Archaeogenetics of the Near East

For a non-technical introduction to genetics in general, see Introduction to genetics.

The archaeogenetics of the Near East is the study of the genetics of past human populations (archaeogenetics) in the Ancient Near East using DNA from ancient remains. Researchers use Y-DNA, mtDNA and other autosomal DNAs to identify haplogroups and haplotypes in ancient populations of Egypt, Persia, Mesopotamia, Anatolia, Arabia, the Levant and other areas.

Developments in DNA sequencing in the 1970s and 1980s provided researchers with the tools needed to study human genetic variation and the genetics of human populations to discover founder populations of modern people groups and human migrations. In 2005, National Geographic launched The Genographic Project, led by 12 prominent scientists and researchers, to study and map historical human migration patterns by collecting and analyzing DNA samples from hundreds of thousands of people from around the world.

Egypt

Main article: DNA history of Egypt

Contamination from handling and intrusion from microbes create obstacles to the recovery of Ancient DNA.[1] 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.[2][3][4][5]

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,[6] though Egypt’s NRY frequency distributions 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.[7][8][8][9][10][11]

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[12] 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.[13]

In 2013, Nature announced the publication of the first genetic study utilizing next-generation sequencing to ascertain the ancestral lineage of an Ancient Egyptian individual. The research was led by Carsten Pusch of the University of Tübingen in Germany and Rabab Khairat, who released their findings in the Journal of Applied Genetics. DNA was extracted from the heads of five Egyptian mummies that were housed at the institution. All the specimens were dated between 806 BC and 124 AD, a timeframe corresponding with the late Dynastic period. The researchers observed that one of the mummified individuals likely belonged to the mtDNA haplogroup I2, a maternal clade that is believed to have originated in Western Asia.[14]

Iran (Persia)

Further information: Origin of the Azerbaijanis

The latest comparative study (2013) on the complete mitochondrial DNA diversity in Iranians has indicated that Iranian Azeris are more related to the people of Georgia, than they are to other Iranians (Persians, Armenians, etc.), while the Persians, Armenians and Qashqai on the other hand were more related to each other.[15]

It furthermore showed that overall, the complete mtDNA sequence analysis revealed an extremely high level of genetic diversity in the Iranian populations studied which is comparable to the other groups from the South Caucasus, Anatolia and Europe.[15] The same 2013 research further noted that "the results of AMOVA and MDS analyses did not associate any regional and/or linguistic group of populations in the Anatolia, Caucasus and Iran region pointing to strong genetic affinity of Indo-European speaking Persians and Turkic-speaking Qashqais, thus suggesting their origin from a common maternal ancestral gene pool.[15] The pronounced influence of the South Caucasus populations on the maternal diversity of Iranian Azeris is also evident from the MDS analysis results."[15]

The study also notes that " It is worth pointing out the position of Azeris from the Caucasus region, who despite their supposed common origin with Iranian Azeris, cluster quite separately and occupy an intermediate position between the Azeris/Georgians and Turks/Iranians grouping".[15]

The MtDNA results from the samples overall on average closely resemble those found in the neighbouring regions of the Caucasus, Anatolia, and to a lesser extent (Northern) Mesopotamia.[15]

Among the most common MtDNA lineages in the nation, namely U3b3, appears to be restricted to populations of Iran and the Caucasus, while the sub-cluster U3b1a is common in the whole Near East region.[16]

An earlier genetic research was made by Nasidze et al. (2006) on the North Iranian populations on the Gilaks and Mazandaranis, spanning the southwestern coast of the Caspian Sea, up to the border with neighbouring Azerbaijan. The Gilaks and Mazandaranis comprise 7% of the Iranian population. It has been suggested that their ancestors came from the Caucasus region, perhaps displacing an earlier group in the South Caspian.[17] Linguistic evidence supports this scenario, in that the Gilaki and Mazandarani languages (but not other Iranian languages) share certain typological features with Caucasian languages, and specifically South Caucasian languages.[17] There have been patterns analyzed of mtDNA and Y chromosome variation in the Gilaki and Mazandarani.

Based on mtDNA HV1 sequences tested by Nasidze et al., the Gilaks and Mazandarani most closely resemble their geographic and linguistic neighbors, namely other Iranian groups. However, their Y chromosome types most closely resemble those found in groups from the South Caucasus.[17] A scenario that explains these differences is a south Caucasian origin for the ancestors of the Gilani and Mazandarani, followed by introgression of women (but not men) from local Iranian groups, possibly because of patrilocality.[17] Given that both mtDNA and language are maternally transmitted, the incorporation of local Iranian women would have resulted in the concomitant replacement of the ancestral Caucasian language and mtDNA types of the Gilani and Mazandarani with their current Iranian language and mtDNA types. Concomitant replacement of language and mtDNA may be a more general phenomenon than previously recognized.

The Mazandarani and Gilani groups fall inside a major cluster consisting of populations from the Caucasus and West Asia and are particularly close to the South Caucasus groups—Georgians, Armenians, and Azerbaijanis. Iranians from Tehran and Isfahan are situated more distantly from these groups.[17]

Iraq (Mesopotamia)

The indigenous Pre-Arab and Pre-Islamic Assyrian (aka Chaldo-Assyrian) Christian population, descendants of the ancient Mesopotamians, are fairly closely related to other Iraqis,[18][19] and also to some modern Jordanians, Mandeans and Near Eastern Jewish populations, yet due to religious, ethnic and cultural endogamy have a very distinct genetic profile that distinguishes their population from Iraqi Arabs, Kurds, Armenians, Turcomans, and Shabakis.[20] "The Assyrians are a fairly homogeneous group of people, believed to originate from the land of old Assyria in northern Iraq [..] they are Christians and are bona fide descendants of their ancient namesakes."[21]

In a 2006 study of the Y chromosome DNA of six regional populations, including, for comparison, Assyrians and Syrians, researchers found that, "the two Semitic populations (Assyrians and Syrians) are very distinct from each other according to both [comparative] axes. This difference supported also by other methods of comparison points out the weak genetic affinity between the two populations with different historical destinies." [22]

A 2008 study on the genetics of "old ethnic groups in Mesopotamia," including 340 subjects from seven ethnic communities ("Assyrian, Jewish, Zoroastrian, Armenian, Turkmen, the Arab peoples in Iran, Iraq, and Kuwait") found that Assyrians were homogeneous with respect to all other ethnic groups sampled in the study, regardless of religious affiliation.[23]

A study published in 2011 looking at the relationship between Iraq's Marsh Arabs and ancient Sumerians concluded "the modern Marsh Arabs of Iraq harbour mtDNAs and Y chromosomes that are predominantly of Middle Eastern origin. Therefore, certain cultural features of the area such as water buffalo breeding and rice farming, which were most likely introduced from the Indian sub-continent, only marginally affected the gene pool of the autochthonous people of the region. Moreover, a Middle Eastern ancestral origin of the modern population of the marshes of southern Iraq implies that, if the Marsh Arabs are descendants of the ancient Sumerians, also Sumerians were not of Indian or Southern Asian ancestry."[18]

In a 2011 study focusing on the genetics of the Maʻdān people of Iraq, researchers identified Y chromosome haplotypes shared by Marsh Arabs, Arabic speaking Iraqis, Assyrians and Mandeans "supporting a common local background."[18]

Levant (Israel, Syria, Palestine, Lebanon, Jordan)

Further information: Phoenicia, Phoenicianism, Canaanites, Jews, and Samaritans

Zalloua and Wells (2004), under the auspices of a grant from National Geographic Magazine, examined the origins of the Canaanite Phoenicians. The debate between Wells and Zalloua was whether haplogroup J2 (M172) should be identified as that of the Phoenicians or that of its "parent" haplogroup M89 on the YDNA phylogenetic tree.[24] Initial consensus suggested that J2 be identified with the Canaanite-Phoenician (Northwest Semitic) population, with avenues open for future research.[25] As Wells commented, "The Phoenicians were the Canaanites"[26] It was reported in the PBS description of the National Geographic TV Special on this study entitled "Quest for the Phoenicians" that ancient DNA was included in this study as extracted from the tooth of a 2500-year-old Phoenician mummy.[27]

Wells identified the haplogroup of the Canaanites as haplogroup J2 which originated in northern Mesopotamia.[28] The National Geographic Genographic Project linked haplogroup J2 to the site of Jericho, Tel el-Sultan, ca. 8500 BCE and indicated that in modern populations, haplogroup J2 is found primarily in the Middle East, but also along the coasts of North Africa and Southern Europe, with especially high distribution among present-day Jewish populations (30%), Southern Italians (20%), and lower frequencies in Southern Spain (10%).[29]

In a 2005 study of ASPM gene variants, Mekel-Bobrov et al. found that the Israeli Druze people of the Carmel region have among the highest rate of the newly evolved ASPM haplogroup D, at 52.2% occurrence of the approximately 6,000-year-old allele.[30] While it is not yet known exactly what selective advantage is provided by this gene variant, the haplogroup D allele is thought to be positively selected in populations and to confer some substantial advantage that has caused its frequency to rapidly increase. According to DNA testing, Druze are remarkable for the high frequency (35%) of males who carry the Y-chromosomal haplogroup L, which is otherwise uncommon in the Mideast (Shen et al. 2004).[31] This haplogroup originates from prehistoric South Asia and has spread from Pakistan into southern Iran.

Cruciani in 2007 found E1b1b1a2 (E-V13) [one from Sub Clades of E1b1b1a1 (E-V12)] in high levels (>10% of the male population) in Turkish Cypriot and Druze Arab lineages. Recent genetic clustering analyses of ethnic groups are consistent with the close ancestral relationship between the Druze and Cypriots, and also identified similarity to the general Syrian and Lebanese populations, as well as a variety of Jewish lineages (Ashkenazi, Sephardi, Iraqi Jewish, and Moroccan Jews) (Behar et al. 2010).[32]

A study published by the National Academy of Sciences found that "the paternal gene pools of Jewish communities from Europe, North Africa, and the Middle East descended from a common Middle Eastern ancestral population", and suggested that "most Jewish communities have remained relatively isolated from neighbouring non-Jewish communities during and after the Diaspora".[33] Researchers expressed surprise at the remarkable genetic uniformity they found among modern Jews, no matter where the diaspora has become dispersed around the world.[33] Skorecki and colleague wrote that "the extremely close affinity of Jewish and non-Jewish Middle Eastern populations observed ... supports the hypothesis of a common Middle Eastern origin".[34]

This research has suggested that, in addition to Israelite male, significant female founder ancestry might also derive from the Middle East-with 40% of Ashkenazim descended from four women lived about 2000–3000 years ago in the Middle East.[35] In addition, Behar (2006) suggested that the rest of Ashkenazi mtDNA is originated from about 150 women, most of those were probably of Middle Eastern origin.[36] However, more recent research suggests that the four founding maternal lineages of Ashkenazi Jews originate in Europe and that only ~8% of Ashkenazi mtDNA can confidently be assigned a Near Eastern origin, while >80% of Ashkenazi maternal lineages have a likely European origin [37]

In 2004, a team of geneticists from Stanford University, the Hebrew University of Jerusalem, Tartu University (Estonia), Barzilai Medical Center (Ashkelon, Israel), and the Assaf Harofeh Medical Center (Zerifin, Israel), studied the modern Samaritan ethnic community living in Israel in comparison with modern Israeli populations to explore the ancient genetic history of these people groups. The Samaritans or Shomronim (singular: Shomroni; Hebrew: שומרוני) trace their origins to the Assyrian province of Shomron (Samaria) in ancient Israel in the period after the Assyrian conquest circa 722 BCE. Shomron was the capital of the Northern Kingdom of Israel when it was conquered by the Assyrians and gave the name to the ancient province of Samaria and the Samaritan people group. Tradition holds that the Samaritans were a mixed group of Israelites who were not exiled or were sent back or returned from exile and non-Israelites relocated to the region by the Assyrians. The modern-day Samaritans are believed to be the direct descendants of the ancient Samaritans.

Their findings reported on four family lineages among the Samaritans: the Tsdaka family (tradition: tribe of Menasseh), the Joshua-Marhiv and Danfi families (tradition: tribe of Ephraim), and the Cohen family (tradition: tribe of Levi). All Samaritan families were found in haplogroups J1 and J2, except the Cohen family which was found in haplogroup E3b1a-M78.[31] This article predated the E3b1a subclades based on the research of Cruciani, et al.[38]

Turkey (Anatolia)

See also: Khazar theory

In population genetics the question has been debated whether the modern Turkish population is significantly related to other, mainly Central Asian, Turkic peoples, or whether they are rather derived from the indigenous largely Indo-European and Semitic speaking populations of Anatolia which were, with the exception of Greeks, Armenians, Assyrians, Jews and Kurds, culturally assimilated during the late Middle Ages. The contribution of the Central Asian genetics to the modern Turkish people has been debated and become the subject of several studies. As a result, several studies have concluded that the historical (pre-Islamic) and indigenous Anatolian groups are the primary source of the present-day Turkish population,[39][40][41][42][43][44][45] in addition to neighboring peoples,[41] such as Balkan peoples (such as Phrygians and Macedonian Greeks),[46] and Central Asian Turkic people, from the Turkic homelands in modern Kazakhstan, Uzbekistan, Turkmenistan and Kirghizstan.[41]

The Pre-Islamic and Pre-Turkic speaking population of Anatolia consisted of, at different periods, a vast patchwork of Caucasoid peoples, including Language Isolate speakers such as Hurrians, Hattians and Urartians among others, who were eventually absorbed by Indo-European populations, the most prominent being Greeks, Kaskians, Persians, Luwians, Hittites, Mitanni, Phrygians, Lydians, Cimmerians, Scythians, Medes, Lycians, Cilicians, Armenians, Celts and Kurds. Semitic speaking peoples such as Assyrians, Amorites, Eblaites, Phoenicians, Arameans and Jews also maintained a longstanding presence across southern Anatolia, and to a lesser degree, Kartvelian (Georgian) and Northwest Caucasian speaking peoples in the north east.

See also

References

  1. ↑ Encyclopedia of the Archaeology of Ancient Egypt By Kathryn A. Bard, Steven Blake Shubert. Books.google.com. pp. 278–279. Retrieved 2014-06-30.
  2. ↑ Keita, S. O. Y.; Boyce, A. J. (2005). "Genetics, Egypt, and History: Interpreting Geographical Patterns of Y Chromosome Variation". History in Africa. 32: 221–46. doi:10.1353/hia.2005.0013. JSTOR 20065742.
  3. ↑ Keita, S. O. Y. (2005). "Explanation of the Pattern of P49a,f TaqI RFLP Y-Chromosome Variation in Egypt". African Archaeological Review. 22 (2): 61–75. doi:10.1007/s10437-005-4189-4. JSTOR 25130819.
  4. ↑ Keita SO (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". American Journal of Human Biology. 17 (5): 559–67. doi:10.1002/ajhb.20428. PMID 16136533.
  5. ↑ "Shomarka Keita: What genetics can tell us". Ngm.nationalgeographic.com. Retrieved 2014-06-30.
  6. ↑ Cavalli-Sforza, History and Geography of Human Genes, The intermediacy of North Africa and to a lesser extent Europe is apparent
  7. ↑ Luis, J; Rowold, D; Regueiro, M; Caeiro, B; Cinnioglu, C; Roseman, C; Underhill, P; Cavallisforza, L; Herrera, R (2004). "The Levant versus the Horn of Africa: Evidence for Bidirectional Corridors of Human Migrations". The American Journal of Human Genetics. 74 (3): 532–44. doi:10.1086/382286. PMC 1182266Freely accessible. PMID 14973781.
  8. 1 2 Cavalli-Sforza, L.L., P. Menozzi, and A. Piazza. 1994. The History and Geography of Human Genes. Princeton: Princeton University Press.
  9. ↑ Arredi B, Poloni ES, Paracchini S, et al. (August 2004). "A predominantly neolithic origin for Y-chromosomal DNA variation in North Africa". American Journal of Human Genetics. 75 (2): 338–45. doi:10.1086/423147. PMC 1216069Freely accessible. PMID 15202071.
  10. ↑ Manni F, Leonardi P, Barakat A, et al. (October 2002). "Y-chromosome analysis in Egypt suggests a genetic regional continuity in Northeastern Africa". Human Biology. 74 (5): 645–58. doi:10.1353/hub.2002.0054. PMID 12495079.
  11. ↑ Cavalli-Sforza. "Synthetic maps of Africa". The History and Geography of Human Genes. ISBN 0691087504.The present population of the Sahara is Sudan in the extreme north, with an increase of Negroid component as one goes south
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  14. ↑ Marchant, Jo (12 April 2013). "Egyptian mummies yield genetic secrets". Nature. Retrieved 13 January 2014.
  15. 1 2 3 4 5 6 Derenko, M.; Malyarchuk, B.; Bahmanimehr, A.; Denisova, G.; Perkova, M.; Farjadian, S.; Yepiskoposyan, L. (2013). "Complete Mitochondrial DNA Diversity in Iranians". PLOS ONE. 8 (11): e80673. doi:10.1371/journal.pone.0080673. PMC 3828245Freely accessible. PMID 24244704.
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  17. 1 2 3 4 5 "Concomitant Replacement of Language and mtDNA in South Caspian Populations of Iran". Curr. Biol. 16: 668–73. April 2006. doi:10.1016/j.cub.2006.02.021. PMID 16581511. Retrieved 4 April 2014.
  18. 1 2 3 Al-Zahery N, Pala M, Battaglia V, et al. (2011). "In search of the genetic footprints of Sumerians: a survey of Y-chromosome and mtDNA variation in the Marsh Arabs of Iraq". BMC Evolutionary Biology. 11: 288. doi:10.1186/1471-2148-11-288. PMC 3215667Freely accessible. PMID 21970613.
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  20. ↑ Dr. Joel J. Elias, Emeritus, University of California, The Genetics of Modern Assyrians and their Relationship to Other People of the Middle East
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  23. ↑ Banoei et al., Human Biology. February 2008, v. 80, no, I, pp. 73-81., "Variation of DAT1 VNTR alleles and genotypes among old ethnic groups in Mesopotamia to the Oxus region" PMID 18505046 "The relationship probability was lowest between Assyrians and other communities. Endogamy was found to be high for this population through determination of the heterogeneity coefficient (+0,6867), Our study supports earlier findings indicating the relatively closed nature of the Assyrian community as a whole, which as a result of their religious and cultural traditions, have had little intermixture with other populations."
  24. ↑ http://ycc.biosci.arizona.edu/nomenclature_system/fig1.html. Retrieved September 16, 2007. Missing or empty |title= (help)
  25. ↑ National Geographic Magazine, October 2004. Available online: http://ngm.nationalgeographic.com/features/world/asia/lebanon/phoenicians-text/1; and http://www.independent.com.mt/news.asp?newsitemid=57215 [accessed: March 10, 2008]
  26. ↑ "Who Were the Phoenicians? - National Geographic Magazine". Ngm.nationalgeographic.com. Retrieved 2014-06-30.
  27. ↑ "National Geographic Special 'Quest for the Phoenicians'". PBS. 2004. Archived from the original on 2004-09-23.
  28. ↑ and http://www.independent.com.mt/news.asp?newsitemid=57215 [Accessed April 11, 2008
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  30. ↑ Mekel-Bobrov N, Gilbert SL, Evans PD, et al. (September 2005). "Ongoing adaptive evolution of ASPM, a brain size determinant in Homo sapiens". Science. 309 (5741): 1720–2. Bibcode:2005Sci...309.1720M. doi:10.1126/science.1116815. PMID 16151010.
  31. 1 2 Shen P, Lavi T, Kivisild T, et al. (September 2004). "Reconstruction of patrilineages and matrilineages of Samaritans and other Israeli populations from Y-chromosome and mitochondrial DNA sequence variation". Human Mutation. 24 (3): 248–60. doi:10.1002/humu.20077. PMID 15300852.
  32. ↑ Behar DM, Yunusbayev B, Metspalu M, et al. (July 2010). "The genome-wide structure of the Jewish people". Nature. 466 (7303): 238–42. Bibcode:2010Natur.466..238B. doi:10.1038/nature09103. PMID 20531471.
  33. 1 2 Hammer MF, Redd AJ, Wood ET, et al. (June 2000). "Jewish and Middle Eastern non-Jewish populations share a common pool of Y-chromosome biallelic haplotypes". Proceedings of the National Academy of Sciences of the United States of America. 97 (12): 6769–74. Bibcode:2000PNAS...97.6769H. doi:10.1073/pnas.100115997. PMC 18733Freely accessible. PMID 10801975.
  34. ↑ Skorecki K; Selig S; Blazer S; et al. (January 1997). "Y chromosomes of Jewish priests". Nature. 385 (6611): 32. Bibcode:1997Natur.385...32S. doi:10.1038/385032a0. PMID 8985243. Archived from the original on February 9, 2007.
  35. ↑ Wade, Nicholas (January 14, 2006). "New Light on Origins of Ashkenazi in Europe". The New York Times. Retrieved 24 May 2006.
  36. ↑ Behar DM, Metspalu E, Kivisild T, et al. (March 2006). "The matrilineal ancestry of Ashkenazi Jewry: portrait of a recent founder event". American Journal of Human Genetics. 78 (3): 487–97. doi:10.1086/500307. PMC 1380291Freely accessible. PMID 16404693.
  37. ↑ Costa MD, Pereira JB, Pala M, et al. (2013). "A substantial prehistoric European ancestry amongst Ashkenazi maternal lineages". Nature Communications. 4: 2543. Bibcode:2013NatCo...4E2543C. doi:10.1038/ncomms3543. PMC 3806353Freely accessible. PMID 24104924.
  38. ↑ Cruciani F; La Fratta R; Torroni A; Underhill PA; Scozzari R (August 2006). "Molecular dissection of the Y chromosome haplogroup E-M78 (E3b1a): a posteriori evaluation of a microsatellite-network-based approach through six new biallelic markers". Human Mutation. 27 (8): 831–2. doi:10.1002/humu.9445. PMID 16835895.
  39. ↑ Arnaiz-Villena, A.; Karin, M.; Bendikuze, N.; Gomez-Casado, E.; Moscoso, J.; Silvera, C.; Oguz, F.S.; Sarper Diler, A.; De Pacho, A.; Allende, L.; Guillen, J.; Martinez Laso, J. (2001). "HLA alleles and haplotypes in the Turkish population: Relatedness to Kurds, Armenians and other Mediterraneans". Tissue Antigens. 57 (4): 308–17. doi:10.1034/j.1399-0039.2001.057004308.x. PMID 11380939.
  40. ↑ Schurr, Theodore G.; Yardumian, Aram (2011). "Who Are the Anatolian Turks?". Anthropology & Archeology of Eurasia. 50 (1): 6–42. doi:10.2753/AAE1061-1959500101.
  41. 1 2 3 Hodoğlugil U; Mahley RW (March 2012). "Turkish population structure and genetic ancestry reveal relatedness among Eurasian populations". Annals of Human Genetics. 76 (2): 128–41. doi:10.1111/j.1469-1809.2011.00701.x. PMID 22332727.
  42. ↑ Rosser, Zoë H.; Zerjal, Tatiana; Hurles, Matthew E.; Adojaan, Maarja; Alavantic, Dragan; Amorim, António; Amos, William; Armenteros, Manuel; Arroyo, Eduardo; Barbujani, Guido (2000). "Y-Chromosomal Diversity in Europe is Clinal and Influenced Primarily by Geography, Rather than by Language". The American Journal of Human Genetics. 67 (6): 1526–43. doi:10.1086/316890. PMC 1287948Freely accessible. PMID 11078479.
  43. ↑ Nasidze, I; Sarkisian, T; Kerimov, A; Stoneking, M (2003). "Testing hypotheses of language replacement in the neighbouring Caucasus: evidence from the Y-chromosome". Human Genetics. 112 (3): 255–61. doi:10.1007/s00439-002-0874-4 (inactive 2015-02-02). PMID 12596050. INIST:14599190.
  44. ↑ Cinnioğlu, Cengiz; King, Roy; Kivisild, Toomas; Kalfoğlu, Ersi; Atasoy, Sevil; Cavalleri, Gianpiero L.; Lillie, Anita S.; Roseman, Charles C.; Lin, Alice A.; Prince, Kristina; Oefner, Peter J.; Shen, Peidong; Semino, Ornella; Cavalli-Sforza, L. Luca; Underhill, Peter A. (2004). "Excavating Y-chromosome haplotype strata in Anatolia". Human Genetics. 114 (2): 127–48. doi:10.1007/s00439-003-1031-4. PMID 14586639.
  45. ↑ Wells RS, Yuldasheva N, Ruzibakiev R, et al. (August 2001). "The Eurasian heartland: a continental perspective on Y-chromosome diversity". Proceedings of the National Academy of Sciences of the United States of America. 98 (18): 10244–9. Bibcode:2001PNAS...9810244W. doi:10.1073/pnas.171305098. JSTOR 3056514. PMC 56946Freely accessible. PMID 11526236.
  46. ↑ Comas D, Schmid H, Braeuer S, et al. (March 2004). "Alu insertion polymorphisms in the Balkans and the origins of the Aromuns". Annals of Human Genetics. 68 (Pt 2): 120–7. doi:10.1046/j.1529-8817.2003.00080.x. PMID 15008791.

Bibliography

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