The Genetic Origin of the Turko-Mongols and Review of The Genetic Legacy of the Mongols. Part 1: The Y-chromosome Lineages of Chinggis Khan The Russian Journal of Genetic Genealogy. Volume 4, No 2 (2012)/Volume 5, No 1 (2013). P. 1-8.

The Russian Journal of Genetic Genealogy (Русская версия): Том 4, №2, 2012 год ISSN: 1920-2997 http://ru.rjgg.org © Все права защищены The Genetic Origin of the Turko-Mongols and Review of The Genetic Legacy of the Mongols. Part 1: The Y-chromosomal Lineages of Chinggis Khan RJGG Kherlen Batbayar Zhaxylyk M. Sabitov Abstract Batu-Mungke Dayan Khan, a direct descendant of Chinggis Khan (commonly known as Genghis Khan) proved himself to be one of the most influential and powerful rulers in Mongolian history. After experiencing decades of East and West Mongolian political strife and incessant battling, Dayan Khan became the sole survivor of Chinggis Khan’s lineage in Mongolia Proper. Chinggis Khan raised five sons, all of whom reached adulthood. Besides Dayan Khan’s descendants residing in Mongolia Proper, only two other offspring of Chinggis Khan’s family line prevailed in the Central Asian countries of Kazakhstan and Uzbekistan. These two lineages’ modal haplotypes belong to haplogroup C3 under different subgroups. However, according to a genetic research study, The Genetic Legacy of the Mongols, Chinggis Khan’s descendants number approximately 16 million. One may only hypothesize that this calculation is unreasonably inaccurate when studying both the historical and genetic, or genealogical, points-of-view. In this study, we have determined Batu-Mungke Dayan Khan’s haplotype by analyzing 12 Ychromosomal STR loci of seven surviving elders of the House of Dayan Khan. Our findings differ significantly from those provided in The Genetic Legacy of Mongols. Chinggis Khan’s three surviving lineages’ Y-chromosome DNA test results suggest the near-extinction, not the proliferation, of the direct male descendant line of Chinggis Khan. Introduction In 2003, Tatiana Zerjal and her colleagues identified a Y-chromosomal lineage that profoundly affects mankind’s genealogical history. This Y-chromosomal lineage presented itself in approximately 8% of the men living in the vast region stretching from the Pacific to the Caspian Sea [1]. It is referred to as the star-cluster C3*, also celebrated as Chinggis Khan’s Ychromosome DNA. The central star-cluster C3* profile is: 10-16-25-10-11-13-14-12-11-11-11-12-8-1010, for the loci DYS389I-DYS389b-DYS390-DYS391DYS392-DYS393-DYS388-DYS425-DYS426DYS434-DYS435-DYS436-DYS437-DYS438DYS439 [1]. ___________________________________________________________ Received: September 6 2011; accepted: September 7 2012; published: September 12 2012 Correspondence: altinsarana@gmail.com babasan@yandex.ru 1 According to their calculations, this pattern variation occurred in Mongolia around one thousand years ago. Therefore, the conclusion was that the Y-chromosome lineage was restricted to Chinggis Khan and his close male-line relatives [1]. However, none of the legitimate descendants of Chinggis Khan were actually tested in the research study by Zerjal and colleagues. In 2009, Sabitov Zh. M. demonstrated that the Kazakh Tores, whose ancestor is Chinggis Khan’s oldest son Jochi, possessed a different C3 subhaplogroup while the star-cluster C3* was the modal haplotype of two Kazakh Kerey clans (Abakand Ashamaily) [2]. Later, in 2012, the same result was obtained by Abilevet al after testing 51 Kazakh Kereys, in which 76.5% of the Kereys possessed the star-cluster C3* [3]. Contrary to their statement, Kerey is not the dominant clan in Kazakhstan; it is indeed one of the numerically few tribes. Therefore, the unusual distribution of the star-cluster C3* Y-chromosome cannot be fully explained by its presence in the Kereys The Russian Journal of Genetic Genealogy (Русская версия): Том 4, №2, 2012 год ISSN: 1920-2997 http://ru.rjgg.org © Все права защищены alone. In the preceding year, Turuspekov et al identified the star-cluster C3* in a minor branch of Chinggis Khan’s grandson Shiban, son of Jochi [4]. However, the same DNA sequence was also dominant in many other Kazakh tribes that are not directly related to Chinggis Khan. These contradicting findings resulted in testing the offspring of Batu-Mungke Dayan Khan, descendant of Chinggis Khan’s fourth son Tolui. Tolui’s descendants live in The State of Mongolia (Outer Mongolia) and Inner Mongolia (an autonomous region in The People’s Republic of China). RJGG 19th century. Therefore, today, only three officially recognized branches of Chinggisids (Chinggis Khan’s line of descent) are known. One branch is descended from Jani-Beg Khan, who fathered the Kazakh royal house (Figure 1). Jani-Beg Khan’s ancestor is Tuka-Timur, one of the younger sons of Jochi [2]. The Kazakh Chinggisids are also known by Tore in Kazakhstan. Tuka-Timur fathered the Crimean Tatar royal family (the Gireys) as well. However, the Gireys were not tested in this study. Currently, another minor lineage of Jochi exists in Kazakhstan and Uzbekistan. The predecessor of this royal house is Shiban, Jochi’s fifth son (Figure 1). The Shibanids were once the rulers of the Khanate of Khiva (Khwarezm). The Kazakh Shibanids’ common ancestor is Shah-Temir Khan (18th century). The better known branch is the Mongolian royal house, which was founded by Batu-Mungke Dayan Khan. By various accounts, Batu-Mungke Dayan Khan was Chinggis’s grandson Kublai Khan’s direct male-line descendant [7, 8]. In the 15th century, BatuMungke Dayan Khan reunited the majority of the warring Mongol tribes, successfully continuing Chinggis Khan’s State with the help of his Queen-consort, Mandukhai-the-Wise. After the death of his uncle, Mandoul Khan, and his father, Bayan-Mungke Jinong, young BatuMungke became the only surviving descendant of Kublai Khan in Mongolia [7]. However, the number of Kublai’s descendants was significantly reduced, long before Batu-Mungke’s birth, during the reign of the Western Mongolian ruler Esen-Taishi (d. 1455), who mercilessly massacred the Mongolian Chinggisids [7, 8]. In this study, seven patrilineal descendants of Batu-Mungke Dayan Khan and two of Chinggis Khan’s brothers’ descendants were chosen for Y-chromosome DNA sequencing. Rather than testing a multitude of subjects, for the sake of accuracy, the most legitimate and proven descendants of Dayan Khan were selected. The DNA donors were selected based upon their official Mongol and Manchu titles and ranks, which were precisely recorded in Mongolian, Manchu, and Soviet documents. Background Chinggis Khan’s Legacy Chinggis Khan had five recognized sons who reached adulthood: Jochi, Chagatai, Ogetei, Tolui, and Khulgen [5, 6]. Ogetei’s and Khulgen’s descendants had completely disappeared by the end of the 15th century. The last descendants of Chagatai presided over a sector of modern-day Tashkent, Uzbekistan, until the second half of the 18th century, completely disappearing in the 2 The Russian Journal of Genetic Genealogy (Русская версия): Том 4, №2, 2012 год ISSN: 1920-2997 http://ru.rjgg.org © Все права защищены RJGG Figure 1. Chinggis Khan's Mini Family Tree Demonstrating the Roots of Uzbek-Kazakh Shibanids, Kazakh Khans, and Mongolian Khans. single adult male of the Dayan Khan noble family was massacred by the orders of Joseph Stalin and the Communist Party of the Soviet Union [9]. The mass murdering began in October 1937 and ended in April 1939, with a total of 25,824 Mongolian men dead, including the nobles, Buddhist lamas, and commoners [9]. Distribution of the Borjigin Clan and the Dayan Khanid Aristocracy in Mongolia During his lifetime, Batu-Mungke sired eleven sons with three wives. His grandchildren eventually took the place of most of the former tribal chiefs, and a new aristocracy was formed to ensure the unity and continuity of Chinggisid sovereignty. Dayan Khan and his senior wife, Mandukhai-the-Wise, raised seven sons: Torebolat, Ulusbolat, Arsubolat, Barsbolat, Ochirbolat, Alchubolat, and Arabolat (Albor) [8]. Today, the progeny of Dayan Khan−Torebolat, Barsbolat, Ochirbolat, and Alchubolat − live in Inner Mongolia. Dayan Khan also fathered four sons, Prince Gerebolat, Prince Geresenz, Prince Ching, and Prince Geretu, with his two younger wives [8]. One of these four sons, Geresenz, outlived the other three, all of whom have left no descendants [8]. According to the official archival data from 1921, a total of 13,274 (approximately 2.65% of the total population in 1921) direct male descendants of Prince Geresenz were recorded living in Outer Mongolia and 8,996 of these men were over 18-years-old [9]. However, during the Great Purge of 1936-1939, every Chinggis Khan’s royal family belonged to the Borjigin sub-clan of the Kiyat clan, which is one of the 19 Nirun clans [5, 6, 10]. As recorded by the historians from the Mongol Ilkhanate period (1256-1335), only Chinggis Khan’s father Yesukhei’s sons inherited the royal clan name Borjigin [5]. However, currently 20% of the Mongolians are officially bearing Borjigin as their clan name notwithstanding the fact that the Borjigins were massively murdered twice, first during Esen Taishi’s reign and then the Great Purge. Therefore, a more scrupulous method was needed in order to select the authentic biological descendants of Dayan Khan from one fifth of the population carrying the same clan name. All male descendants of Batu-Mungke Dayan Khan were granted the title Taij (baron) by birth until 1939. Four particular lineages possessed the title Khan (king) and ruled the commoners as 3 RJGG The Russian Journal of Genetic Genealogy (Русская версия): Том 4, №2, 2012 год ISSN: 1920-2997 http://ru.rjgg.org © Все права защищены well as the Taij nobles. These titles were inheritable only on a descending scale. Furthermore, Mongolian noblemen were also granted Manchu ranks from the Qing (Manchu) Empire. When the Qing Empire subjugated the Mongolians, they introduced the Manchu banner (Gusha) system. By this system, all former Borjigin tribal leaders became hereditary banner chiefs and received the top four Qing (Manchu) ranks: Hoshoi Qinwang, Doroi Junwang, Doroi Beile, or Gusha Beizi. By 1911, at the end of the 230 (287 years in Inner Mongolia) years of Qing (Manchu) Dynasty’s domination, Outer Mongolia had 4 Khansand 86 banner chiefs; and Inner Mongolia had 49 banner chiefs. Some Mongolian noblemen also received the Manchu title Efu (prince consort) through marriage. Method Buccal mucosal cell samples were collected meticulously between June 2010 and February 2012. Genealogical relationships were recorded prior to sample collection, confirming that all participants were unrelated within at least the last six to seven generations. Furthermore, in order to minimize genetic distancing between the most recent common ancestor of the participants and the progenitor, Dayan Khan, the descendants of three different sons of Dayan Khan were studied (Table 1). To reduce the possibility of any skewing of the data due to alternative paternity, the progeny of two sons of the queen-consort, Mandukhai-the-Wise, and the progeny of Prince Geresenz, a son from a different wife, were selected. Table 1. Traditional Social Status of the Nine Borjigin DNA Donors: Mongolian Appellations and Manchu Ranks are shown. Identification Appellation Manchu Rank Meaning in English Geresenz’sProgeny 1 Khan HoshoiQinwang Prince of First Rank Geresenz’sProgeny 3 Geresenz’sProgeny 4 Taij Taij GushaBeizi GushaBeizi Banner Chief Banner Chief Geresenz’sProgeny 2 Torebolat’sProgeny Alchubolat’sProgeny 1 Alchubolat’sProgeny 2 Khasar’sProgeny Belgutei’sProgeny Taij DoroiJunwang Taij DoroiBeile Taij Efu Taij GushaBeizi Wang GushaBeizi Tawnang unknown Prince of Second Rank Lord Prince Consort Banner Chief Banner Chief unknown DYS393-DYS390-DYS394-DYS391-DYS385aDYS385b-DYS426-DYS388-DYS439-DYS389iDYS392-DYS389ii. Moreover, one descendant of Chinggis Khan’s full-brother Khasar and a descendant of Chinggis Khan’s paternal half-brother Belgutei were tested as a control group. The participants were tested by the standard male-line testing kits from Genealogy By Genetics DBA DNA Findings. A total of 12 Y-chromosome short tandem repeat (STR) loci were analyzed for each sample: For STR haplotype analysis, Dayan Khan’s YDNA result was compared with the Y-STR haplotypes of two separate lineages of Jochi for the same loci. 4 RJGG The Russian Journal of Genetic Genealogy (Русская версия): Том 4, №2, 2012 год ISSN: 1920-2997 http://ru.rjgg.org © Все права защищены Figure 2. Batu-Mungke Dayan Khan's Mini Family Tree Illustrating the Ancestry of the Y-DNA Donors. despite the fact that Torebolat’s descendant belongs to haplogroup C3 under different subgroup. The rest of the samples belong to haplogroups D3a; O3; Q; and R1a1a. Participants of high hereditary ranks demonstrated matching results, including the participant with the highest Mongolian title Khan. All three C3 (M407+) participants possessed Manchu princely titles: Hoshoi Qinwang (Prince of First Rank), Doroi Junwang (Prince of Second Rank), and Efu (Prince Consort). Results Genotyping revealed that Batu-Mungke Dayan Khan belonged to haplogroupC3 (M407+). Analysis of 12 fast evolving Y-STRs provided that three out of the nine samples are identical, i.e. they share the same haplotype as shown in Table 2. Two of them are from Prince Geresenz’s lineage and one sample is from Alchubolat’s lineage. The remaining samples did not demonstrate any close patrilineal heritage; DYS393 DYS390 DYS19 DYS391 DYS 385 DYS426 DYS388 DYS439 DYS389i DYS392 DYS389ii Haplogroup Identification Table 2. Y-chromosomal Short Tandem Repeat (Y-STR) Haplotypes of the Borjigin DNA Donors from Outer and Inner Mongolia. Geresenz's Progeny 1 C3 14 23 15 10 11;11 11 12 11 13 11 28 Geresenz's Progeny 3 Q 13 23 14 12 16;16 12 12 12 13 16 30 Torebolat's Progeny C3 13 24 15 Alchubolat's Progeny 2 D3a 13 25 15 Belgutei's Progeny R1a1a 14 25 16 Geresenz's Progeny 2 Geresenz's Progeny 4 Alchubolat's Progeny 1 Khasar's Progeny C3 O3 C3 O3 14 14 14 13 23 24 23 23 15 15 10 10 9 11;11 14;21 12;15 11 11 11 12 9 13 11 11 11 13 12 13 11 13 11 28 28 29 15 10 11;11 11 12 11 13 11 28 15 10 12;15 11 12 13 12 12 27 5 10 11 11;11 11;14 11 12 12 12 12 10 14 13 7 11 30 30 RJGG The Russian Journal of Genetic Genealogy (Русская версия): Том 4, №2, 2012 год ISSN: 1920-2997 http://ru.rjgg.org © Все права защищены When the results were compared with the entire database of Genealogy By Genetics DBA DNA Findings, an exact-match rate of 4.1% (24/588) was observed from unknown DNA donors from the State of Mongolia. Moreover, five people from the State of Mongolia showed Genetic Distance-1, i.e. the mismatch is at only one marker and the individuals are tightly related. No matches were detected from other regions that are high in haplogroup C3, such as Kazakhstan, Kyrgyzstan, and Uzbekistan, except for a less-than 0.1% match from Russian Siberian minorities. Finally, Alchubolat’s and Geresenz’s lineages are directly branched from Dayan Khan himself (Figure 2) and these two sons of Dayan Khan were born from different mothers. Thus, we conclude that Dayan Khanid’s modal haplotype is: descendants of Chinggis Khan (Table 3). Genetic heterogeneity was present in the main male-line successors of Chinggis Khanas early as the 15th century. Jani-Beg Khan’s descendants’, i.e. the Kazakh Tore lineage’s, modal haplotype is: 14-24-15-10-12-14-11-13-12-13-11-29 [2]. The characteristic mutation of their haplotype is RecLOH in DYS 448 [11]. On the contrary, the modal haplotype of Shah-Temir’s descendants is: 13-25-16-10-12-13-11-14-10-13-11-29 [4]. Comparison of Dayan Khan’s Y-STR haplotype: 14-23-15-10-11-11-11-12-11-13-11-28, 14-23-15-10-11-11-11-12-11-13-11-28 with the modal haplotypes of the Jochids demonstrated that while all three of them belong to haplogroup C3, no haplotypes were shared when considering the full 12 Y-STR markers. Based upon a mutation rate of 0.002 for Y-DNA STR markers, the three Chinggisids most likely shared a common paternal ancestor more than 150 generations ago, which is approximately 4,500 years or more for a generation time of 30 years. Moreover, Batu-Mungke Dayan Khan’s and Jani-Beg Khan’s Ychromosome sequences each constitute less than 3.5% of the Central Asian gene pool based on various studies from Kazakhstan, Mongolia, and Uzbekistan, as demonstrated in figures 3 and 4. for the loci DYS393-DYS390-DYS394-DYS391DYS385a-DYS385b-DYS426-DYS388-DYS439DYS389i-DYS392-DYS389ii. Both samples from Chinggis Khan’s brothers’ descendants failed to provide any further information. Khasar’s descendant belongs to haplogroup O3 and Belgutei’s descendant belongs to haplogroup R1a1a. Discussion Y-STR haplotype analysis revealed the heterogeneous paternal origins of the modern day Chinggisids DYS393 DYS390 DYS394 DYS391 DYS385 DYS426 DYS388 DYS439 DYS389i DYS392 DYS389ii Haplogroup Table 3. Y-STR Haplotype Results and Haplogroup Classification of the Three Separate Male-line Lineages of Chinggis Khan. Dayan Khanid 14 23 15 10 11;11 11 12 11 13 11 28 C3 Kazakh Tore Shibanid 14 13 24 25 15 16 10 10 12;14 12;13 11 11 13 14 12 10 13 13 11 11 29 29 C3 C3 that Chinggis Khan and his close male-line relatives were the primary distributors of this Ychromosome [1]. However, the members from this particular Shiban lineage are significantly The Shibanids modal haplotype matched the hypothetical Y-chromosome DNA sequence of Chinggis Khan, i.e. the star-cluster C3*. The 2003 research study by Zerjalet al suggested 6 The Russian Journal of Genetic Genealogy (Русская версия): Том 4, №2, 2012 год ISSN: 1920-2997 http://ru.rjgg.org © Все права защищены few, numbering less than a thousand in Kazakhstan and Uzbekistan combined, and they are absent elsewhere including Mongolia, Afghanistan, Kyrgyzstan, etc. Moreover, as we have demonstrated no other Chinggisid lineage possesses the star-cluster C3*. On the contrary, the star-cluster C3* is the defining Y-STR modal haplotype of the Kazakh Kereys. Furthermore, two new studies by Sabitov Zh. M. and Turuspe- RJGG kov et al suggested that besides the Kereys, the primary carriers of the star-cluster C3* are the Uly Zhuz (Great Horde) Kazakhs, i.e. the Uysuns [4, 12]. Approximately one third of the entire Kazakh population belongs to the Uly Zhuz. The star-cluster C3* was also found from many other non-Chinggisid Turko-Mongol tribes from Mongolia, Karakalpakstan (an autonomous republic in Uzbekistan), and India. Conclusion Shibanids are absent elsewhere, including countries that are traditionally linked to the Mongol Empire, such as Afghanistan and Mongolia. Finally, contrary to previous studies by Zerjalet al, the star-cluster C3* is the modal haplotype of various Turko-Mongol tribes that are not directly related to Chinggis Khan. Therefore, the starcluster C3* or Chinggis Khan’s hypothetical Ychromosome was either the dominant haplotype of most Mongolian men in the 13th century or it had been well distributed throughout Central Asia before Chinggis Khan’s time due to the territorial expansions of various nomadic empires. To summarize, Chinggis Khan’s Ychromosome is not as widespread as previously thought. The three remaining lineages of Chinggis Khan belong to three different subgroups of haplogroup C3. Although haplogroup C3 is the dominant haplogroup in both Kazakhstan and Mongolia, Dayan Khan’s and Jani-Beg Khan’s haplotypes each constitute less than 3.5% of the Central Asian (i.e. Kazakhstan, Mongolia, and Uzbekistan) Y-chromosome gene pool. Out of all the haplogroup C3 DNA sequences, these two haplotypes are the least distributed in Central Asia. The primary Mongolian Chinggisid lineage, or the Dayan Khanid aristocracy, possesses haplogroup C3 (M407+). The divergence time between Dayan Khan’s lineage and Jani-Beg Khan’s lineage is more than 4,500 years for a generation time of 30 years. The third Chinggisid branch belongs to the hypothetical haplotype of Chinggis Khan, the star-cluster C3*. However, the number of the members of this particular branch is a lot scarcer than the previously mentioned two major branches. The estimated number of the Shibanids is less than a thousand altogether in Kazakhstan and Uzbekistan. The Acknowledgements A special thanks to all our wonderful DNA donors; Sharaid Uuganbayar, Sartuul Tugulgur, and Khongorzul Enkh-Amgalan, for collecting the DNA samples; and Mary Mills and Daniel Zniderzicfor editing the manuscript and giving incredibly valuable advice. 7 The Russian Journal of Genetic Genealogy (Русская версия): Том 4, №2, 2012 год ISSN: 1920-2997 http://ru.rjgg.org © Все права защищены References RJGG 6. Muizz Al Ansab. History of Kazakhstan in Persian Sources. T. 3. 2006. Almaty, Kazakhstan: Daik Press. 7. Atwood C.P. (2004) Encyclopedia of Mongolia and the Mongol Empire. New York: Facts on File. 8. Shagdar Ts. (1990) Luvsandanzan Altan Tovch. Ulaanbaatar, Mongolia: State Press Agency. 9. Natsagdorj Ts. (2008) Altan Ourgiin Soulchiin Toin. Ulaanbaatar, Mongolia: Bit Press. 10. Miles C. (1838) The Shajrat Ul Atrak or the Genealogical Tree of the Turks and Tatars. London, England: W M. H. Allen, and Co. 11. Sabitov Zh. M. (2009) In Search of Genghis Khan. 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