The genetic history of Russian old settlers of polar northeastern Siberia

ISSN 1022�7954, Russian Journal of Genetics, 2010, Vol. 46, No. 11, pp. 1386–1394. © Pleiades Publishing, Inc., 2010.Original Russian Text © R.I. Sukernik, N.V. Volodko, I.O. Mazunin, N.P. Eltsov, E.B. Starikovskaya, 2010, published in Genetika, 2010, Vol. 46, No. 11, pp. 1571–1579.

1386

The history of the first Russian settlements in polarregions of eastern Siberia begins from Cossack troopsacting upon the orders of the Moscow State to “seekfor Yukaghir lands and bring yasaq�free people tounder the rule of the high lord.” Soon after the LenaFort (the future city of Yakutsk) was founded in 1632,a unit under the command of Ivan Rebrov, havingtraveled about 1000 km through the Arctic Ocean bykoches (small sailing boats), founded two settlementsat the mouth of the Indigirka River, which was referredto as the “dog’s river” in that period. One of them gaverise to the modern Russkoe Ust’e [1]. Almost simulta�neously with the “sea path” to the Indigirka, a landroute was opened from Yakutsk over the VerkhoyanskRidge. In 1642–1643, a unit of 23 Cossacks reachedthe Alazeya and Kolyma by sea. On the banks of theserivers, they built two forts: Nizhnekolymskii (a littleupstream of the modern Cherskii town) and Sred�nekolymskii (the modern Srednekolymsk). This iswhere the sea part of Semen Dezhnev’s travel, whichended with the discovery of a strait between Asia andAmerica, began. Having led the remnants of his mento the upper Anadyr’ River, Semen Dezhnev foundedthe Anadyrskii winter shelter (later, a fort) somewhatupstream of the Markovo village in 1694. Thus, per�manent Russian settlements appeared at the delta ofthe Indigirka, on the lower Kolyma, and on the upperAnadyr’ in the mid�17th century. Most hunters andCossacks are likely to have originated from Pomor’eon the White Sea [2, 3]. The settlers were called Indi�girka (or Russkoe Ust’e) people, Kolyma (Pokhodskor Porech’e) people, and Anadyr’ (or Markovo) peo�

ple. They were characterized by the same type of eco�nomic activities, including fishery, gathering, fur (Arc�tic fox) hunting and trapping, and sled dog breeding.Their small number determined a narrow range ofintermarriage shaped by their relations with Yuk�aghirs, the largest ethnic group in the area [4]. Sincethat time, the descendants of the first Russian settlersand Yukaghir women have remained an isolated popu�lation among the surroundings ethnic groups. TheRussian old settlers are extremely heterogeneous withrespect to anthropological types. In some old familiesof former Cossacks, mainly in Pokhodsk and RusskoeUst’e, persons of the European type, with fair hair,gray or blue eyes, a straight nose and relatively narrowcheekbones could be seen as recently as the late 19thcentury; however, mixed types were prevailing [5].According to Bogoraz [5], “Russian migrants mixingwith indigenous aborigines have formed a specifictribal mixture that, on the one hand, has retained theRussian language with many archaic words andexpressions; Russian songs, tales, superstitions, andmany traditions and, on the other hand, has assimi�lated the material life style of indigenous tribes and thehabits of ichthyophagous hunters like Yukaghirs, wholived there before Russians appeared” [5]. As timepassed, the settlements founded by the first Russianmigrants became increasingly more multiethnic due tolater migrants, including Evens, Yakuts, Chukchi, andexiled Russians Tatars [2, 6–9].

The purpose of this study was to reconstruct thehistory of the formation of anthropologically isolatedRussian old settlers of polar Siberia, with unique prop�

The Genetic History of Russian Old Settlersof Polar Northeastern Siberia

R. I. Sukernika, N. V. Volodkoa, I. O. Mazunina, N. P. Eltsovb, and E. B. Starikovskayaa

a Institute of Chemical Biology and Fundamental Medicine, Siberian Branch, Russian Academy of Sciences,Novosibirsk, 630090 Russia

e�mail: [email protected] Institute of Cytology and Genetics, Siberian Branch, Russian Academy of Sciences, Novosibirsk, 630090 Russia

Received March 5, 2010

Abstract—The mtDNA variation has been studied in representatives of the Russkoe Ust’e (n = 30), Kolyma(n = 31), and Markovo (n = 26) ethnic subgroups originating from Russian military men, hunters, and fisherswho married local Yukaghir women and settled at the Arctic Ocean coast and on the Anadyr’ River more than350 years ago. The mtDNA haplotypes characteristic of indigenous Siberian peoples have been demonstratedto form the basis of the mitochondrial gene pool of Russian old settlers of the region. Only one of 30 identifiedhaplotypes belonging to 11 haplogroups (H2a) is characteristic of European populations. The C and D hap�logroups are the most diverse. The analysis has revealed the characteristics of the population structure of theRussian old settlers and allowed them to be interpreted in terms of recent historical and environmental pro�cesses.

DOI: 10.1134/S1022795410110153

HUMAN GENETICS

RUSSIAN JOURNAL OF GENETICS Vol. 46 No. 11 2010

THE GENETIC HISTORY OF RUSSIAN OLD SETTLERS 1387

erties of mitochondrial DNA (mtDNA), such asstrictly maternal inheritance, a relatively high muta�tion rate, and the absence of recombinations, servingas tools.

MATERIALS AND METHODS

Populations

Russkoe Ust’e (Indigirka) people. There were sev�eral settlements with a total population of ~275 peoplein the delta of the Indigirka, on the banks of its threemain tributaries (Russkoustinskaya, Srednyaya, andKolymskaya) almost until the mid�20th century. TheRusskoe Ust’e village, which still exists, have beenmarked in all geographic maps and mentioned in sci�entific literature since the time when a unit under thecommand of lieutenant D. Laptev (who had todescribe the Arctic Ocean coast east of the Lena on theorder by V. Bering, the head of the Great NorthernExpedition) was forced to spent the winter of 1739–1740 at the mouth of the Indigirka. At that time, nomore than 500 people lived in Russkoe Ust’e andneighboring villages and hunter’s lodges. However,after a number of epidemics of smallpox in the 19thcentury (especially the 1884 epidemic), the ethniccomposition of Russkoe Ust’e residents substantiallychanged due to the inflow of new immigrants, namelyEvens and Yakuts living in the delta of Indigirka.

Kolyma (Porech’e) people. Kolyma people origi�nate from members of Cossack troops who originallysettled on the Malaya Chukoch’ya River but wereforced out of that place by Chukchi and founded a newsettlement, Pokhodsk, on the left creek of the Kolymadelta. Kolyma people used to intermarry with RusskoeUst’e people, which was facilitated by geographic andclose economic relations based on fishery and fur(Arctic fox) hunting in the Arctic Ocean coastalregion. According to the 1897 census, about 680Porech’e people, including 85 Yukaghirs, and 77Yakuts, lived in Pokhodsk, as well as neighboring andremote hunter’s lodges on the lower Kolyma River [5].

The geographic remoteness together with adverseenvironment favored the preservation of Indigirka andKolyma peoples as anthropologically and linguisti�cally isolated populations until recent time. Under theconditions of long�term social isolation, they pre�ferred to marry women from their own territorialgroup. Therefore, marriages between cousins, half�sibs, and relatives with multiple kinship relations wererather common. Among the few “locally Russian, pre�settling” surnames, Chikachev, Kiselev, Strukov,Antonov, and Shkulev have remained the most preva�lent until the present time. These surnames are fre�quently mentioned in petitions and other commercialand official documents of the 17th century [Sukernik,unpublished data; 2, 8, 10, 11].

Markovo people. Although many individual familygroups were forced to move after the Anadyr’ fort was

evacuated because of military encounters with“unpeaceful” Chukchi and Koryaks, and Anadyr’people moved to Gizhiga in 1771, several hundreds ofRussians mixed with other ethnic groups, includingChuvans, Lamuts (Evens), and Chukchi, settled in thevillages of Markovo, Eropol, and Chuvanskoe in thefirst half of the 19th century. At present, their ancestorsare called Markovo people.

Samples

The original material (pedigrees and blood sam�ples) was obtained in the course of several expeditionsto the Russkoe Ust’e and Chokurdakh villages ofAllaikhovskii raion (the Russkoe Ust’e sample, n = 32)and the Pokhodsk and Cherskii villages of Nizh�nekolymskii raion (the Pokhodsk sample, n = 31) ofthe Republic of Sakha (Yakutia) and the Markovo vil�lage of Anadyrskii raion of the Chukchi AutonomousRegion (the Markovo sample, n = 26) (Fig. 1). Thesamples were collected by R.I. Sukernik, E.B. Starik�ovskaya, and N.V. Volodko in 2005–2008. To obtain arepresentative set of blood samples for molecular anal�ysis, a pedigree covering four or five generations andincluding data on the ethnicity and mother spokenlanguage was compiled for each family. An informedwritten consent to participate in the study wasobtained in each particular case.

Molecular Genetic and Statistical Analyses

Genomic DNA was isolated from a buffy coat bythe standard method using a QIAamp Blood Kit(QIAGEN). The first hypervariable segment (HVS1)of the mtDNA control region was sequenced, and theRFLP analysis of the mtDNA coding region was car�ried out. We also sequenced a number of diagnosticsites of the coding region. The sequencing was per�formed using fluorescent labeling of double�strandedPCR products by means of a Big Dye Terminators kit(ABI/Perkin�Elmer Cetus) followed by separationusing an ABI Prism 3100 automated sequencer. Theresults were treated using the Sequencher version 4.2software (GenCode). The list of endonucleases andprimers for both complete sequencing of mtDNA andRFLP analysis was published earlier [12, 13]. Eachhaplotype was classified with a phyletic lineage on thebasis of the set of specific polymorphic substitutions inthe nucleotide sequences of the coding and controlregions according to the permanently renewed classi�fication [13–16]. Complete sequencing of two sam�ples was performed, and the sequences were depositedin the GenBank database under the access numbersGU903055 and GU903056.

Bernstein’s equation [17] was used to estimate the geneinflow rate: M = Pm – Pb/Pa – Pb, where M and (1 – M)are the gene contributions of two original populations,Pa and Pb are the initial allele frequencies in the popu�lations a and b before admixture, and Pm is the fre�

1388


SUKERNIK et al.

quency of the same allele in the mixed population. Thealleles for which the difference Pa – Pb is the largest arethe most informative.

The phylogenetic tree of the complete mtDNAsequences for the B5b2 and H2a haplogroups wasdrawn by the maximum parsimony method using themtPhyl software developed by N.P. Eltsov. It is avail�able at the Web site http://www.bonet.nsc.ru/labs/mtgenome/programs.html. The divergence ofmtDNA clusters was dated using the ρ statistic[18].The standard deviation (σ) was determined by thestandard method [19]. The relative estimates ρ and σwere expressed in absolute time units using the calibra�tion mutation rate of 5138 years per transition in themtDNA coding region [20].

RESULTS AND DISCUSSION

Table 1 shows the diversity of mtDNA haplotypesidentified in Russian old settlers; Table 2 shows thefrequencies of mitochondrial haplogroups in Russianold settlers as compared to those in Yukaghirs, Evens,and Chukchi. Note the diversity of haplotypes andhaplogroups distributed among the three isolated pop�ulations varying at a relatively low frequency. At the

same time, the C2a haplogroup characteristic of Yuk�aghirs [14] has been found at frequencies of 25.0, 35.5,and 23.1% in Russkoe Ust’e, Pokhodsk, and Markovo,respectively. Hence, the gene contributions of Yuk�aghir women in the Russkoe Ust’e, Pokhodsk, andMarkovo populations are 61.0, 86.6, and 56.3%,respectively. To estimate the contribution of Chukchi,the A2 lineage originating from hunters of sea mam�mals of ancient Beringia is the most informative [14,19, 21, 22]. The A2 haplogroup is absent in Yukaghirs,Evens, and Koryaks whose populations have not beenaffected by an inflow of genes from the neighboringReindeer Chukchi populations [14, 23]. In Russianold settlers, the A2 concentration distinctly decreaseswith an increase in the distance from the area origi�nally inhabited by Chukchi (Table 2). Correspond�ingly, the gene contributions of Chukchi to the Mark�ovo, Pokhodsk, and Russkoe Ust’e populations were35.5, 17.7, and 0.0%, respectively. The G1 haplogroupwas the most prevalent in the Markovo population(30.8%), which also suggests a gene flow from Evens andChukchi.

Group D haplotypes have low frequencies in Rus�sian old settlers, each population being characterizedby a different set of haplotypes. Four D haplotypes

Fig. 1. The region of expeditions for sample collection.

A R C T I C O C E A N

East SiberianSea

Laptev Sea

Chokurdakh

Russkoe Ust’e

Pokhodsk

CherskiiMarkovo

Sea ofOkhotsk

Yakutsk

Lena

Yana

Indigirka

Kolyma

Ana

Nizhnyaya

dyr’

Tunguska



Tab

le 1

.T

he d

iver

sity

of m

tDN

A h

aplo

type

s in

Rus

sian

old

set

tler

s

Hap

logr

oup

RF

LP

(S

NP

)H

VS

1 (–

16 0

00)

Rus

skoe

U

st’e

Pok

hods

kM

arko

vo

H2a

(–/–

) 476

9a�7

025a

(14

38 2

706)

CR

S2

––

U4a

1(–

/–) 4

643k

113

29a

1230

8g (

8818

)13

4 31

1 35

6 51

91

––

А2а

(–/–

) 663

e (3

330)

11

1 19

2 22

3 2

90 3

11 3

19 3

62–

14

A2b

(–/–

) 663

e (1

1365

)11

1 22

3 26

5 29

0 31

9 36

2–

–1

(–/–

) 663

e (1

1365

)11

1 17

6 22

3 26

5 29

0 31

9 36

2–

2–

B5b

2(+

/–) (

8281

d9)

111

140

189

234

243

463

519

–1

–F

1b(–

/–) �

6389

w�1

2406

h�12

629b

172

189

232A

249

304

311

384

519

–1

–(–

/–) �

6389

w�1

2406

h�12

629b

189

232A

245

249

304

311

519

3–

–Y

1a(+

/–) 7

933j

�83

91e

126

189

231

266

519

2–

–С

2a(+

/+) �

1325

9o (

3816

)22

3 29

8 32

7 51

98

116

C2b

1(+

/+) �

1715

c �1

3259

o (1

2672

)12

9 2

23 2

98 3

27 5

193

21

(+/+

) �17

15c�

1325

9o (

1267

2)09

3 12

9 22

3 32

7 51

9–

–1

C2b

2(+

/+) �

1325

9o (

1267

2)17

1 22

3 29

8 32

7 34

4 35

7 51

94

––

C3

(+/+

) �13

259o

09

3 18

9 22

3 26

1 28

8 29

8 51

91

2–

(+/+

) �13

259o

093

223

288

298

327

390

519

–1

–D

2b(+

/+) �

5176

a (1

935

8683

112

15 1

4905

)12

9 18

9 22

3 36

21

––

D3a

1(+

/+) �

951j

�51

76a�

1018

0l 1

5437

e 22

3 31

9 36

21

––

D3a

2a(+

/+) �

5176

�101

80l 1

4923

c 15

437e

(68

81 1

4815

)22

3 31

9 32

0 36

2–

1–

D3a

2a1

(+/+

) �51

76a�

1018

0l 1

3717

a 14

923c

154

37e

(688

1 14

815)

093

172

173

223

255

319

362

––

2D

5a(–

/–) �

5176

a�12

026h

(10

397

1270

5)

092

172

189

223

266

362

–2

–D

8a(+

/+) �

1715

c�51

76a

(876

2 12

651C

)04

2 09

3 21

4 22

3 36

21

––

D9a

(+/+

) 483

0n�5

176a

22

3 2

94 3

621

1–

G1b

(+/+

) 483

0n 8

198a

01

7 09

3 12

9 22

3 51

93

41

(+/+

) 483

0n 8

198a

01

7 09

3 12

9 17

2 22

3 26

5 5

19–

–1

(+/+

) 483

0n 8

198a

017

129

223

519

1–

5(+

/+) 4

830n

819

8a

017

093

129

153

223

319

519

––

1G

2a(+

/+) 4

830n

�759

8f

223

227

278

362

––

1(+

/+) 4

830n

�759

8f

223

227

234

278

362

456

–1

–Z

1a(+

/+) �

6749

v (1

0325

) 12

9 18

5 22

3 22

4 26

0 29

8 51

9–

–2

M7c

(+/+

) �48

48c

982

0g (

5442

)14

5 22

3 29

5 30

4 51

9–

1–

Tot

al:

32

3126

Not

e: T

he d

iagn

osti

c si

tes

in th

e co

ding

reg

ion

and

HV

S1

are

bold

face

d. R

estr

icti

on e

ndon

ucle

ases

are

des

igna

ted

by s

ingl

e le

tter

s: a

= A

luI,

c =

Dde

I, e

= H

aeII

I, f

= H

haI,

g =

Hin

fI, h

=H

paI,

j =

Mbo

I, k

= R

saI,

l =

Taq

I, n

= H

aeII

, o =

Hin

cII,

v =

Avr

II, a

nd w

= T

sp50

9I. T

he p

osit

ions

of t

he m

utat

ions

hav

e be

en d

eter

min

ed r

elat

ive

to th

e rC

RS

[33

]. T

rans

vers

ions

are

indi

cate

d by

cap

ital

lett

ers

corr

espo

ndin

g to

the

new

bas

e; th

e ot

her

mut

atio

ns a

re tr

ansi

tion

s. T

he p

rese

nce

of m

utat

ions

in th

e co

ding

reg

ion

unde

tect

able

by

rest

rict

ion

anal

ysis

was

det

erm

ined

by

sequ

enci

ng; t

hese

mut

atio

ns a

re in

dica

ted

in p

aren

thes

es. T

he p

rese

nce

or a

bsen

ce o

f the

103

94 D

deI/

1039

7 A

luI

rest

rict

ion

site

s is

indi

cate

d as

(+

/+),

(+

/–),

or

(–/–

). 8

281d

9 is

a 9

�bp

dele

tion

in th

e in

terg

enic

seg

men

t CO

II/t

RN

AL

ys.

1390


SUKERNIK et al.

have been identified in the Russkoe Ust’e population;three haplotypes, in the Pokhodsk population; andone haplotype, in the Markovo population. It is note�worthy that the D5a haplotype identified in two repre�sentatives of the Pokhodsk population is identical tothat found in 400�year�old burial sites in central Yaku�tia [24]. Apparently, the D5a haplogroup is formed by a

gene flow from the Yakut population, where its fre�quency is the highest in Siberia (15%) [25, 26].

The fact that representatives of Russian old settlerscarry haplotypes that are untypical of Yukaghirs agreeswith recent historical events in the lower Indigirka andKolyma basins, including a series of devastating small�pox and measles epidemics and expansion of Tungus�

Table 2. The spectrum of mitochondrial haplogroups in Russian old settlers as compared to Yukaghir, Even, and Chukchi pop�ulations

Haplogroup

Frequency, %

Russian old settlersYukaghirs* (100) Evens** (87) Chukchi* (182)

Russkoe Ust’e (32) Pokhodsk (31) Markovo (26)

H2a 6.3 – – – – –

U4a1 3.1 – – – – –

A2 – 9.7 19.2 – – 54.9

B5b2 – 3.2 – – – –

F1b 9.4 3.2 – – 5.7 –

Y1a 6.3 – – – 5.7 –

C2a 25.0 35.5 23.1 41.0 12.6 7.1

C2b1 9.4 – 7.7 8.0 5.7 –

C2b2 12.5 6.5 – 5.0 10.3 –

C3 3.1 9.7 – 13.0 3.4 7.1

C4 – – – – 1.1 –

Z1a – – 7.7 2.0 2.3 –

D2a – – – – – 8.2

D2b 3.1 – – 1.0 3.4 –

D2c – – – – 1.1 –

D3a1 3.1 – – 2.0 2.3 1.1

D3a2a – 3.2 – 1.0 – 3.8

D3a2a1 – – 7.7 – – –

D4a – – – – 1.1 –

D5a – 6.5 – 5.0 3.4 –

D6 – – – 1.0 19.5 –

D7 – – – 2.0 4.6 –

D8 3.1 – – 2.0 2.3 –

D9 3.1 3.2 – 5.0 3.4 –

D* – – – – 1.1 –

G1 12.5 12.9 30.8 12.0 10.3 17.6

G2 – 3.2 3.8 – – –

M7c – 3.2 – – – –

Notes: * [14].** Unpublished data.



speaking Evens and Turkic�speaking Yakuts [5, 7]. Forexample, the total proportion of the B5b2, D5a, F1b,G2a, and M7c mitochondrial haplogroups is 9.4, 19.3,and 3.8% in the Russkoe Ust’e, Pokhodsk, and Mark�ovo populations, respectively.

In this study, we have performed completesequencing of two rare mtDNAs. One of them, foundin the Pokhodsk population, is a derivative of the east�ern Eurasian haplogroup B5b2 (Fig. 2), which mostlyincludes Japanese sequences, as well as several Chi�nese, one Philippine, and one Negidal sequences con�taining a common motif of five mutations (4895,15850, 16111, 16234, and 16463). Its coalescence timeis 19.85 ± 4.04 years [13, 27]. The Pokhodsk variant

has a unique set of seven mutations in the codingregion (3305, 5964, 6737, 7211, 7990, 8994, and9152). The Pokhodsk variant of B5b2, as well as aNegidal variant that we sequenced earlier [13], is anisolated subcluster.

Another rare haplotype found on the Indigirkabelongs to the western Eurasian haplogroup H2a,which is less common in European populations thanthe H1 and H3 haplogroups [28, 29]. As seen in thephylogenetic tree (Fig. 3), the H2a haplogroup has acoalescence time of 15.32 ± 3.39 years and consists ofseveral subclusters (H2a1–H2a5). The Indigirka vari�ant of H2a is characterized by a unique set of muta�tions (73, 4350, 5460, and 8709) and forms a separate

Fig. 2. The phylogenetic tree of complete mtDNA sequences of the B5b2 haplogroup. The positions of the mutations have beendetermined relative to the rCRS [33]. Transversions are indicated by capital letters corresponding to the new base; deletions aredenoted del; insertions, ins; the other mutations are transitions. Back mutations are underlined. The point mutations at positions16182 and 16183, poly(C) tract length polymorphism (309–315 bp), ant the point mutation at position 16519 are hypervariableand have not been used for constructing the phylogenetic tree. The coalescence time and its standard deviation are indicated (inthousands of years) for the main nodes. The ethnic populations where the samples were taken and individual GenBank accessnumbers of the samples are indicated.

R11'B

9bpdel

B

522delCA709

85849950

1039816140

B5

103204

15988829

12361152231550815662158511592716243

B5b

489515850161111623416463

B5b2

204481

236141619299

1110112451

B5b2c

8664200

15766

37864907

AP008465AP008584

Japanese 16456С

AP008323

16181insC

AP010689

19511647G

AP008737

6249

AY255179

8551119081520916362

AP008847

195

AP011059

57441143716291

B5b2a

AP008546

39721370814693

AP008488

6767

DQ112865

19.85 ± 4.04

4.57 ± 1.61Japanese

Japanese Japanese

Japanese

Japanese

Japanese

Japanese

Japanese

Chinese Asian

103

B5b2a' b

8894T14750

B5b2b1

AP008910

1819

AP008403

AP008491

322

AP008580

20715758

B5b2b

42517058

AP008371

33985918

12696

AP008846

16265

AP009425

55493

522delCA1193144045616032

7590G8829

1091510972152231612916244

AY519489Negidal

Japanese

Japanese

Japanese

Japanese

JapaneseJapanese

Japanese

2073335754278538021

10807147501612916399

GQ119041Philippine

204

146364037668065

9755delG1233216256

GU377081

3305596467377211799089949152

GU903056

Russian Old Settler

131

16.02 ± 4.85

17.98 ± 4.25

8.81 ± 4.02

Chinese

1392


SUKERNIK et al.

subcluster that is apparently typical for migrants fromPomor’e. Indeed, as evident from pedigrees, H2a hasbeen inherited from a single ancestor in the maternalline; it is present in at least one quarter of RusskoeUst’e people with the “locally Russian” surnamesKiselev and Chikachev.

In addition, Indigirka people have inherited a hap�lotype of the U4 haplogroup. The frequency of the U4haplogroup in eastern European and western Siberianpopulations is polymorphic [13, 30–32]. In general,the rare haplogroups H2a U4a1, and Y1a on the Indi�girka and B5b2 and M7c on the Kolyma indicate ran�dom events in the course of formation of the modernlong�term Russian resident population.

Thus, the molecular genetic analysis of mtDNAdiversity has shown that the mitochondrial gene poolof Russkoe Ust’e, Pokhodsk, and Markovo peopleswas mostly inherited from Yukaghirs, which entirelyagrees with the population history and geography ofthe region [7]. According to historical documents, in

the late 17th century, “over the entire vast Yukaghirland from the Lena to the Anadyr’, wives of both mil�itary men and hunters were Yukaghir women.” [4].The contribution of the Chukchi maternal gene poolin the Markovo and Pokhodsk populations (the A2haplogroup) is also considerable. The G1 haplogroup,which is frequent in Koryaks and Evens, has left a sub�stantial trace in each of the three samples from Rus�sian old settlers. The contribution of later immigrants(Yakuts) is apparently reflected by the presence of therare mtDNA haplogroups D5a, F1b, G2a, and M7c.

ACKNOWLEDGMENTS

We are grateful to the residents of the RusskoeUst’e, Chokurdakh, Pokhodsk, Cherskii, and Mark�ovo villages who took part in the project. We thankAcademician A.L. Derevyanko for intellectual sup�port and assistance in the course of the study. We thankD.G. Solodovnikov, the head of administration of theMarkovo village; I.P. Suzdalov, the head of the Nizh�

Fig. 3. The phylogenetic tree of complete mtDNA sequences of the H2a haplogroup. Designations are the same as in Fig. 2.

H

H2

H2a

H2a2b

H2a2b1

rCRS

H2a3

H2a4H2a1

H2a5

H2a1a

H2a2

1438

4769

20.46 ± 6.15

15.32 ± 3.39

1081016274

73435054608709

15556038

15784

EF657353EuropeanGU90305

Russian Old Settler34381089913911

EF657571European

15216519

AY738962Italian

1462EF657524European

8242259

16519

AY339429Finnish

3388AEF657238European

1555

EF657289European

750

EF657416EuropeanEF657556European

2638860

15326

EF657704European

EU130575Canadian of French origin

69987561

1024515340

EF657438European

10700EF657579European

16291

408016235

AY339426

525insAC7853

EU719115unknown

3388A1629316400

EF418606unknown

38825460

13749

EU443605

1468634

EU444119104461447016270

EU284181European

72113958C

EF657705

Finnish

unknownRussian

European

15216129

EU714299unknown

2636716

1290314311

EU716647unknown

929912366

EU330412Finnish

8182

EF657287European

15314

16195

EU795361European

1891625916519

EU157923Swedish

11140

161871622716362

AM263181Dargin

55576221

AM263179Dargin

95116354

EF657247EuropeanEF657661EuropeanFJ238053German

13708

8843

EF657627European

DQ112834European

11260

EF657441European

1123314323

AY738963Spanish

184216291

4592

FJ527772Spanish

FJ527773Spanish

FJ527774Spanish

FJ527775Spanish

FJ527776Spanish

10391

FJ527779Spanish

8285insC

FJ527778Spanish

9033

FJ527777Spanish

62608679

4769

EF657470EuropeanFJ161702unknown

73

FJ800808 Irish

14646598084

EU780223Danish

914511914

FJ496869unknown

575751

633916124

FJ842614Danish

38345177

16519

AY339427Finnish

101721031016291

EU597521unknown

6173

1309515932161571620916519

FJ668389unknown

15216178

AY339428Finnish

72G2195

3395C5978

1339516256

EU233278unknown

310102351426616129

EF661009

6.85 ± 1.82

11.25 ± 4.32

6.17 ± 2.52

9.59 ± 2.3710.7 ± 5.4

Italian



nekolymskii ulus municipal unit; Z.M. Robek, thedirector of the museum of the Cherskii village;V.F. Sokovikov, the head of administration of thePokhodsk village; I.P. Borisova, the director of themuseum of the Pokhodsk village; A.S. Dronov, thehead of the Allaikhovskii ulus municipal unit;A. Kiselev, the head of administration of the RusskoeUst’e village; and G.V. Koshkarskaya, the first deputyhead of the Department for Ethnic Groups and Feder�ative Relations of the Republic of Sakha (Yakutia) fortheir assistance in collecting the material.

This study was supported by the Russian Founda�tion for the Humanities (project no. 08�01�00356a)and the Russian Foundation for Basic Research(project no. 09�04�00183a).

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The genetic history of Russian old settlers of polar northeastern Siberia