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91
Introduction
Yakutia, or the Sakha Republic of the Russian
Federation, spans an immense territory across the
Central and Northeastern Siberia, with a land mass
of about 3 million km (Figure 1). According to the
2010 census, among a total population of around
1 million people, Yakuts (or Sakhas) constitute
49.9%, hence the most populous ethnic entity in
the republic, followed by Russians (37.8%), Evenks
(2.2%), Ukrainians (2.2%), Evens (1.6%) and Tatars
(0.9%) [1].
Sakhas have a number of distinguishing ethno-
graphic features over the other Northeastern Si-
Abstract. Sakhas are Turkic-speaking people from Northeastern Siberia, constituting the largest ethnic
population in Yakutia. According to popular legends, two heroes who arrived from the Asian Steppe during
the late medieval ages, namely Elley Bootur and Omogoy Baay, are the progenitors of all Sakhas. While there
is ample historical evidence towards the existence of such legendary characters, archaeological ndings and
ancient DNA studies provide further insights on actual Sakha ethnogenesis. is study aims to establish the
genetic basis of the legendary characters Elley and Omogoy, at least through their paternal lineages, and
then to reveal the prevalence of these Y-chromosomes among the contemporary Yakut population. To this
end, an attempt was made to delineate fact from ction with respect to the Sakhas’ paternal lineages through
a reconciliation of population genetics data on contemporary and ancient Sakhas, along with archaeological
evidence and well-recorded historical narratives. To achieve this, 17-loci Y-chromosomal STR and hap-
logroup analyses were conducted on a contemporary Sakha who was presumably a direct descendant of
Elley’s paternal line. Furthermore, 367 Sakha Y-chromosomal STR haplotypes were compiled from the lit-
erature and elsewhere, and searched against the Y-chromosome STR Haplotype Reference Database to nd
potential matches with non-Sakha populations. Sakhas’ paternal lineages were found to comprise 6 major
descent clusters, each corresponding to an ancient clan. e most prevalent haplotype indeed corresponded
to that of the contemporary Elley descendant. Furthermore, data presented in the current work suggests a
Khitan origin for this paternal line. As shown before, Sakhas’ paternal lineages were found to be very ho-
mogenous and exhibit signs of a strong population bottleneck. Reconciled genetic and archaeological data
agree well with Sakhas’ historical narratives, whereby, at least from a paternal lineage perspective, only a few
individuals may have arrived from Central Asia and had reproductive success that led to the Sakha Y-chro-
mosomal diversity today.
Key words: Geneology, Elley and Omogoy, haplogroup N, founder eect, Xiongnu, autochthons.
For citations: Tikhonov D.G., Gurkan C., Demirdov K. D., Beyoglu E. On the origins of the Sakhas’
paternal lineages: Reconciliation of population genetic / ancient DNA data, archaeological ndings and
historical narratives // Siberian Research. 2019. 1(1). P. 91 -111. DOI: 10.33384/26587270.2019.01.004e
Received December 3, 2018; accepted for publication March 01, 2019; published April 15, 2019.
ETHNOGENESIS AND POPULATION GENETICS
On the origins of the Sakhas' paternal lineages:
Reconciliation of population genetic / ancient DNA data,
archaeological ndings and historical narratives
Tikhonov D.G., Gurkan C., Demirdov K.D., Beyoglu E
SIBERIAN RESEARCH | 1 01 | 2019
DOI: 10.33384/26587270.2019.01.004e
92
berian populations, such as their Turkic language,
a traditional economy based on semi-nomadic
horse- and cattle-breeding, and other customs that
bear closer resemblances with those from the Asian
Steppe instead. Yet, results from various population
genetics investigations of contemporary Sakhas
suggest a more complex picture. Sakhas’ maternal
lineages are heteregenous and have an admixed or-
igin, featuring close anities with both the autoch-
thonous Northeastern Siberian populations, such
as Evenks, who were traditionally Tungusic-speak-
ing reindeer-herders and hunter-gatherers, and
Southern Siberian Turkic-speaking Tuvans [2, 3].
In sharp contrast, Sakhas’ paternal lineages are very
homogenous, both geographically and chronolog-
ically, possibly due to a population bottleneck or
founder eect [3 – 6 ].
According to Sakhas’ historical narratives, two
legendary characters named Elley Bootur and Omo-
goy Baay arrived from the Asian Steppe during the
late middle ages and revolutionarily changed the
way of life in the middle Lena River. rough his
monumental work titled Elleyada, Gavriel V. Kse-
nofontov, a well-known Siberian social anthropol-
ogist and folklore specialist, reported the story of
Elley as it was vividly described in the Sakha oral
tradition [7]. Apparently, behind such a choice for
the title of his work lied not only an impression
with Homer’s Iliad and the Odyssey, but also a deep
fascination with the outcome of the excavations
by Schliemann, which had formally conrmed the
actual existence of otherwise mythical Troy and
Mycenae. In the preface of Elleyada, which could
only be published posthumously, the famous eth-
nographer Alexey P. Okladnikov highlighted that
‘e name in itself of a new publication on Yakut
folklore was reminiscent of Homer’s stories on the
Trojan War. Now all he needed was to nd his Troy
like Schliemann!’ [7].
Based on substantial historical evidence current-
ly available, as well as popular legends that are oen
based at least in part on actual historical events and
at times ornate with ctional anecdotes [8], Elley
and Omogoy were not only legendary characters
[9]. Some investigators hypothesize that Elley was
indeed a real person, perhaps from the Khitan Dy-
nasty [10, 11]. is study attempts to ascertain the
genetic evidence towards the existence of the leg-
endary Sakha characters such as Elley and Omogoy
through the use of Y-chromosomal short tandem
repeat (Y-STR) and haplogroup data on contempo-
rary and ancient Sakhas, along with that of a con-
temporary descendant of Elley’s paternal lineage
according to the historical evidence available.
Fig 1. A map showing the Russian Federation wherein the Sakha Republic and its capital Yakutsk,
the Lena River and Lake Baikal are highlighted.
TIKHONOV D.G., GURKAN C., DEMIRDOV K. D., BEYOGLU E
93
Zerjal et al. [12] were the rst to describe a
Y-chromosomal base substation comprising a TC
transtition at the RBF5 locus, subsequently named
as the Tat C allele, at a very high-level among the
Yakut population (86% of those tested). ese au-
thors also conducted 9-loci Y-STR analysis on the
individuals bearing the Tat C allele, and were the
rst to reveal the presence of three main haplo-
types among the Yakuts. rough Y-chromosomal
SNP and STR (9-loci) analyses, Pakendorf et al. [5]
also showed the presence of a strong founder-eect
among the paternal lineages of the Yakut popula-
tion. ese ndings were all later further conrmed
through the archaeogenetic studies, such as by 17-
loci Y-STR analysis [6, 13]. Yet, despite all of these
ndings of immense signicance, no attempt has
so for been made to link the key Y-STR haplotypes
with perhaps known historical personalities.
In the current study, an attempt is thus made for
the very rst time to establish the genetic aliation
of the main paternal lineages observed among the
Yakut population, such as with the legendary Sakha
characters of Elley and Omogoy, through the use of
Y-STR and haplogroup data on contemporary and
ancient Sakhas, along with a putative direct descen-
dant of Elley’s paternal lineage. Establishment of
Fig. 2. Megino-Khangalassky branch of the Elley pedigree. Arrow indicates the actual individual sampled
for the Y-chromosomal STR and haplogroup analyses.
ON THE ORIGINS OF THE SAKHAS' PATERNAL LINEAGES
94
these genetic connections would allow us to help
clarify some important and hitherto unclear as-
pects on the Yakut ethnogenesis.
Materials and Methods
A putative pedigree focusing on the paternal
lineage of the Megino-Khangalassky Ulus of the
Elley clan, starting with the progenitor Elley was
constructed (Figure 2). e genealogical data were
derived from various sources in the literature, his-
torical documents and research from the current
study: the immediate descendents of Elley were rst
described in the 18 century [14], from Bolotoi to
Mengebil Byugyu, and further on to Kirtey (Kintey)
and then Khisirges was according to anthropolog-
ical literature [15, 16], from Khisirges to Oyunnut,
and further on to yiachan was according to an-
thropological literature and archived documents
[15, 17], and nally from yiachan down to the
last four generations (for whom only the initials for
the rst names are provided) was according to re-
search from the current study.
A contemporary representative of Elley’s pater-
nal lineage corresponding to a 59 year old Sakha was
identied in the current study (Figure 2) and a buc-
cal swab sample was collected along with informed
consent and in full accordance with the principles
of the Declaration of Helsinki by the World Medical
Association. 17 Y-chromosomal STR loci analysis
(DYS456, DYS389I, DYS390, DYS389II, DYS458,
DYS19, DYS385a/b, DYS393, DYS391, DYS439,
DYS635, DYS392, Y-GATA-H4, DYS437, DYS438,
and DYS448; the Life Technologies AmpFLSTR®
YlerTM Kit) and Y-chromosomal SNP-based hap-
logroup assignment were carried out at the Gentis
Laboratory (Moscow, e Russian Federation).
Table 1 provides the sources for the 367 contem-
porary Sakha Y-chromosomal STR haplotypes com-
piled from the literature, YHRD, and the current
study [5, 13, 18 – 22]. Table 2 lists the corresponding
nine-loci Sakha Y-chromosomal STR haplotypes,
their frequencies and the associated Y-chromo-
somal haplogroup assignments. When Y-chromo-
somal SNP based haplogroup assignments were not
readily available, the online 21-haplogroup version
of the Whit Athey in silico haplogroup assignment
algorithm was used instead (http://www.hprg.com/
hapest5/index.html) [21].
Each Y-chromosomal STR haplotype listed in
Table 2 were queried against Y chromosome STR
Haplotype Reference Database (YHRD) for po-
tential matches with those from non-Sakha popu-
lations using the ‘search the database’ option [21].
Accordingly, Table 3 lists the 17-loci Y-chromo-
somal STR haplotypes corresponding to the three
most prevalent Sakha haplotypes observed in the
current study and their near-perfect matches with
Population Data Number of haplotypes Source or YHRD Accession No.:
Sakha (Elley’s line) 1 e present study
Sakha 4 [18]
Sakha 4 [19]
Sakha 8 [22]
Sakha 21 [20]
Sakha 24 YA004128 & YA00397
Sakha 133 [13]
Sakha 172 [5]
Tota l 367
Table 1. Sakha Y-chromosomal STR haplotypes used in the current study
TIKHONOV D.G., GURKAN C., DEMIRDOV K. D., BEYOGLU E
95
those from non-Sakha population datasets.
Median-joining network analyses were carried
out with the full 17-loci (Yler), 14-loci [Yler, mi-
nus DYS385a, DYS386b and DYS390], and nine-loci
[DYS389I, DYS389II, DYS19, DYS385a, DYS385b,
DYS390, DYS391, DYS392 and DYS393] Y-chro-
mosomal STR datasets using the Network v.5.0.0.0
soware (www.uxusengineering.com). Either the
whole network or a specic descent cluster in a giv-
en network was analyzed with the Network Time
Estimates sub-program of the same soware by se-
lecting a proposed central ancestral node and then
all the other descendant nodes. Y-chromosom-
al STR mutation rates of 0.003257, 0.003365 and
0.003307 per locus per generation were used for
the nine-, 14- and 17-loci median-joining network
analyses, in that order, and along with a generation
time of 32 years [23].
Results
Among the 367 nine-loci Y-chromosomal STR
haplotypes corresponding to the contemporary
Sakhas (Table 2), 66 dierent haplotypes were ob-
served, thus aording a discrimination capacity of
17.9%, as well as only 35 unique haplotypes (9.5%).
ese results are in line with earlier reports on the
high homogeneity of the Sakha paternal lineages
[4 – 6]. e contemporary representative of Elley’s
paternal lineage (Ellyayevsky family), a 59 years old
Fig. 3 Nine-loci median-joining network analysis of Sakha Y-chromosomal STR haplotypes belonging to
the N haplogroup.
Each haplotype is denoted by a yellow circle, except for the proposed ancestral nodes for each of the ve major
descent clusters that are shown in dierent colours according to the legend inset provided. e size of each the
node is indicative of the number of haplotypes represented. Median vectors are shown as tiny red circles.
ON THE ORIGINS OF THE SAKHAS' PATERNAL LINEAGES
96
Fig. 4. e distribution of the Sakha Y-chromosomal STR haplotype clusters/groups
male individual (M.) (Figure 2), was found to have
a Y-chromosomal haplogroup assignment of N1c1
[dened by the single nucleotide polymorphism
(SNP) M178+] and a full 17-loci Y-chromosomal
STR haplotype as shown in Table 3. At nine loci res-
olution, this haplotype corresponded to the most
prevalent haplotype (37.3%) among all Sakhas. Tak-
ing into consideration the fact that the genealogical
paternal lineage of the male M. tested in the current
study presumably represented that of the legendary
character Elley, it was classied as ‘the original El-
ley line’. Once again, all generations of the paternal
lineage of the male individual M. were documented
and conrmed by archival data and publications.
e rst information on the genealogy of the El-
ley family was described by the participant of the
Academic Detachment of the Second Kamchatka
Expedition (Russia) Yakov Lindenau in 1741-1745.
e second most prevalent haplotype (16.9%) [13]
with unknown origin, was classied as “dominant
line Unknown”. e third most prevalent haplotype
(13.9%) was supposedly attributed to Omogoy ge-
nus by Adamov [24]. Determination of the Y chro-
mosomal STR haplotype to the mummied frozen
body (ID: YAKa 69) from the Kous Tcharbyt site,
conducted by Crubezy et al. [6] coincides with the
third common haplotype of Sakha. e excavated
thomb at the Kous Tcharbyt site, was identied as
that of a tojon (clan chief) and dated to the 16
Century/early 17 Century CE, presumably from
the Bayaginskii family. According to Aprosimov
and Popov [15], Bayaginskii nasleg (sub-district)
belonged to the Omogoisky family, thus this hap-
lotype was classied as ‘the dominant Omogoy
line’. Figure 3 depicts the median-joining network
analysis for the nine-loci Y-chromosomal STR hap-
lotypes belonging to the N haplogroup among the
contemporary Sakhas.
As a direct comparison of the contemporary
and ancient Sakha populations, out of the 27 suc-
cessful Y-chromosomal STR proles that could be
obtained from mummied frozen bodies belonging
to Sakhas from the 15 to 19 centuries, the two
most prevalent haplotypes that were observed eight
(29.6%) and seven (25.9%) times corresponded to
‘the original Elley line’ and the ‘dominant Omogoy
line’ from the current study, respectively [6]. When
the ancient DNA dataset was eventually increased
to comprise data from 62 ancient Sakhas, the 17-
loci Y-chromosomal STR haplotype corresponding
to that of ‘the original Elley line’ from the current
study was found to correspond to as high as 46.8%
of all the male lineages analyzed [27].
TIKHONOV D.G., GURKAN C., DEMIRDOV K. D., BEYOGLU E
97
Haplotype ID
DYS389 I
DYS390
DYS389 II
DYS19
DYS385a/b
DYS393
DYS391
DYS392
Number of
Haplotypes
Haplogroup*
Source
Elley Clan - - - - - - - 173 N
«the original
Elley line» 14 23 32 14 11, 13 14 11 16 137 N1c1,
1–8
E1 14 23 32 14 11, 13 14 10 16 4 N1c1, N 4, 6
E2 14 23 32 14 12, 13 14 11 16 3 N1c14, 5
E4 14 23 32 14 11, 13 14 12 16 3 N 6
E5 14 23 32 15 11, 13 14 11 16 3 N1c14, 6
E6 14 24 32 14 11, 13 14 11 16 2 N1c1,
N1c1al, 1, 3
E3 14 23 32 14 11, 12 14 11 16 2 N1c14
E13 14 23 32 14 11, 13 14 11 17 2 N1c11, 4
E8 14 23 32 14 11, 13 13 11 16 1 N 6
E9 14 22 32 14 11, 13 14 11 16 1 N1c14
E10 14 23 32 14 11, 14 14 11 16 1 N1c14
E14 14 23 32 14 11, 13 14 11 14 1 N1c14
A2 14 23 32 15 11, 12 14 11 16 3 N 6
A7 14 23 33 14 11, 13 14 11 16 3 N1c1, N 4, 6
A12 15 23 33 14 11, 13 14 11 16 2 N1c14, 5
A4 13 23 31 14 11, 14 14 11 16 2 N1c14
A14 14 23 32 14 11, 12 14 12 16 1 N1c14
A17 14 23 32 14 11, 13 12 10 16 1 N 6
A19 14 23 33 14 12, 13 14 11 16 1 N 6
Unknown Clan - - - - - - - 82 N
«the dominant
Unknown line» 14 23 31 14 11, 13 14 11 16 62 N1c1, N 4, 6,
8
Table 2. Frequencies of the 9-loci Y-STR haplotypes observed in the Sakha population
ON THE ORIGINS OF THE SAKHAS' PATERNAL LINEAGES
98
B1 14 23 31 14 11, 13 14 10 16 5 N1c1, N 1, 4,
6, 7
B2 14 24 31 14 11, 13 14 11 16 3 N1c14
B3 14 23 31 14 11, 13 13 11 16 3 N1c1, N 4, 6
B5 13 23 31 14 11, 14 14 11 16 2 N1c14
B4 14 23 31 14 11, 12 14 11 16 1 N1c14
B6 14 23 31 14 11, 13 14 12 16 1 N 6
A 14 23 30 14 11, 13 14 11 16 5 N1c1, N 2, 4,
6
Omogoy Clan - - - - - - - 71 N
«the dominant
Omogoy line» 14 23 31 14 11, 13 14 11 15 51 N1c1, N 1, 4,
6, 8
Om1 14 23 31 14 12, 13 14 11 15 4 N1c1, N 4, 6
Om2 14 23 31 14 11, 13 14 10 15 3 N1c14
Om3 14 23 31 14 11, 13 13 11 15 2 N1c14
Om4 14 23 31 14 11, 14 14 11 15 2 N 6
Om5 14 23 31 15 11, 13 14 11 15 1 N 6
A3 13 23 30 14 11, 13 14 11 15 3 N1c1, N 4, 6
A6 14 23 30 14 11, 13 14 11 15 2 N 6
A13 15 23 32 14 11, 13 14 11 15 1 N1c14
A15 15 23 32 15 11, 13 13 11 15 1 N1c14
A22 13 23 30 14 11, 14 14 11 15 1 N 6
Eurasian Clan - - - - - - - 10 N
«the dominant
Eurasian line» 14 23 30 14 11, 13 14 10 14 7 N 6, 7
A5 14 23 30 14 11, 13 14 10 15 2 N1c14
A16 14 22 30 14 11, 13 14 10 14 1 N1c14
ontinuation of table 2
TIKHONOV D.G., GURKAN C., DEMIRDOV K. D., BEYOGLU E
99
Xioungnu line - - - - - - - 4 N
Xuo 13 23 29 14 11, 13 14 11 14 1 N 6
Xuo1 14 23 30 14 11, 13 14 11 14 1 N 6
Xuo3 13 24 29 14 11, 13 14 11 14 1 N 7
A35 13 23 29 14 11, 13 15 11 14 1 N1c14
Admixed - - - - - - - 10 various
A8 14 22 30 15 9, 19 13 10 13 2 L 6
A9 13 24 30 16 14, 15 13 11 11 2 12a (xl2a1) 6
A21 12 23 28 14 14, 15 13 10 11 1 l1 6
A28 13 25 31 16 11, 14 13 11 11 1 R1a 6
A29 13 25 29 16 12, 13 13 10 11 1 l2a1 6
A30 13 25 29 16 12, 13 13 10 12 1 l2a (xl2a1) 6
A31 13 25 30 16 11, 15 13 12 11 1 R1a 6
A36 13 23 28 15 11, 19 14 10 11 1 Jl 8
Autochtons - - - - - - - 17 various
A1 10 24 27 15 11, 12 13 9 11 4 E1blb 6
A11 10 24 27 15 11, 12 13 9 10 2 E1blb 6
A10 14 23 30 17 11, 18 13 10 11 1 l2a (xl2a1) 2
A18 14 23 32 14 13, 19 14 11 16 1 Q 6
A1 10 24 27 15 11, 12 13 9 11 1 E1blb 6
A20 12 23 28 14 12, 13 13 10 14 1 N 6
A23 13 23 28 15 11, 17 16 10 12 1 G2a 6
A24 13 23 31 15 11, 16 13 11 11 1 R1a 6
A25 10 24 27 15 12, 12 13 9 7 1 l2a (xl2a1) 6
A26 10 24 27 15 12, 19 13 9 11 1 E1blb 6
ontinuation of table 2
ON THE ORIGINS OF THE SAKHAS' PATERNAL LINEAGES
100
A27 10 24 27 15 12, 13 13 9 10 1 G2a 6
A32 13 23 30 14 13, 13 13 10 14 1 N2a1a (N1b) 3
A34 13 24 30 15 15, 15 13 11 11 1 l2a (xl2a1) 2
Tota l 367
* Table 1 in the Supporting information le contains the parallel haplogroup assignments by the V. Urasin’s Pedictor
and that by D. Adamov.
a Haplotype observed in the current study, the putative Elley line M., XVII:1 (Fig. 2).
b Haplogroup assignments made with the Whit Athey algorithm.
с the name of the haplogroup according to ISOGG 2016
‡ Alleles shown in italics correspond to those that dier from the original/dominant haplotype in the same cluster/
group by only a few allelic repeat values.
Table 3. 17-loci Y-STR near-matches at YHRD between the original/dominant Elley, Unknown and
Omogoy haplotypes and non-Sakha haplotypes (excluding the Athabaskan Indian haplotype).
Notably, the following haplotype pairs have the same 17-loci Y-STR prole:
Mg4 and Mg5; LiMo and ShC1.
ID Haplotype *
DYS456
DYS389 I
DYS390
DYS389 II
DYS458
DYS19
DYS385a/b
DYS393
DYS391
DYS439
DYS635
DYS392
GATA_H4
DYS437
DYS438
DYS448
YHRD
Accession
No.:
Ell0 the original
Elley line 14 14 23 32 16 14 11,
13 14 11 10 22 16 12 14 11 19 Current
study
Vil the dominant
Unknown line 14 14 23 31 16 14 11,
13 14 11 10 22 16 12 14 11 19 [13]
Omo1 the dominant
Omogoy line 14 14 23 31 16 14 11,
13 14 11 10 22 15 12 14 11 19 [6]
TCy1 Turkish Cypriot 15 14 23 32 17 14 11,
13 14 11 10 22 16 12 14 11 19 YA003850
LiKo Liaoning,
China (Korean) 14 14 23 32 17 14 11,
13 14 11 10 22 16 12 14 11 20 YA003759
Afg Afghanistan
(Afghan) 14 14 23 32 16 14 10,
13 14 10 10 22 16 12 14 11 19 YA003842
JilH Julin, Chian
(Han) 14 14 23 30 16 14 11,
13 14 11 10 22 16 12 14 11 19 YA004124
LiMo
Liaoning,
China
(Mongolian)
14 14 23 30 17 14 11,
13 14 11 10 22 16 12 14 11 19 YA003758
ontinuation of table 2
TIKHONOV D.G., GURKAN C., DEMIRDOV K. D., BEYOGLU E
101
ShC1 Shandong,
China (Han) 14 14 23 30 17 14 11,
13 14 11 10 22 16 12 14 11 19 YA004081
SSK1 Seoul, South
Korea (Korean) 14 14 23 30 16 14 11,
13 14 11 10 21 16 12 14 11 19 YC000251
SSK2 Seoul, South
Korea (Korean) 14 14 23 31 17 14 11,
13 14 11 10 22 16 12 14 11 20 YC000251
SKK South Korea
(Korean) 14 14 23 31 17 14 11,
13 14 11 10 22 16 12 14 11 20
YA003728,
YC000239,
YA003441
Mg1
Ulaanbaatar,
Mongolia
(Khalkh)
14 14 23 31 17 14 10,
13 14 11 10 22 16 12 14 11 19 YA003733
Mg4
Northern Mon-
golia (Khalkh,
Darkhad, Uri-
ankhai)
14 14 23 31 17 14 11,
13 14 11 10 22 16 12 14 11 19 YA003734
Mg5
Ulaanbaatar,
Mongolia
(Khalkh)
14 14 23 31 17 14 11,
13 14 11 10 22 16 12 14 11 19 YA003733
LiHa
Liaoning,
China
(Northern Han)
14 14 23 31 17 14 11,
13 13 11 10 22 16 12 14 11 19 YA003756
ChHa
Changchun,
Jilin, China
(Han)
14 14 23 31 16 14 11,
13 15 11 10 22 15 12 14 11 19 [31]
a - Haplotype observed in the current study, the putative Elley line M., XV:1 (g. 2).
* - Allelic values shown in italics denote allelic repeat variations from ‘the original Elley line’
ontinuation of table 3
Based on Y-chromosomal SNP or in silico hap-
logroup assignments (Table 2), nearly all Y-chro-
mosomal STR haplotypes in the Elley, Unknown,
Omogoy, Eurasian and Xiongou descent clusters
exclusively comprised haplogroup N (92.7% of all
haplotypes), while those from the ‘Admixed’ and
‘Autochthons’ groups exhibited more heterogenous
haplogroup distributions. Table 4 shows the ‘Time
to Most Recent Common Ancestor’ or TMRCA
estimates for each of the ve major Sakha clans
classied based on the descent clusters described
in Table 2 and Figure 3. In close agreement with
the hypotheses of the current study, the Xiongnu
clan seems to be the oldest, followed by Omogoy,
Elley, Unknown and Eurasian clans, in that order.
However, it should be noted that, TMRCA calcula-
tions can only be rough estimates due to the uncer-
tainty with respect to the precise Y-chromosomal
STR mutation rate(s) that should be used during
such calculations [28]. When each of the dierent
nine-loci Y-chromosomal STR haplotypes in Ta-
ble 2 were queried against YHRD, the following
major observations could be made: (i) haplotypes
associated with the Elley, Omogoy, Unknown and
Autochtons descent clusters had 20, seven, three
and two matches in total and in that order (2.5%
of the total matches) with the population datasets
corresponding to other ethnic groups from Yakutia,
namely Evenks and Yakut-speaking Evenks, (ii) the
haplotypes associated with the Elley, Omogoy and
ON THE ORIGINS OF THE SAKHAS' PATERNAL LINEAGES
102
Sakha Clan(s) TMRCA (in years) SD (in years) Time period (Mean)
Elley 283 77 1656 - 1807 CE (1733)
Unknown 271 117 1628 - 1862 CE (1745)
Omogoy 360 121 1535 - 1777 CE (1656)
Elley / Unknown /
Omogoy 767 166 1083 - 1465 CE (1249)
Eurasian 334 249 1433 - 1931 CE (1682)
Xiongnu 1334 588 94 - 1270 CE (682)
Table 4. TMRCA analyses for the nine-loci Sakha Y-chromosomal STR clusters
Unknown descent clusters had 4, eleven and twen-
ty ve matches in total and in that order (2.9% of
the total matches) with outside-Yakutia population
datasets, largely in the Eurasian context, (iii) vast
majority of haplotype matches (94.7% of the total
matches) were associated with the Eurasian, Xiong-
nu and Admixed/Autochthons descent clusters/
groups and the outside-Yakutia population data-
sets, and (iv) 67.0% of the haplotype matches with
the outside-Yakutia population datasets were asso-
ciated with the Admixed and Autochthons groups,
where the matches were either over the entire 17-
loci or had dierences at only a few loci by one-to-
two allelic repeat(s).
Notably, the original Elley line and the domi-
nant Omogoy line haplotypes themselves merely
diered by a single allelic repeat at two loci, namely
DYS389II and DYS392 (Tables 2 and 3). In turn, the
dominant Unknown line haplotype diered only by
a single allelic repeat at one locus each with those
from the Omogoy and Elley lines at DYS389II and
DYS392, respectively (Tables 2 and 3). Upon que-
rying the 17-loci Y-chromosomal STR haplotypes
corresponding to ‘the original Elley line’ and ‘the
dominant Unknown’ and ‘the dominant Omogoy
lines’ against YHRD, 14 near-perfect matches were
observed in total, each of which diered from the
queried haplotypes by single allelic value dier-
ences at one-to-four dierent loci and largely orig-
inating from populations spanning a wide range of
Eurasian geography (Table 3). One further match
corresponded to a haplotype from the Athabaskan
Indian dataset from Central Alaska, U.S.A. (YHRD
Accession No.: YA003683), which had a single al-
lelic repeat dierence at only one locus from ‘the
original Elley line’, hence bringing about the possi-
bility for the former to be also a descendant of the
Elley family.
Figure 5 depicts the 17-loci median-joining net-
work for the 14 near-perfect YHRD matches with
the Y-chromosomal STR haplotypes corresponding
to the original Elley, the dominant Unknown and
Omogoy lines, and those from the outside-Yakutia
population datasets. TMRCA is estimated at 1127 ±
317 years or 889 ± 317 CE, covering the period of
572 – 1206 CE, which coincides with the rise and
fall of Turkic, Uyghur and Rouran Khaganates.
When only 9-loci Y-chromosomal STR hap-
lotypes corresponding to the original Elley and
the dominant Omogoy lines were queried against
YHRD, the original Elley line haplotype had perfect
matches with two non-Sakha haplotypes, namely
one from the Turkish Cypriot and another from
the Korean minority in Liaoning, China datasets
(haplotype IDs TCy1 and LiKo, respectively, in Ta-
ble 3) [29, 30], while the dominant Omogoy line
had none. In comparison, among the 13 dierent
nine-loci Y-chromosomal STR haplotypes in the
Autochthons group, only three had 19 matches in
total with the outside-Yakutia datasets at YHRD
(Table 2). Notably, one of these Sakha haplotypes
(A20) had nine such matches in total: three with
the Central Anatolia, Turkey (YHRD Accession
No.: YP000080), three with the Zhejiang, Chi-
TIKHONOV D.G., GURKAN C., DEMIRDOV K. D., BEYOGLU E
103
Fig. 5. 17-loci median-joining network analysis of the original/dominant Elley, Unknown and Omogoy
Y-chromosomal STR haplotypes with the YHRD matches from outside Yakutia populations.
Each haplotype is denoted by a yellow circle, except for the Elley, Unknown and Omogoy haplotypes shown
in dierent colored circles according to the legend provided. e size of each circle is indicative of the number
of haplotypes represented. Median vectors are shown as small black circles and the proposed ancestral node is
marked with *. Refer to Table 4 for the haplotype IDs, etc.
na [Han] (YHRD Accession No.: YP000506), two
with the Phillipines [Fillipino] (YHRD Accession
No.: YP000229) and one with the Taiwan [Paiwan]
(YHRD Accession No.: YP000552) datasets. YHRD
matches between the two other nine-loci Y-chro-
mosomal STR haplotypes from the Autochthons
group and outside-Yakutia datasets corresponded
to those from U.S.A., ailand, China and East-
ern European countries. When repeated at 17-loci,
none of the 13 Y-chromosomal STR haplotypes
from the Autochthons group had any matches at
YHRD with the outside-Yakutia datasets.
Discussion
Yakut ethnogenesis has been investigated by sev-
eral generations of scientists. ese investigations
comprised ethnographic, archaeological, anthro-
pological, and genetic studies, as well as those on
comparative linguistics and folklore of Sakha. Nev-
ertheless, up until now, these studies have not al-
lowed us to fully understand many aspects of the
Yakut ethnogenesis. A breakthrough in the study
of this issue was made with the discovery of the
Kulun-Atakh culture (13-15 centuries CE) [32].
A.I. Gogolev was the rst to prove the emergence of
Sakha as a nationality in the territory of the middle
Lena, and his work still remains as the only scientif-
ic work that comprehensively analyzed the ethno-
genesis of the Yakuts using a wide range of scientic
disciplines: historical, cultural, linguistic, archaeo-
logical and anthropological [32]. Notably, the au-
thor nally refuted the possibility for the migration
of Sakha from the south to an already formed eth-
ON THE ORIGINS OF THE SAKHAS' PATERNAL LINEAGES
104
nos in Middle Lena. Yet, population genetic analy-
ses still remain a discipline not reconciled with the
established facts of Yakut ethnogenesis. is is per-
haps partly due to the fact that genetic studies have
begun to be involved in the study of the ethnogene-
sis of the Yakuts only relatively recently [6, 33].
In the present work, an attempt is made to uti-
lize the accumulated Y-chromosomal data that has
already been in use in population and forensic ge-
netics to help identify the paternal ancestors of the
Yakut people. e solution of these questions is of
great importance in understanding the ethnogene-
sis of the Sakha. Each population is heterogeneous
in a genetic sense, and the elucidation of the pop-
ulation genetic structure can help determine the
precise origins and potential contributors in a giv-
en population. According to the views of a number
of authoritative researchers, the Yakut ethnos was
formed in the territory of Yakutia as a result of the
mixing of people from the south and the autoch-
thonous population [34].
Starting with Zerjal et al. [12], a number of stud-
ies on the Yakut Y-STR haplotypes were carried out,
but none of these studies can cover a truly repre-
sentative diversity of the haplotypes of the studied
population. us, in the current work, an attempt
was made to compile from all these sources 367
9-loci Y-STR haplotype data in total (see Table 1),
currently the most complete collection of such data
on the Sakha population. Such an approach enabled
the grouping of similar Sakha haplotypes in an at-
tempt to identify their ancestral paternal lineages
(see Table 2).
e relationship among the Sakha Y-STR hap-
lotypes belonging to the haplogroup N are now
shown by a median-joining network analysis (Fig-
ure 3). e presence of three, closely related star-
shaped clusters is immediately notable in the net-
work. Similar structures in phylogenetic patterns
reect the facts of the long historical isolation of
relatively small groups of the population. In addi-
tion, two small centers of divergence are noted in
the peripheral part of the star-shaped structures.
Data from the current study suggest that the Ya-
kut modal Y-STR haplotype, which belonged to the
haplogroup N1c1, could be that of the legendary
character Elley. e remaining four major clusters
could also be classied in line with the anthropo-
logical/archaeological/archaeogenetic data avail-
able in the literature. Yet, even prior to that, each
Y-STR haplotype could be classied as mixed and
autochthons based on the fact that whether match-
ing haploptypes could be observed or not, respec-
tively, with the populations outside the territory of
modern Yakutia.
In forensic genetics, 17-loci Y-STR haplotypes
analyses are commonly used to identify the pa-
ternal lineage of a given individual. In an eort to
help identify similar Y-STR haplotypes to the three
main Yakut haplotypes, the search function of the
YHRD database was used. Table 3 lists the 14-loci
haplotypes that are presumably related to these
three major Yakut haplotypes, which were then an-
alyzed through the construction of median-joining
networks (Figure 5). According to the results pre-
sented here, very similar Y-STR haplotypes to that
of the original Elley line were found in the west:
Afghanistan and northern Cyprus, and in the east:
Liaoning Province, China and Ulaanbaator, North-
ern Mongolia. In the case of the dominant Omo-
goy line, very closely matching haplotypes diering
by a single mutational step were found in the city
of Chifen of the Jirin Province, China. e widest
range of similar haplotypes was found for the Ya-
kut haplotype Unknown: In Mongolia, China and
South Korea. For instance, haplotypes diering by a
single step mutation were found in Northern Mon-
golia (Khalk, Darhad, Uryankhai populations),
Ulaanbaator (Khalk) and in the province of Jirin,
China (Han population).
Notably, Tat-C-bearing Y-chromosomes were
also observed in ancient DNA samples from the
2700-3000 years-old Upper Xiajiadian culture in
Inner Mongolia, as well as those from the Serteya
II site at the Upper Dvina region in Russia and the
‘Devichyi gory’ culture of long barrow burials at the
Nevel’sky district of Pskovsky region in Russia [35,
36]. A 14-loci Y-chromosomal STR median-join-
ing network of the most prevalent Sakha haplo-
types and a Tat-C-bearing haplotype from one of
the ancient DNA samples recovered from the Up-
per Xiajiadian culture in Inner Mongolia (DSQ04)
revealed that the contemporary Sakha haplotype
‘Xuo’ (Table 2, Haplotype ID “Xuo”) classied as
TIKHONOV D.G., GURKAN C., DEMIRDOV K. D., BEYOGLU E
105
that of ‘the Xiongnu clan’ in our current study, was
the closest to the ancient Xiongnu haplotype (Fig-
ure 6). TMRCA estimate for this 14-loci Y-chromo-
somal STR network was 4357 ± 1038 years or 2341
± 1038 BCE, which correlated well with the Upper
Xiajiadian culture that was dated to the Late Bronze
Age (700-1000 BCE).
e three most prevalent 17-loci Y-chromosom-
al STR haplotypes observed among contemporary
Sakhas (79% in total), namely ‘the original Elley
line’, ‘the dominant Unknown line’ and ‘the dom-
inant Omogoy line’ themselves seem to be closely
related aer all (Table 3). is is because from ‘the
original Elley line’ to ‘the dominant Omogoy line’,
through the apparent intermediary of ‘the domi-
nant Unknown line’, there are only two incremen-
tal one-step mutations at only two loci, namely
DYS389II and DYS19, or vice versa. A TMRCA esti-
mate for the corresponding descent clusters around
these three haplotypes (Figure 3) was calculated as
767 ± 166 years or 1083 - 1415 CE (Table 4).
ese three major Sakha paternal lineages may
have also arrived in Yakutia at dierent times and/
or from dierent places and/or with a dierence in
several generations instead, or perhaps Y-chromo-
somal STR mutations may have taken place in situ
in Yakutia. Nevertheless, the immediate common
ancestor(s) from the Asian Steppe of these three
most prevalent Sakha Y-chromosomal STR haplo-
Fig. 6. 14-loci median-joining network analysis for the original/dominant Elley (Ell), Unknown Clan
(Vil), Omogoy (Omo), Eurasian (Eur) and Xiongnu (Xuo) Y-chromosomal STR haplotypes and that
for a representative ancient DNA sample (Ch0 or DSQ04) from the Upper Xiajiadian Culture
recovered from the Inner Mongolia Autonomous Region, China.
Each haplotype is denoted by a yellow circle, while the median vectors are shown as small red diamonds. e
proposed ancestral node is marked with *. A scale bar whose length is roughly proportional to a single mutation
event between any of the two neighboring haplotypes is also provided.
types possibly lived during the prominence of the
Turkic Khaganates, hence the near-perfect matches
observed across a wide range of Eurasian geogra-
phy, including as far as from Cyprus in the West
to Liaoning, China in the East, then Middle Lena
in the North and Afghanistan in the South (Ta-
ble 3 and Figure 5). ere may also be haplotypes
closely-related to ‘the dominant Elley line’ among
Karakalpaks, Uzbeks and Tajiks, however, limita-
tions in the loci coverage for the available dataset
(only eight Y-chromosomal STR loci) precludes
further conclusions on this matter [25].
A potential focus for the Westward prolifera-
tion of ‘the original Elley line’ could be the Zhetysu
territory in Central Asia. Aer the fall of the Liao
Dynasty in 1125 CE following the Jurchen invasion,
Elyui Dashi’s together with 20 thousand warriors
of the Chotan fortress, ed from the Jurchen and
westward to establish the Qara Khitai Khaganate or
Western Liao Empire in the Zhetysu territory [37].
Apparently, with the descendants of this army, the
related Elley haplotype reached Afghanistan and
Cyprus. In Balaresque P. et al. [25] we found poly-
morphisms close to Elley haplotype in Karakalpaks,
Uzbeks and Tajiks. Liao Empire of the Elyui Dynas-
ty lasted from 916 to 1125 CE, mentions about Kh-
itan people in the Chinese chronicles gradually dis-
appear by the time of the Ming Dynasty (1638-44
CE). Ancient DNA studies based on mitochondrial
ON THE ORIGINS OF THE SAKHAS' PATERNAL LINEAGES
106
DNA on the remains belonging to the members of
the Elyui Dynasty of the Liao Empire near towns
Chifens in Inner Mongolia and Fuxin in the Liaon-
ing province of China revealed that Khitans were
genetically closest to Daurs, Northern Chinese and
Mongols, and also bore similarities to Orochons,
Buryats and Evenks [38]. Daurs are considered to
be direct descendants of Khitans, and the former
were also found to have paternal lineages belonging
to the N1c haplogroup at a frequency of 7.7% [39].
Although nuclear DNA has so far not been success-
fully isolated from the burials of the Khitan nobil-
ity, data from contemporary populations in Liaon-
ing already allows us to postulate a potential Khitan
origin for the most prevalent Y-chromosomal STR
haplotypes among Sakhas (Table 3).
In one way or the other, the arrival of perhaps
only a few people from the Asian Steppe in Yakutia
between the 12 and 14 centuries CE coincides
with the introduction hitherto unknown technol-
ogies and cultural traditions to Northeastern Sibe-
ria, which ranged from advanced animal husband-
ry and milk production (including preparation
and consumption of koumiss, a form of fermented
product made with mare’s milk), transition to a
semi-nomadic lifestyle and formation of farms, new
house building technologies and introduction of
new household equipments [7]. ese revolution-
ary changes in Yakutia coincided with emergence
of the Kulun-Atakhskaya Culture of Sakhas around
the 13 - 14 centuries CE [32].
A novel Y-chromosomal STR haplotype that was
observed at a high frequency across Eurasia cor-
responds to those Y-chromosomal lineages again
belonging to haplogroup N1 [25]. Balarsque et
al. concluded that this high frequency haplotype
spread along the Silk Road in around 700 CE and
corresponded to the reproductive success of men
from the Rouran and Uighur Khaganates, as well as
the West Liao Empire of the Khitan Dynasty. Sim-
ilar haplotypes were also observed among contem-
porary Sakhas and classied under ‘the Eurasian
Clan’ (Table 2), whose descent cluster (Figure 3)
was dated as 334 ± 249 years old and correspond-
ed to a period between 1433 - 1931 CE (Table 4).
is brings about the possibility for the prolifera-
tion of these haplotypes as a result of the 200 years
old tea trade between Russia and China. e Tea
Trade Route stretched from Hankow, China and
passed through Mongolia, then Kyakhta in modern
day Buryatia, and nally reached Moscow [40]. A
Northern branch of the Tea Route also stretched
from Kyakhta to Yakutsk. Support for this theory
comes from the observation of 35 perfect and near
perfect-matches at YHRD (diering by only one
allelic dierence at a single locus of 17-loci haplo-
types) from China, Mongolia and Russia (at central
Russian cities of Sverdlovsk, Bryansk and Vologda),
and as far as Switzerland, Poland, Norway and Fin-
land, in a way recapitulating the tea trading path.
A potential connection between Sakhas and
Xiongnu culture was also observed, albeit in the
maternal lineages instead, whereby the mtDNA
hypervariable sequence 1 of a women from grave
25A from the Egyin Gol necropolis was observed in
contemporary Sakhas, as well as in an ancient DNA
sample belonging to a male individual recovered
from the 2200-2400-years-old Pokrovsk grave site
in Yakutia [41, 42].
Another line of evidence towards potential an-
cestral links between Xiongnu and Sakhas came
from 3D geometric-morphometric analyses based
on 44 craniofacial biometric points on 1558 skulls
from the collections of various museums from
around the world, which included 68 Xiongnu and
31 Sakha skulls from the Musée de l’Homme col-
lection in Paris, France [43]. In this study, Xiongnu
skulls were found to be closest to those of Mongols,
as well as those from the Iron Age (2nd - 3 centu-
ry BCE) proto-Mongolic Xianbei people from the
Liaoning province in China and Sakhas, albeit the
direct connection between Xiongnu and Sakha re-
mained mysterious at the time. Although the cur-
rent evidence from ancient DNA studies may so far
fail to substantiate such a claim, our results from
median-joining network and associated TMRCA
analyses for the most prevalent Y-chromosomal
STR Sakha haplotypes (Table 2-4) suggest that the
penetration of haplogroup N to the Northeastern
Siberia possibly started as early as the period of
Xiongnu domination in the Asian Steppe around
200 BCE. In addition to the matches between four
contemporary Sakha Y-chromosomal STR haplo-
types (‘the Xiongnu clan’) and that from the remains
TIKHONOV D.G., GURKAN C., DEMIRDOV K. D., BEYOGLU E
107
of a Xiongnu nobility from the Egyin Gol necrop-
olis (grave 25A), other lines of scientic evidence
have also been accumulating towards cultural, eco-
nomic and historical links between Middle Lena
and Xiongnu dominated Asian Steppe [34, 44, 45].
According to the Sakha legends and this study,
Elley and Omogoy were not Turkic per se. e core
population forming the basis for Sakha ethnogen-
esis may be represented by the ‘autochthonic’ de-
cent cluster as classied in the current study instead
(Table 2 and Figure 3). Such a classication for a
subset of the contemporary Sakha paternal lineag-
es was made based on an earlier hypothesis, which
also considered the contribution of an aboriginal
stratum of unknown origin during the ethnogen-
esis of Sakhas [34]. Among such ‘Autochthons’
haplotypes observed among Sakhas, there were 21
matches at YHRD, 23.8% of which were Turkic,
along with 3 matches from Turkey and 2 matches
with Yakut-speaking Evenks. One ‘autochthons’
Y-chromosomal STR haplotype had matches at
4 Y-chromosomal STR loci with an ancient DNA
sample from the 2200-2400-years-old Pokrovsk
grave site in Yakutia [41]. Ancient DNA analyses
from the Balychtakh site dated between 1420-1470
CE in Yakutia revealed a 12-loci Y-chromosomal
STR haplotype and a haplogroup assignment of K
(excluding N, O and P) [6]. While a correspond-
ing paternal lineage has not so far been detected
among contemporary Sakhas, there were 28 match-
es in total at YHRD from Mongolia, China, Russia
and Turkey, for the last one of which there were 10
matches from 4 dierent geographic regions. ere-
fore, the ‘autochtons’ cluster from the current study
may correspond to a core Sakha population where
the agglutinative language structure was based on.
Current consensus is that the Yakut language arose
as a result of Turkicisation of an unknown language
that also went Mongolization in the process of de-
velopment. Loaning of Mongolian words to the Ya-
kut language could only be dated as far back as 12
to 13 centuries CE somewhere in Central Yakutia
[46], curiously coinciding with the legend of Elley
and Omogoy and data from population genetics of
ancient and contemporary Sakhas.
e scientic reconciliation of all the data from
genetic studies on contemporary and ancient
Sakhas, as well as the current archaeological evi-
dence towards the emergence of the Sakha culture
seemingly correlate well with the popular historical
narratives, such as the legends of Elley and Omogoy.
According to these legends, Elley is also credited for
the introduction of novel handicras and farming
technologies to the Middle Lena. As in the case of
traditional fairy tales [47], legends are known to
be based at least in part on some historical anec-
dotes [48]. In any case, these stories were possibly
associated with the events that occurred in the Asia
Steppe spanning the Southern part of Siberia from
the Okhotsk coast to the Caspian Sea and in peri-
ods of the dawning and decline of the state admin-
istrative formations of Xiongnu, Turkic Khaganates
and the Khitan Empire. A large proportion of the
contemporary Sakha paternal lineages belong to
the haplogroup N and seemingly fall under ve ma-
jor descent clusters based on 9-loci Y-chromosomal
STR data as described in Table 2 and Figure 2. e
beginning of these paternal lineages was perhaps
laid by inuential Xiongnu nobility, closely related
to a territory of the middle Lena River since the an-
cient times. In one way or the other, 79 to 89% of
all contemporary Sakha paternal lineages could be
traced down to only three haplotypes in total, which
are in turn closely related themselves and exhibit
only minute dierentiations of an ancestral 17-loci
Y-chromosomal STR haplotype (Table 4). Curious-
ly, even when the number of Y-chromosomal STR
loci analyzed were increased up to 23, no signicant
increase in the resolution of the Sakha paternal lin-
eages could be observed, which constituted a fur-
ther proof towards the actual homogeneity of these
lineages [27]. Unfortunately, the current data still
does not have the resolution to help deduce wheth-
er such Y-chromosomal STR mutation among the
three most prevalent Sakha paternal lineages took
place in situ in Yakutia or prior to their arrival in
the Asian Steppe. In either case, the legends of Elley
and Omogoy and how they have together changed
the way of life in Middle Lena forever are still in
agreement with the reconciled scientic data in the
current study. Finally, around 8% of the contempo-
rary Sakha Y-chromosomal STR haplotypes, which
can not be classied under the ve major Sakha pa-
ternal lineages discussed in this manuscript may be
ON THE ORIGINS OF THE SAKHAS' PATERNAL LINEAGES
108
considered as a mixed Sakha cluster instead, more
than half of which (4,9%) constitute autochthonic
polymorphisms. Apparently these paternal lineages
may have constituted a core for forming an aggluti-
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TIKHONOV D.G., GURKAN C., DEMIRDOV K. D., BEYOGLU E
111
About the authors
TIKHONOV Dmitry Gavrilievich, MD, Professor, Senior Research Ocer of the Scientic research
Center of the Medical Institute of the North-Eastern Federal University, 677009, Yakutsk, St. Bld. 8, Russia,
https://orcid.org/0000-0003-3385-9471, e-mail: Tikhonov.dmitri@yandex.ru.
GURKAN Cemal, Director of the Turkish Cypriot DNA Laboratory / Academic Sta, Dr Fazil Kucuk
Faculty of Medicine, Eastern Mediterranean University, Nicosia (North Cyprus), Turkey, http://orcid.
org/0000-0001-7379-4559, e-mail: cemal.gurkan@gmail.com.
DEMIRDOV Damla, Turkish Cypriot DNA Laboratory, Committee on Missing Persons in Cyprus
Turkish Cypriot Member Oce, Nicosia (North Cyprus), Turkey, damla0330@hotmail.com.
BEYOGLU Erdem, Baris Mental and Neurological Disorders State Hospital, Nicosia (North Cyprus),
Turkey erdembey@yahoo.com.
Supporting Online Material
Refer to Web version of the journal for supplementary material.
ON THE ORIGINS OF THE SAKHAS' PATERNAL LINEAGES
