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The Russian Journal of Genetic (Русская версия): Том 4, №2, 2012 год ISSN: 1920-2997 http://ru.rjgg.org © Все права защищены RJGG

The Genetic Origin Kherlen Batbayar of the Turko- and Review Zhaxylyk M. Sabitov of The Genetic Legacy of the Mongols. Part 1: The Y-chromosomal Lineages of Chinggis

Abstract

Batu-Mungke Dayan Khan, a direct descendant of Chinggis Khan (commonly known as ) 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 Proper. Chinggis Khan raised five , all of whom reached adulthood. Besides Dayan Khan’s descendants residing in Mongolia Proper, only two other offspring of Chinggis Khan’s 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 hypothes- ize 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 Y- chromosomal 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 According to their calculations, this pattern In 2003, Tatiana Zerjal and her colleagues variation occurred in Mongolia around one thou- identified a Y-chromosomal lineage that pro- sand years ago. Therefore, the conclusion was foundly affects mankind’s genealogical history. that the Y-chromosome lineage was restricted to This Y-chromosomal lineage presented itself in Chinggis Khan and his close male-line relatives approximately 8% of the men living in the vast [1]. However, none of the legitimate descen- region stretching from the Pacific to the Caspian dants of Chinggis Khan were actually tested in Sea [1]. It is referred to as the star-cluster C3*, the research study by Zerjal and colleagues. In also celebrated as Chinggis Khan’s Y- 2009, Sabitov Zh. M. demonstrated that the Ka- chromosome DNA. The central star-cluster C3* zakh Tores, whose is Chinggis Khan’s profile is: oldest , possessed a different C3 sub- haplogroup while the star-cluster C3* was the 10-16-25-10-11-13-14-12-11-11-11-12-8-10- modal haplotype of two Kazakh Kerey (Ab- 10, akand Ashamaily) [2]. Later, in 2012, the same result was obtained by Abilevet al after testing for the loci DYS389I-DYS389b-DYS390-DYS391- 51 Kazakh Kereys, in which 76.5% of the Kereys DYS392-DYS393-DYS388-DYS425-DYS426- possessed the star-cluster C3* [3]. Contrary to DYS434-DYS435-DYS436-DYS437-DYS438- their statement, Kerey is not the dominant DYS439 [1]. in Kazakhstan; it is indeed one of the numerical-

______ly few tribes. Therefore, the unusual distribution

Received: September 6 2011; accepted: September 7 2012; of the star-cluster C3* Y-chromosome cannot be published: September 12 2012 fully explained by its presence in the Kereys Correspondence: [email protected] [email protected] 1

The Russian Journal of (Русская версия): Том 4, №2, 2012 год ISSN: 1920-2997 http://ru.rjgg.org © Все права защищены RJGG alone. In the preceding year, Turuspekov et al 19th century. Therefore, today, only three offi- identified the star-cluster C3* in a minor branch cially recognized branches of Chinggisids of Chinggis Khan’s grandson Shiban, son of Jochi (Chinggis Khan’s line of descent) are known. [4]. However, the same DNA sequence was also One branch is descended from Jani-Beg Khan, dominant in many other Kazakh tribes that are who fathered the Kazakh royal house (Figure 1). not directly related to Chinggis Khan. These con- Jani-Beg Khan’s ancestor is Tuka-Timur, one of tradicting findings resulted in testing the the younger sons of Jochi [2]. The Kazakh offspring of Batu-Mungke Dayan Khan, descen- Chinggisids are also known by Tore in Ka- dant of Chinggis Khan’s fourth son Tolui. Tolui’s zakhstan. Tuka-Timur fathered the Crimean Ta- descendants live in The State of Mongolia (Outer tar royal family (the Gireys) as well. However, Mongolia) and Inner Mongolia (an autonomous the Gireys were not tested in this study. Cur- region in The People’s Republic of China). rently, another minor lineage of Jochi exists in Kazakhstan and Uzbekistan. The predecessor of In this study, seven patrilineal descendants this royal house is Shiban, Jochi’s fifth son (Fig- of Batu-Mungke Dayan Khan and two of Ching- ure 1). The Shibanids were once the rulers of gis Khan’s ’ descendants were chosen the of Khiva (Khwarezm). The Kazakh for Y-chromosome DNA sequencing. Rather than Shibanids’ common ancestor is Shah-Temir testing a multitude of subjects, for the sake of Khan (18th century). The better known branch is accuracy, the most legitimate and proven des- the Mongolian royal house, which was founded cendants of Dayan Khan were selected. The DNA by Batu-Mungke Dayan Khan. By various ac- donors were selected based upon their official counts, Batu-Mungke Dayan Khan was Ching- Mongol and Manchu titles and ranks, which were gis’s grandson ’s direct male-line precisely recorded in Mongolian, Manchu, and descendant [7, 8]. In the 15th century, Batu- Soviet documents. Mungke Dayan Khan reunited the majority of the warring Mongol tribes, successfully continuing Chinggis Khan’s State with the help of his Background Queen-consort, Mandukhai-the-Wise. After the death of his , Mandoul Khan, and his fa- Chinggis Khan’s Legacy ther, Bayan-Mungke Jinong, young Batu- Mungke became the only surviving descendant Chinggis Khan had five recognized sons who of Kublai Khan in Mongolia [7]. However, the reached adulthood: Jochi, Chagatai, Ogetei, To- number of Kublai’s descendants was significantly lui, and Khulgen [5, 6]. Ogetei’s and Khulgen’s reduced, long before Batu-Mungke’s birth, dur- descendants had completely disappeared by the ing the reign of the Western Mongolian ruler end of the 15th century. The last descendants of Esen-Taishi (d. 1455), who mercilessly massa- Chagatai presided over a sector of modern-day cred the Mongolian Chinggisids [7, 8]. Tashkent, Uzbekistan, until the second half of the 18th century, completely disappearing in the

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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.

Distribution of the Clan single adult male of the Dayan Khan noble fami- and the Dayan Khanid Aristocracy ly was massacred by the orders of Joseph Stalin in Mongolia and the Communist Party of the Soviet Union [9]. The mass murdering began in October 1937 During his lifetime, Batu-Mungke sired ele- and ended in April 1939, with a total of 25,824 ven sons with three . His grandchildren Mongolian men dead, including the nobles, eventually took the place of most of the former Buddhist lamas, and commoners [9]. tribal chiefs, and a new aristocracy was formed to ensure the unity and continuity of Chinggisid Chinggis Khan’s royal family belonged to the sovereignty. Dayan Khan and his senior , Borjigin sub-clan of the Kiyat clan, which is one Mandukhai-the-Wise, raised seven sons: Tore- of the 19 Nirun clans [5, 6, 10]. As recorded by bolat, Ulusbolat, Arsubolat, Barsbolat, Ochirbo- the historians from the Mongol period lat, Alchubolat, and Arabolat (Albor) [8]. Today, (1256-1335), only Chinggis Khan’s Ye- the progeny of Dayan Khan−Torebolat, Barsbo- sukhei’s sons inherited the royal clan name Bor- lat, Ochirbolat, and Alchubolat − live in Inner jigin [5]. However, currently 20% of the Mongo- Mongolia. Dayan Khan also fathered four sons, lians are officially bearing Borjigin as their clan Prince Gerebolat, Prince Geresenz, Prince Ching, name notwithstanding the fact that the and Prince Geretu, with his two younger wives were massively murdered twice, first during [8]. One of these four sons, Geresenz, outlived Esen Taishi’s reign and then the Great Purge. the other three, all of whom have left no des- Therefore, a more scrupulous method was cendants [8]. According to the official archival needed in order to select the authentic biological data from 1921, a total of 13,274 (approximate- descendants of Dayan Khan from one fifth of the ly 2.65% of the total population in 1921) direct population carrying the same clan name. All male descendants of Prince Geresenz were rec- male descendants of Batu-Mungke Dayan Khan orded living in Outer Mongolia and 8,996 of were granted the title Taij (baron) by birth until these men were over 18-years-old [9]. Howev- 1939. Four particular lineages possessed the er, during the Great Purge of 1936-1939, every title Khan (king) and ruled the commoners as

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The Russian Journal of Genetic Genealogy (Русская версия): Том 4, №2, 2012 год ISSN: 1920-2997 http://ru.rjgg.org © Все права защищены RJGG well as the Taij nobles. These titles were inherit- Method able only on a descending scale. Furthermore, Mongolian noblemen were also granted Manchu Buccal mucosal cell samples were collected ranks from the Qing (Manchu) Empire. When meticulously between June 2010 and February the Qing Empire subjugated the Mongolians, 2012. Genealogical relationships were recorded they introduced the Manchu banner (Gusha) prior to sample collection, confirming that all system. By this system, all former Borjigin tribal participants were unrelated within at least the leaders became hereditary banner chiefs and last six to seven generations. Furthermore, in received the top four Qing (Manchu) ranks: Ho- order to minimize genetic distancing between shoi Qinwang, Doroi Junwang, Doroi Beile, or the most recent common ancestor of Gusha Beizi. By 1911, at the end of the 230 the participants and the , Dayan (287 years in Inner Mongolia) years of Qing Khan, the descendants of three different sons of (Manchu) ’s domination, Outer Mongolia Dayan Khan were studied (Table 1). To reduce had 4 Khansand 86 banner chiefs; and Inner the possibility of any skewing of the data due to Mongolia had 49 banner chiefs. Some Mongolian alternative paternity, the progeny of two sons of noblemen also received the Manchu title Efu the queen-consort, Mandukhai-the-Wise, and (prince consort) through . 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 2 Taij DoroiJunwang Prince of Second Rank Geresenz’sProgeny 3 Taij GushaBeizi Banner Chief Geresenz’sProgeny 4 Taij GushaBeizi Banner Chief Torebolat’sProgeny Taij DoroiBeile Lord Alchubolat’sProgeny 1 Taij Efu Prince Consort Alchubolat’sProgeny 2 Taij GushaBeizi Banner Chief Khasar’sProgeny Wang GushaBeizi Banner Chief Belgutei’sProgeny Tawnang unknown unknown

Moreover, one descendant of Chinggis Khan’s DYS393-DYS390-DYS394-DYS391-DYS385a- full- Khasar and a descendant of Chinggis DYS385b-DYS426-DYS388-DYS439-DYS389i- Khan’s paternal half-brother Belgutei were DYS392-DYS389ii. tested as a control group. The participants were tested by the standard male-line testing kits For STR haplotype analysis, Dayan Khan’s Y- from Genealogy By DBA DNA Findings. DNA result was compared with the Y-STR haplo- A total of 12 Y-chromosome short tandem re- types of two separate lineages of Jochi for the peat (STR) loci were analyzed for each sample: same loci.

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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 2. Batu-Mungke Dayan Khan's Mini Family Tree Illustrating the Ancestry of the Y-DNA Donors.

Results despite the fact that Torebolat’s descendant be- longs to haplogroup C3 under different sub- Genotyping revealed that Batu-Mungke group. The rest of the samples belong to haplo- Dayan Khan belonged to haplogroupC3 groups D3a; O3; Q; and R1a1a. Participants of (M407+). Analysis of 12 fast evolving Y-STRs high hereditary ranks demonstrated matching provided that three out of the nine samples are results, including the participant with the highest identical, i.e. they share the same haplotype as Mongolian title Khan. All three C3 (M407+) par- shown in Table 2. Two of them are from Prince ticipants possessed Manchu princely titles: Ho- Geresenz’s lineage and one sample is from Alc- shoi Qinwang (Prince of First Rank), Doroi Jun- hubolat’s lineage. The remaining samples did wang (Prince of Second Rank), and Efu (Prince not demonstrate any close patrilineal heritage; Consort).

Table 2. Y-chromosomal Short Tandem Repeat (Y-STR) Haplotypes of the Borjigin DNA Donors from Outer and Inner Mongolia.

385 DYS DYS19 DYS393 DYS390 DYS391 DYS426 DYS388 DYS439 DYS392 DYS389i DYS389ii entification Haplogroup Id

Geresenz's Progeny 1 C3 14 23 15 10 11;11 11 12 11 13 11 28 Geresenz's Progeny 2 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 Geresenz's Progeny 4 O3 14 24 15 10 14;21 11 9 11 12 13 28 Torebolat's Progeny C3 13 24 15 9 12;15 11 13 11 13 11 29

Alchubolat's Progeny 1 C3 14 23 15 10 11;11 11 12 11 13 11 28 Alchubolat's Progeny 2 D3a 13 25 15 10 11;11 11 12 12 14 7 30 Khasar's Progeny O3 13 23 15 10 12;15 11 12 13 12 12 27

Belgutei's Progeny R1a1a 14 25 16 11 11;14 12 12 10 13 11 30

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The Russian Journal of Genetic Genealogy (Русская версия): Том 4, №2, 2012 год ISSN: 1920-2997 http://ru.rjgg.org © Все права защищены RJGG

When the results were compared with the descendants of Chinggis Khan (Table 3). Genetic entire database of Genealogy By Genetics DBA heterogeneity was present in the main male-line DNA Findings, an exact-match rate of 4.1% successors of Chinggis Khanas early as the 15th (24/588) was observed from unknown DNA do- century. Jani-Beg Khan’s descendants’, i.e. the nors from the State of Mongolia. Moreover, five Kazakh Tore lineage’s, modal haplotype is: people from the State of Mongolia showed Ge- netic Distance-1, i.e. the mismatch is at only 14-24-15-10-12-14-11-13-12-13-11-29 [2]. one marker and the individuals are tightly re- lated. No matches were detected from other re- The characteristic of their haplotype gions that are high in haplogroup C3, such as is RecLOH in DYS 448 [11]. On the contrary, the Kazakhstan, Kyrgyzstan, and Uzbekistan, except modal haplotype of Shah-Temir’s descendants for a less-than 0.1% match from Russian Sibe- is: rian minorities. Finally, Alchubolat’s and Gere- senz’s lineages are directly branched from 13-25-16-10-12-13-11-14-10-13-11-29 [4]. Dayan Khan himself (Figure 2) and these two sons of Dayan Khan were born from different Comparison of Dayan Khan’s Y-STR haplo- . Thus, we conclude that Dayan Khanid’s type: modal haplotype is: 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 dem- for the loci DYS393-DYS390-DYS394-DYS391- onstrated that while all three of them belong to DYS385a-DYS385b-DYS426-DYS388-DYS439- haplogroup C3, no haplotypes were shared when DYS389i-DYS392-DYS389ii. considering the full 12 Y-STR markers. Based upon a mutation rate of 0.002 for Y-DNA STR Both samples from Chinggis Khan’s brothers’ markers, the three Chinggisids most likely descendants failed to provide any further infor- shared a common paternal ancestor more mation. Khasar’s descendant belongs to haplo- than 150 generations ago, which is approx- group O3 and Belgutei’s descendant belongs to imately 4,500 years or more for a genera- haplogroup R1a1a. tion time of 30 years. Moreover, Batu-Mungke Dayan Khan’s and Jani-Beg Khan’s Y- chromosome sequences each constitute less Discussion than 3.5% of the Central Asian gene pool based on various studies from Kazakhstan, Mongolia, Y-STR haplotype analysis revealed the hete- and Uzbekistan, as demonstrated in figures 3 rogeneous paternal origins of the modern day and 4.

Table 3. Y-STR Haplotype Results and Haplogroup Classification of the Three Separate Male-line Lineages of Chinggis Khan.

DYS393 DYS390 DYS394 DYS391 DYS385 DYS426 DYS388 DYS439 DYS392 DYS389i DYS389ii Chinggisids Haplogroup

Dayan Khanid 14 23 15 10 11;11 11 12 11 13 11 28 C3

Kazakh Tore 14 24 15 10 12;14 11 13 12 13 11 29 C3 Shibanid 13 25 16 10 12;13 11 14 10 13 11 29 C3

The Shibanids modal haplotype matched the that Chinggis Khan and his close male-line rela- hypothetical Y-chromosome DNA sequence of tives were the primary distributors of this Y- Chinggis Khan, i.e. the star-cluster C3*. The chromosome [1]. However, the members from 2003 research study by Zerjalet al suggested this particular Shiban lineage are significantly

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The Russian Journal of Genetic Genealogy (Русская версия): Том 4, №2, 2012 год ISSN: 1920-2997 http://ru.rjgg.org © Все права защищены RJGG few, numbering less than a thousand in Ka- kov et al suggested that besides the Kereys, the zakhstan and Uzbekistan combined, and they primary carriers of the star-cluster C3* are the are absent elsewhere including Mongolia, Afgha- Uly Zhuz (Great Horde) , i.e. the Uysuns nistan, Kyrgyzstan, etc. Moreover, as we have [4, 12]. Approximately one third of the entire demonstrated no other Chinggisid lineage pos- Kazakh population belongs to the Uly Zhuz. The sesses the star-cluster C3*. On the contrary, the star-cluster C3* was also found from many oth- star-cluster C3* is the defining Y-STR modal er non-Chinggisid Turko-Mongol tribes from haplotype of the Kazakh Kereys. Furthermore, Mongolia, Karakalpakstan (an autonomous re- two new studies by Sabitov Zh. M. and Turuspe- public in Uzbekistan), and India.

Conclusion

To summarize, Chinggis Khan’s Y- Shibanids are absent elsewhere, including coun- chromosome is not as widespread as previously tries that are traditionally linked to the Mongol thought. The three remaining lineages of Ching- Empire, such as Afghanistan and Mongolia. Fi- gis Khan belong to three different subgroups of nally, contrary to previous studies by Zerjalet al, haplogroup C3. Although haplogroup C3 is the the star-cluster C3* is the modal haplotype of dominant haplogroup in both Kazakhstan and various Turko-Mongol tribes that are not directly Mongolia, Dayan Khan’s and Jani-Beg Khan’s related to Chinggis Khan. Therefore, the star- haplotypes each constitute less than 3.5% of the cluster C3* or Chinggis Khan’s hypothetical Y- Central Asian (i.e. Kazakhstan, Mongolia, and chromosome was either the dominant haplotype Uzbekistan) Y-chromosome gene pool. Out of all of most Mongolian men in the 13th century or it the haplogroup C3 DNA sequences, these two had been well distributed throughout Central haplotypes are the least distributed in before Chinggis Khan’s time due to the ter- Asia. The primary Mongolian Chinggisid lineage, ritorial expansions of various nomadic empires. or the Dayan Khanid aristocracy, possesses hap- logroup C3 (M407+). The divergence time be- tween Dayan Khan’s lineage and Jani-Beg Acknowledgements Khan’s lineage is more than 4,500 years for a generation time of 30 years. The third Chinggi- A special thanks to all our wonderful DNA sid branch belongs to the hypothetical haplotype donors; Sharaid Uuganbayar, Sartuul Tugulgur, of Chinggis Khan, the star-cluster C3*. Howev- and Khongorzul Enkh-Amgalan, for collecting the er, the number of the members of this particular DNA samples; and Mary Mills and Daniel Znider- branch is a lot scarcer than the previously men- zicfor editing the manuscript and giving incredi- tioned two major branches. The estimated num- bly valuable advice. ber of the Shibanids is less than a thousand al- together in Kazakhstan and Uzbekistan. The

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The Russian Journal of Genetic Genealogy (Русская версия): Том 4, №2, 2012 год ISSN: 1920-2997 http://ru.rjgg.org © Все права защищены RJGG

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