.20876. PMID 18618658.
Categories: