Ancient History of the Arbins, Bearers of Haplogroup R1b, from Central Asia to Europe, 16,000 to 150

Advances in Anthropology

2012. Vol.2, No.2, 87-105

Published Online May 2012 in SciRes (http://www.SciRP.org/journal/aa) http://dx.doi.org/10.4236/aa.2012.22010

Ancient History of the Arbins, Bearers of Haplogroup R1b, from

Central Asia to Europe, 16,000 to 1500 Years before Present

Anatole A. Klyosov

The Academy of DNA Genealogy, Newton, USA

Email: aklyosov@comcast.net

Received March 14th, 2012; revised April 13th, 2012; accepted April 23rd, 2012

This article aims at reconstructing the history of R1b ancient migrations between 16,000 and 1500 years

before present (ybp). Four thousand four hundred eight (4408) haplotypes of haplogroup R1b (with sub-

clades) were considered in terms of base (ancestral) haplotypes of R1b populations and the calculated

time to their common ancestors. The regions considered are from South Siberia/Central Asia in the east

(where R1b haplogroup arose ~16,000 ybp) via the North Kazakhstan, South Ural to the Russian Plain

and further west to Europe (the northern route entering Europe around 4500 ybp); from the Russian Plain

south to the Caucasus (6000 ybp), Asia Minor (6000 ybp) and the Middle East (6000 - 5500 ybp) to the

Balkans in Europe (the southern route, entering Europe around 4500 ybp); along North Africa and the

Mediterranean Sea (5500 - 5000 ybp) via Egypt to the Atlantic, north to Iberia (the North African route

with arrival to the Pyrenees 4800 ybp). The Arbins (bearers of R1b haplogroup) along their migration

route to the Middle East and South Mesopotamia apparently have established the Sumer culture (and the

state), moving westward to Europe (5000 - 4500 ybp) carrying mainly the R-M269 subclade and its

downstream L23 subclade. This last subclade was nearly absent along the North African route, and/or did

not survive the migration to Iberia or evidenced later. At the arrival to Iberia (4800 ybp) the M269 sub-

clade split off M51 and soon thereafter the L11 downstream subclades. These populations became known

as the Bell Beakers and moved north, along with the newly arisen subclades of P312 and L21 (which split

off within a few centuries after P312). Those subclades and their downstream clades have effectively,

without major interruptions, populated Europe (the smooth haplotype trees demonstrate the near non-stop

proliferation of R1b haplotypes in Europe). They are evidenced from the Atlantic eastward to the Balkans,

Carpathian Mountains, present day Poland to the western border of the Russian Plain and up to the Baltic

Sea. The Isles had a different history of R1b migrations. The bearers of L11, P312 and L21 moved to the

Isles by land and sea concurrently with those Arbins who were populating Europe between 4000 and 2500

ybp and formed the respective “local” subclades of P314, M222, L226, which largely populated the Isles.

As a result, a significant part of the Isles is populated almost exclusively by the Arbins, whose frequency

reaches 85% - 95% among the current population. In general, the frequency of Arbins in Western and

Central Europe, reaches—albeit not uniformly—some 60% of the population. This study essentially pre-

sents an example of application of DNA genealogy in studying the history of mankind.

Keywords: Y Chromosome; Mutations; Haplotypes; Haplogroups; TMRCA; STR; SNP; Arbins; Aryans;

R1b

Introduction

The origin and history of haplogroup R1b, bearers I refer to

as the Arbins (see explanation below), currently populate nearly

60% of Western and Central Europe. Additionally Arbins

populate significant parts of the Caucasus, Anatolia and Asia

Minor, Middle East, and many locations in Central Asia, in-

cluding South Siberia, Altay, Tuva, North-Western China,

Middle Asia, some Ural and Middle Volga regions with ethnic

groups and populations such as the Bashkirs, Tatars, Chuvash,

and other.

History of R1b was significantly distorted from the begin-

ning of “genetic genealogy” at the end of 1990-s, when it was

claimed, quite groundlessly, that R1b arose in Europe some

30,000 years before present (ybp). Groundlessly because, in-

deed, the claims were based on no data. Such data never existed.

Nevertheless, statements and claims such as “Around 30,000

years ago, a descendant of the clan making its way into Europe

gave rise to marker M343, the defining marker of haplogroup

R1b. These travelers are direct descendants of the people who

dominated the human expansion into Europe, the Cro-Magnon”

(Spencer Wells, “Deep Ancestry”, 2006). This and similar claims,

such as R1b (and its M269 subclades) were “well established

throughout Paleolithic Europe”, “contemporaneous with Aurig-

nacian culture”, “the earliest expansion into Europe, during the

Upper Paleolithic ~30,000 years ago” by Wells, Semino, Un-

derhill, Cavalli-Sforza, Cinnioglu, Kivisild, Wiik and many

others (e.g., Semino et al., 2000; Wells et al., 2001; Cinnioglu

et al., 2004; Wiik, 2008) were essentially based on “thoughts”

that if people lived in Europe some 30,000 years ago, they nec-

essarily were of the R1b haplogroup, and not of I, G, J, E, F or

any other haplogroups. Were any haplotypes analyzed? Their

mutations counted? Any chronological evaluations? There was

nothing of the above.

“Paleolithic origin” of R1b in Europe, or their “Paleolithic

migrations” to Europe around 10,000 - 8000 ybp are still claimed

A. A. KLYOSOV

in recent academic papers, such as (Myres et al., 2010; Balares-

que et al., 2010; Morelli et al., 2010). These assumptions and

resulting calculations are based, typically, on “population muta-

tion rates” (Zhivotovsky et al., 2004; Hammer et al., 2009;

Underhill et al., 2009), which commonly exaggerate the chro-

nological estimates of migrations and events by 200% - 400%

since they are based on crude, artificial and unrealistically naïve

and generalized reasoning (for critique, see Klyosov, 2009a,

2009b, 2009c; Rozhanskii & Klyosov, 2011). The “academic

papers”, placing origin of R1b in Asia Minor or nearby, have

not considered regions east of Asia Minor as well as R1b hap-

lotypes of those eastern regions. In short, the whole story of

R1b migrations and their history is in disarray, and the “popu-

lation geneticists” continue to advance misleading conclusions

due to their methodology.

This study’s methodology/analysis includes considerations of

Y chromosome extended 67 and 111 marker haplotypes, when

available. The methodology was described in detail in the pre-

ceding papers in this journal (Rozhanskii & Klyosov, 2011;

Klyosov & Rozhanskii, 2012a) and elsewhere (e.g., Klyosov,

2009a, 2009c, 2009d), and in Materials and Methods section of

this article.

As described (Klyosov & Rozhanskii, 2012b), Europeoids

(Caucasoids) appeared ~58,000 ybp. They gradually branched

to downstream haplogroups and their subclades, and migrated

to the north, west, south and east. Haplogroup NOP, which was

among them, arose ~48,000 ybp, and moved eastward, pre-

sumably towards South Siberia and/or adjacent regions. Hap-

logroup P split off ~38,000 ybp, presumably in South Siberia,

and gave rise to haplogroup R and then R1 ~30,000 - 26,000

ybp (see the diagram in Klyosov and Rozhanskii, 2012b). Hap-

logroup R1b arose ~16,000 ybp, as it will be shown further in

this paper.

The timing may be reconstructed from a series of R1b hap-

lotypes, made available from the databases (see Appendix). The

most distant R1b haplotypes (those exhibiting the greatest mu-

tational differences) from European R1b haplotypes, were

found in Siberia and Middle Asia (a part of Central Asia)

populations. Central Asian R1b haplotype bearers have the

most ancient common ancestors with European R1b bearers,

and those ancient common ancestors lived ~16,000 ybp in Cen-

tral Asia. We do not know as yet whether in South Siberia or

Middle Asia; however, the evidence will demonstrate that it

was somewhere in that vast region.

In this endeavor both terms, “haplogroup” and “subclade”,

are employed as near equivalents because all haplogroups are

essentially subclades of other upstream haplogroups, and usage

of one or the other of these terms is suggested by the context.

This is shown in the following diagram (ISOGG-2012, a frag-

ment, http://www.isogg.org/tree/ISOGG_HapgrpR.html) which

is related to the most ancient subclades of R1b haplogroup:

R1b M343

R1b1 L278, M415, P25_1, P25_2, P25_3

R1b1* -

R1b1a P297, L320

R1b1a1 M73, M478

R1b1a2 L265, M269, M520, S3, S10

R1b1b M335

R1b1c V88

R1b1c* -

R1b1c1 M18

R1b1c4 V69

Due to lack of a common name for bearers of R1b hap-

logroup (with subclades) and their languages in ancient times,

which were carried for millennia and eventually brought to

Europe as non-Indo European languages, I refer to them as the

Arbins (from R1b), both the people and their original languages,

on the analogy with the Aryans, who essentially belonged to

haplogroup R1a (Klyosov & Rozhanskii, 2012a).

A Bird’s-Eye View at R1b Haplotypes, and the

Most Ancient R1b Populations

Figure 1 presenting an overview of an R1b haplotype tree

includes 338 haplotypes in a short 8 marker format. The pur-

pose of the presentation is to show the complex pattern of the

R1b haplogroup and identify the most ancient branch on the

tree.

The haplotype tree as shown next is arranged by a computer

program which combines branches based on the similarity of

their alleles in the respective markers (or loci) in Y chromo-

some, and dynamics of their alleles (Klyosov, 2009c and refer-

ences therein). Haplotypes identical to each other and prevail-

Figure 1.

The 8-marker 338 haplotype tree for R1b1 haplotypes, composed from

data listed in (Zhong et al., 2010). The tree includes haplotypes of sub-

clade R1b* (8 haplotypes), R1b1* (2 haplotypes), R1b1c (2 haplotypes),

R1b (140 haplotypes), R1b1a1 (12 haplotypes), R1b2b (21 haplotypes),

R1b1a2 (153 haplotypes). Regarding ethnicity and/or location, the

haplotypes belong to 40 Western Europeans, 17 Iberians, 84 Turks, 48

Albanians, Macedonians and Romanians, 40 Italians, 30 Cretans, 13

Greeks and Macedonians, 9 Romanians from a different region, 13

Pakistanis, 4 Indians, 2 Japanese, 2 Mongolians (one of them, No. 176,

belongs to R1b1b1-M73 subclade and stands out in the lower part of

the tree), 2 Tibetans, 1 Kyrgyz from Xinjiang, 7 Han Chinese, 16

Uighur (12 from the Xinjiang Province, six of them belong to R1b*

subclade, three R1b1b1 and three R1b1b2); four Chinese (three R1b2b

and one R1b1b2), 2 Chinese of the Hui tribe, and 6 Chinese of other

ethnicities (Hazak R1b1b1, Tu R1b2b, and four Naxi R1b2b).

A. A. KLYOSOV

ing in the dataset are sitting at the top of the tree; in relatively

1b branch has

its

the tree shows that many of the Asian R1b

however, form separate branches. The

Although the M73 suned a few steps from

recent datasets they typically represent the “base”, or “ances-

tral” haplotype, from which all other haplotype are derived

through mutations. Branches which are located close to the

“trunk” of the tree contain not many mutations from the base

haplotype, hence, they are relatively “young”. The most ancient

branches are those which shoot away from the trunk since they

contain the most mutations in their haplotypes.

In Figure 1 that rather recent and prevailing R

base haplotype 13 24 14 11 X X X 12 X 13 13 29. Thirty-

five of those identical haplotypes sit on the top of the tree. Here

X stand for missing alleles in the haplotype presented in the

standard twelve marker FTDNA format (for definitions see

Materials and Methods). This base haplotype belongs to the

most populous European base haplotypes of the subclade

R-M269, which in turn includes subclade R-P312 and its many

downstream subclades. For example, the base 67 marker hap-

lotype of P312 subclade is as follows (Klyosov, 2011b):

13 24 14 11 11 14 12 12 12 13 13 29 - 17 9 10 11 11 25

15 19 29 15 15 17 17 - 11 11 19 23 15 15 18 17 36 38 12

12 - 11 9 15 16 8 10 10 8 10 10 12 23 23 16 10 12 12 15 8

12 22 20 13 12 11 13 11 11 12 12 (P312)

The alleles identical to those of the apparent “base” haplo-

pe in the tree in Figure 1 are marked in bold. Indeed, the

subclade R1b1a2-M269 takes the largest part of the tree in

Figure 1, since “R1b” as indicated in the legend is largely

R-M269 as well.

The analysis of

plotypes, particularly all the Indian, most of Pakistani, a

quarter of Uighurs, one Japanese, one Tibetan, and two Chinese

Hans are not the “indigenous” haplotypes but those which are

closely associated with European haplotypes and share with

them the same branches (Klyosov, 2010a). In other words, they

are “returns” to the regions with Europeans, or having the Eu-

ropean R1b origin. They are indistinguishable from the Euro-

pean R1b haplotypes.

Some of haplotypes,

ost remarkable branch is most remote from the trunk, hence,

the most ancient, is located in the upper left part of the tree. It

contains 12 haplotypes which all belong to R1b1a1-M73 and

R1b-M343 subclades, and provided by Uighurs and the close

tribes of the Naxi, Han and Tu (Zhong et al., 2010). All the 12

haplotypes are derived from the base haplotype 13 24 15 11/10

X X X 12 X 14/13 12 30, and contain collectively 65 mutations

from it. Using the “linear” formula, we obtain 65/12/.013 = 417

→ 619 conditional generations (the arrow here is a correction

for back mutations), or 15,475 ± 2500 years from their common

ancestor (the calculation is explained in the Materials and

Methods section). This date is in a fair agreement with 16,000 ±

1400 years from a common ancestor of R1b haplotypes (Klyo-

sov, 2008a, 2009d) and will be additionally confirmed later in

this paper. It also indicates that the R1b haplogroup arose in

Central Asia, and, apparently, in the Altay region in South Si-

beria, where their upstream haplogroups NOP → NO + P; P →

R + Q; R → R1 + R2; R1 → R1a + R1b all migrated and lived

there during the time period between 52 and 20-15 thousand

years before present (Klyosov & Rozhanskii, 2012b).

Siberian, Bashkir, and Central Asian R1b-M73

Haplotypes

bclade is positio

e top of the above diagram, it contains the most distant avail-

able haplotypes from the Europeans, both geographically and

by their mutations. Figures 2 and 3 show two M73 haplotype

trees composed from two different datasets; one was obtained

directly from the researchers and published earlier (Klyosov,

2008a), the second was collected from the R1b1b1 Haplogroup

Project (see Figure 3 and references). Both of them have prin-

cipally the same shape, with two primary branches in Figure 2

and three in Figure 3.

Figure 2.

er 14 haplotype tree for R1b1-M73 haplotypes in Central The 25-mark

Asia (Uzbeks, Tajiks, Tuva, Uighurs, Kazakhs) (Klyosov, 2008a).

Figure 3.

er 25 haplotype tree for R1b1a1-M73 haplotypes: the right-

?section=yre

The 25-mark

hand side branch—five Russians, three Kazakhs, one Chinese; the

lower branch—four Uzbeks, two Tajiks, one Turk, one from Luxem-

bourg; the left-hand branch—two English, two Italians, one French and

one Spanish. The data are taken from the FTDNA Project, in which

many haplotypes are available in the 67 marker format:

http://www.familytreedna.com/public/R1b1b1/default.aspx

sults.

A. A. KLYOSOV

The tight 8-haplotype branch close to the “trunk” of the tree

9 11 11 23 15

imilar series of ten

haplo-

e tree

11 13 17 12 12 12 13 13 30 - 15 9 9 11 11 23 15

and

ure 3 has the following base

14 12 12 12 14 13 29 - 16 9 10 11 11 22

, they all are

X 13 14 13 30 - 15 10

d Siberian-Ural-

Figure 2 and the 9-haplotype right-hand branch in Figure 3

both have the same base haplotype as follows:

13 19 14 11 13 13 12 12 13 14 13 30 - 17 9

19 33 12 15 15 16 - 10 10 19 25 15 16 16 17 30 36 12 10

- 11 8 16 16 8 10 10 8 11 10 12 23 23 16 10 12 12 16 8 12

24 21 13 12 11 13 11 11 12 11 (M73, branch)

The remarkable feature of this base haplotype (and ever

plotype in the both branches) is the second allele, DYS390 =

19. In the 25 marker format, the branches contain 15 and 17

mutations respectively, which gives the same 1075 ± 280 years

to the common ancestor in both instances.

It should be mentioned here that a s

-M73 haplotypes (with DYS 390 = 19) were obtained from

the Bashkir population (Myres et al., 2010) near the border

between Europe and Asia, on both sides of the Ural Mountains.

Their base haplotype was 13 19 14 10 X X X 12 13 14 13 29 -

10 (the last marker is DYS461), and all hundred alleles in the

ten haplotypes contained only three mutations; this gives

3/10/.018 = 17 generations, or 425 ± 250 years to their common

ancestor. However, as it will be shown below, a common an-

cestor of these and other R-M73 haplotypes lived more that 10

thousand years ago. What we see in these haplotypes are essen-

tially “tips of the iceberg;” they are tremendously distant from

each other and must be very remote from their common ances-

tor in time. It seems that the “base” of the proverbial iceberg

vanished without leaving their ancient haplotypes behind. In

other words, we see here a series of present-day haplotypes

which either arrived from elsewhere a few centuries earlier, or

went through a severe bottleneck(s) of the population.

The Bashkir’s (similar or slightly mutated) R-M73

pes were identified among Tatars and Mari populations, in the

adjacent regions (ibid.). In the same study, the R-M73 were

found also among Kabardino-Balkars, Mengrels, Turks, with

all speaking Turkic or Uralic languages, which belong to the

Altay language family. Another name for Mari people is

Cheremis, meaning “people from the East” in a neighboring

Komi language. Their base haplotype 13 22 14 11 X X X 12 13

14 13 32 - 10 differs by 7 mutations (!) in ten markers from the

Bashkir base L23 haplotype, which places their common an-

cestor by 9700 ybp (Klyosov, 2011a), and if haplotypes with

DYS390 = 19 are included, their common ancestor moves

deeper in time, to at least 16,000 years before present.

Revisiting the tree in Figure 3, the lower branch in th

s the following base haplotype (when extended to 67 mark-

ers)

13 22 14

20 30 12 15 15 17 - 11 10 19 24 15 14 17 17 35 39 12 10

- 11 8 16 16 8 10 10 8 10 10 12 23 23 15 10 12 12 18 9 12

25 20 13 12 11 13 11 11 12 12 (M73, branch)

its common ancestor lived 3525 ± 600 ybp. It corresponds

to the right branch in Figure 2.

The left-hand side branch in Fig

plotype:

12 25 14 11 13

15 20 32 12 15 16 17 - 11 10 19 25 15 17 17 18 34 36 14

10 - 11 8 16 16 8 11 10 8 11 10 12 22 23 17 10 12 12 15 8

13 22 20 14 12 11 14 11 11 12 12 (M73, branch)

d its common ancestor lived 3050 ± 600 ybp.

All three branches are rather “young”, however

very distant from each other. Their pair-wise mutational differ-

ences are 38, 38 and 42, which translates into the separation

time between their common ancestors as 11,500 - 13,200 years

of the mutational evolution in their haplotypes (we call it the

“lateral time” between two base haplotypes). All three M73

base haplotypes differ from each other collectively by 62 muta-

tions, which place their common ancestor at ~7750 ybp. How-

ever, their mutational difference is even more pronounced when

compared with the base haplotype of R1b1a2-P312 subclade,

ancestral to many European subclades of R1b1a2 haplotypes.

For the above M73 base haplotypes it amounts to 48, 35 and 42

mutations, or up to 16,000 years of the mutational difference

between the two base haplotypes. This places their common

ancestor at 10,600 ybp.

Another series of 26 of 12 marker M73 haplotypes was listed

in (Malyarchuk et al., 2011), and were collected among the

Siberian populations: Shors, Teleuts, Kalmyks, Khakass, Tuva,

and Altayans. Those haplotypes split into two branches. The

first one fits exactly to the first 67 marker base haplotypes with

their available 12 markers; the second deviates from the base

haplotype of the respective right-hand branch in Figure 2 in

two alleles (marked):

13 19 14 11 13 13 X

13 22 14 11 13 16 X X 13 13 13 30 - 15 10

The first 12 marker base haplotype is a mixe

aucasian population (if to consider their current bearers—Bash-

kirs, Kalmyks, Tuva, Tatar, Kabardin, Russian, and Altayan).

All 11 haplotypes of this branch have collectively 20 mutations,

which gives 20/11/.024 = 76 → 83 conditional generations, or,

2075 ± 510 years to a common ancestor. The logarithmic

method gives [ln(11/2)]/.024 = 71 → 77 generations, or 1925

years, which is nearly the same (2 haplotypes are identical in

their base haplotypes among 11 haplotypes in the dataset. .024

is the mutation rate constant for the 12 marker haplotypes). The

same but in 67 markers, Middle Asian M73 haplotypes showed

1075 ± 280 years to the common ancestor. Thus, the Siberia-

Ural-Caucasian dataset is somewhat “older” compared to the

Middle Asian dataset.

The second 12 marker base haplotype was largely from Sibe-

rian populations (Shors, Teleuts, Khakasses), plus a singular

Mari, Turk and Tatar each. All 15 haplotypes of this branch had

22 mutations, which gives 22/15/.024 = 61 → 65 conditional

generations; or 1625 ± 380 years to a common ancestor. The

logarithmic method gives [ln(15/5)]/.024 = 46 → 48 genera-

tions, or 1200 ± 580 years, which is the same within the margin

of error. The 67 marker base haplotype, however showed 3525

± 600 years to the common ancestor. Hence, now the Middle-

Asian M73 branch is “older” compared to the Siberian. Overall,

the Siberian and Middle Asian haplotypes were in about the

same estimate of “age”.

The two 12 marker haplotypes differ by 8 mutations, which

sets their two common ancestors apart by 8/.024 = 333 → 492

generations, or 12,300 years. It gives (12,300+2075+1625)/2 =

7985 ybp to their common ancestor. One can see that the data

obtained with the 67 and 12 marker haplotypes of the M73

subclade from Siberia and Middle Asia are fairly reproducible,

and point to a common ancestor of Central Asian M73 haplo-

types nearly 8000 ybp. It should be kept in mind that these data

A. A. KLYOSOV

are related to currently living descendants of those common

ancestors. We also need to be mindful that the 67 marker M73

Central Asian haplotype examples a distance up to 48 muta-

tions from the European R1b1a2 haplotypes, which is up to

16,000 years of the mutational difference between the two base

haplotypes. This places their (R-M73 in Central Asia and

R-M269 in Europe) common ancestor at 10,600 years before

present.

R1 *

b1-L278/R1b1 Subclade Haplotypes

The the

Projec M73

are represented in

most ancient subclades of R1b are considered by

t R1b1 (xP297); xP297 does not include subclades

d M269, discussed in the preceding section of this article. All

68 haplotypes of the Project which are available in the 67

marker format are shown in the tree in Figure 4. The tree in-

cludes haplotypes of subclade R1b1*, a para-group of R1b1-L278

(see the R1b diagram exampled previously), V88, its down-

stream subclade V69, and a series of haplotypes R1b1a2-M269

which are in the Project apparently by mistake (however, useful

in the tree as a reference branch).

A detailed analysis of the tree was performed in (Klyosov,

2011a). Haplotypes of the R1b1* para-group

e Project by either Ashkenazi Jews (from Hungary, Russia,

Belarus) with their very recent common ancestors a few centu-

ries ago, or by very assorted geography—from Italy, Puerto-Rico,

Germany, Armenia, Turkey. Formally, a triple branch on the

right of Figure 4 (haplotypes 1 - 10) has a common ancestor

who lived about 5800 ybp. It does not carry any certain histori-

cal meaning judging from their composition except, the time

span corresponds to that of R-L23 subclade in the Caucasus and

Anatolia (Klyosov, 2010d); R-L23 was certainly not alone there.

Those assorted haplotypes could have been brought by their

ancestors from Central Asia along with R-M269 followed by

Figure 4.

67 marker haplotype tree of 68 haplotypes of subclades R1b1

1c-V88 (18 - 41), R1b1c4-V88-V69 (42 - 47), R1b1a2

L23

9 9 11 11

in the first 255

-V88 Subclade in Europe and Africa

Hig-

ure 4

16/17 9 10 11 12

(1 - 17), R1b

(48 - 68). The tree was composed from data of the FTDNA

Project

http://www.familytreedna.com/public/R1b1Asterisk/default.asp

x?section=yresults.

haplotypes, and eventually dispersed across Europe.

he next small braTnch of six haplotypes (12 - 17) includes

those from Central Asia (Uzbek), Armenia, Middle East (Iraq

and Bahrain), India, and Germany (the last being relatively

close to the Uzbek haplotype, within 4900 years of the muta-

tional evolution). The Indian haplotype has DYS390 = 18,

which brings it closer to the Central Asian haplotypes with

DYS390 = 19, considered above. Others have DYS390 = 24 or

25. Their mutational deviations (without inclusion of the very

distant haplotype) from base R1b1* haplotype

13 24 15 10 12 13/14 12 13 11/12 13 13 29 - 14

26 14/15 18 30 13/14 14 14/15 16 - 11 11 18 23 15 16 18

16 36 36 13 11 - 11 8 15 16 8 10 10 8 11 10 12 19 21 16

10 12 12 14/15 8 12 24 20/21 14 11 11 13 12 11 12 12

(R1b1*, branch)

, 37, and 67 markers suggests 238, 217, and 23

generations from their common ancestor (without a correction

for back mutations), that is 230 ± 11 generations, a fairly re-

producible value. The most reliable value, for the 67 marker

format, gives ~8400 years to their common ancestor. However,

the Indian haplotype differs from the above base haplotype by

62.5 mutations (!), with its DYS390 = 18 contributing only 6

mutations into the difference. This places a timespan to the

R1b1* common ancestor (of the available haplotypes) to 16,600

years before present. It certainly points to Central Asia, since

the European-Middle Eastern R1b1* haplotypes are only ~8400

years “old”

R1b1c

aplotypes V88 occupy the left-hand side of the tree in F

. Again, a number of their sub-branches are quite “young”.

For example, a part of the branch belongs to Ashkenazi Jews

(Russia, Ukraine, Hungary, Germany, France) with a common

ancestor of only 350 ybp. Haplotypes of England and Scotland

have a common ancestor who lived 650 ybp. However, a com-

mon ancestor of the two sub-branches (Jews and the Isles) lived

6875 ybp, at the very bottom of the V88 subclade. Another

series of V88 haplotypes identifies their route from Saudi Ara-

bia (6225 ybp) via Spain to the Jewish community (1525 ybp,

in the middle of the 1st millennium CE). The base haplotype of

the V88 branch in Figure 4 is as follows:

13 24 16 10 13 14 12 12 12 14 13 29 -

26 14 19 29 12 12 15 15 - 11 12 21 23 15 15 18 19 33

35/36 12 11 - 12 8 15 16 8 10 10 8 9 11 12 22 23 15 11 12

12 15 8 12/13 23 20 13 12 11 13 11 12 12 12 (V88)

d its bearer lived 6575 ± 700 ybp.

V88 haplotypA rather extended series of 72 of 11 marker

from Africa was provided in (Cruciani et al., 2010). They split

into two rather distinct branches and an assorted series of hap-

lotypes (Klyosov, 2010b). One branch of 37 haplotypes has the

following base haplotype (in the format—413a,b 460, 461,

GATA A10, YCAIIa,b after the first standard 12 markers, some

of them are missing): 13 X 15 11 X X X X 12 X X X - 23 21 11

10 13 21 23 All of them contain 111 mutations, which gives

111/37/.02 = 150 → 176 conditional generations, or 4400 ± 610

years to their common ancestor. This African V88 base haplo-

type does not fit with either of the Eurasian V88 base haplo-

types, however, it fits rather well to the R1b1c4-V69 base hap-

lotype (see the next section), whose common ancestor lived

A. A. KLYOSOV

4300 ± 600 ybp. The paper (Cruciani et al, 2010) suggested one

more subclade named R1b1a4, with 29 haplotypes. Their base

haplotype is identical with that shown above; all 29 haplotypes

contain 76 mutations, and gives 76/29/.02 = 131 → 150 genera-

tions, or 3750 ± 570 ybp, which is within the margin of error

with the above date.

Another branch of V88 African haplotypes is recent with a

common ancestor of 750 ± 290 ybp. All 72 African haplotypes

stward along the

This subclade es at the bottom

of the tree in Fig

16 17 19 33

ha r

ison with the entire R1b, and R-V69 is

“y

Subclade

The last brane lower right-

hand side has th

23 15 15 18 17 37 38 12

R-

ublications (Klyosov, 2008a, 2009d,

ting from Central Asia westward by

e trees

15 15 19 17 35 38 12

ve the same base haplotype as shown above with 201 muta-

tions from it, which gives 201/72/.020 = 140 → 163 genera-

tions, or 4075 ± 500 ybp, within the margin of error of that for

the main V88 branch; however, the first is more accurate since

it was calculated for the distinct branch.

This date, 4300 ± 600 ybp, fits well with that for the migration

route of R1b bearers from the Middle East we

editerranean Sea coast to the Atlantic, which took place between

5500 and 4800 ybp (see below). It seems that the R1b-V88-V69

tribe may have split altering migration direction south, to Camer-

oon and Chad, where R1b-V88 bearers live today.

R1b1c4-V69 Subclade

is represented by six haplotyp

ure 4, with the base haplotype

13 23 15 11 13 15 12 12 13 13 13 29 - 16/17 9 10 11 12

27 14 19 29 12 12 14 15 - 11 12 21 23 15

36/37 12 12 - 12 8 15 16 8 10 10 8 10 11 12 21 24 15 11

12 12 14 8 12/13 22 20 13 12 11 13 11 12 12 13 (V69)

All of them collectively have 106 mutations from the bas

plotype, which gives 106/6/.12 = 147 → 172 generations, o

00 ± 600 years from a common ancestor. This is in agree-

ment with the “age” of the upstream V88 subclade, of ~6575

ybp. The V69 base haplotype differs by 16 mutations from the

base V88 haplotype (see above; some mutations are fractional),

which sets apart their common ancestors by 16/.12 = 133 →

154 generations, or 3850 years, and places their common an-

cestor at (3850 + 6575 + 4300)/2 = 7400 ± 800 ybp. This is

within the margin of error with 6575 ± 700 ybp for the V88

upstream subclade.

The analysis demonstrates that R-V88 is a rather “young”

subclade in compar

ounger”. It may have arisen on the R1b general migration

route to the Middle East, for example, north-east or east of the

Caspian Sea, in the Western Iran.

R1b1a2-M269

ch on the tree in Figure 4 in th

e base haplotype

13 24 14 11 11 14 12 12 12 13 13 29 - 17 9 10 11 11 25

15 19 29 15 15 17 17 - 11 11 19

12 - 11 9 15 16 8 10 10 8 10 10 12 23 23 16 10 12 12 15 8

12 22 20 13 12 11 13 11 11 12 12 (M269, a mix)

It is exactly the base haplotype of subclades R-P312 and

L21. They are typical and widespread European haplotypes.

ll 20 haplotypes of the branch contain 365 mutations, which

gives 365/20/.12 = 152 → 179 generations, or 4475 ± 505 years

to their common ancestor. This, again, is a typical “age” for a

mix of haplotypes of subclades R-P312 and R-L21 and their

downstream subclades.

The M269 subclade along with its downstream subclades

were analyzed in many p

10c, 2010d, 2010e, 2011a, 2011b). In short, this subclade

arose around 7000 ybp in Central Asia or in the eastern area of

the Russian Plain—its immediate downstream subclade R-L23

(arose ~6000 ybp) is widespread among the Bashkirs in the

South Urals, North Kazakhstan and adjacent regions, and can

be seen in Russia. It migrated south to the Caucasus and be-

yond, to Anatolia and the Middle East. Nearly all principal

European and Middle Eastern subclades of the R1b haplogroup

(currently more than 80 subclades) are derived from the

R1b1a2-M269 subclade.

It should be mentioned here, that two brother subclades, R1a

and R1b, have been migra

o quite different routes. While R1a were moving along the

southern route from the Altay region across the Himalayas,

Hindustan, the Iranian plateau, Anatolia and the balance of Asia

Minor to the Balkans (Klyosov & Rozhanskii, 2012), R1b were

moving along the northern route, from the same region across

South Urals, Middle Asia, North Kazakhstan, Middle Volga,

the Caucasus, and then split between the southward and west-

ward directions. This migration pattern also explains why R1a,

but not R1b bearers were found in the Eastern Himalayas (Kang

et al., 2011; see also Klyosov & Rozhanskii, 2012).

37 and 67 marker haplotype trees of the R1b1a2*-M269 sub-

clade are shown in Figures 5 and 6, respectively. Th

ntain two distinct branches, one rather complex and obvi-

ously “older”, with the base haplotype

12 25 14 11 11 14 11 12 12 13 13 29 - 17 9 10 11 11 25

15 19 29 15 15 16 17 - 10 11 19 23

Figure 5.

The 37 marker 25-haplotype tree for R1b1a2*-M269 haplotype

6225 ybp) branch on the left and the bottom pre-

The older (~

sent haplotypes from Turkey, Armenia, Italy, France, England.

The younger (1000 ybp) branch in the upper right side repre-

sents Ukraine, Belarus, Poland, and Kazakhstan.

http://www.familytreedna.com/public/ht35new/default.aspx

A. A. KLYOSOV

Figure 6.

The 67 marker haplotype tree for R1b1a2*-M269 haplotypes, the sam

own in Figure 5, however, with fewer haplotypes avail-

- 11 9 15 16 8 10 10 8 11 10 12 23 23 16 10 12 12 15 8

12 12 13 14 29 - 17 9 10 11 11 25

younger

mong

R-L23 appar

Plain, where Eur

irs, a Turkic-language people who live

entral Asia, apparently from the

stic DYS393 = 12). In a

15 16 17 - 11 11 19 23 16 15 17 17 36 37 12

act

11 19 23 16 16 18 17 36 37 12

dataset as sh

able.

13 22 20 13 12 11 13 11 11 12 12 (M269)

d another, a tight, flat, “younger”, and largely Jewish branch,

with the base haplotype.

12 24 14 10 11 14 11

15 19 30 15 15 16 16 - 11 10 19 23 17 16 17 17 38 39 13

12 - 11 9 15 16 8 10 10 8 11 10 12 21 23 16 10 12 12 14 8

12 22 20 13 12 11 13 10 11 12 12 (M269, Jewish)

Mutation deviations between the two are shown in bold. The

branch has 26 mutations in all six 67 marker haplo-

types, which gives 26/6/.12 = 36 → 37 conditional generations,

or 925 ± 200 years to a common ancestor. The 37 marker

branch contains 31 mutation in nine haplotypes, which gives

31/9/.09 = 38 → 40 generations, or 1000 ± 200 years—essen-

tially the same. This Jewish population exemplified with 14 of

37 marker haplotypes was analyzed earlier (Klyosov, 2008b)

and determined that their common ancestor lived 1100 ± 250

ybp; the base haplotype is exactly as that of the “young” branch

listed above.

The older branch has a common ancestor who lived 6200 ±

900 ybp. The two base haplotypes differ by 22 mutations which

separate them by 22/.12 = 183 → 224 generations, or 5600

years, and a common ancestor of both branches lived 6400 ±

900 ybp.

This description gives us a general picture of migration of the

Arbins from Central Asia (~16,000 ybp) westward during the

next 10 - 9 thousand years, to about 7000 - 6000 ybp.

Haplogroup R1b (mainly R1b1a2-L23) a

Bashkirs, and in the Caucasus, Anatolia,

Middle East

ently arose on the eastern side of the Russian

ope meets Asia, ~6200 ybp and migrated to the

Caucasus and further South, to Anatolia and the Middle East.

Another branch of L23 went westward, to Europe, approxi-

mately 4500 ybp.

The most eastern population with the prevailing R-L23 sub-

clade is the Bashk

rgely on both sides of the Ural Mountains and in North Ka-

zakhstan. Frequency of R1b haplogroup reaches 84% in the

Perm Bashkirs, 81% among Baimak Bashkirs, and lower fig-

ures in other Bashkir tribes (Lobov, 2009). 29 of 10 marker

haplotypes of subclade R-L23 of the Bashkirs were published

(Myres et al., 2010), and their base haplotype is 12 24 14 10 X

X X 12 12 13 13 30 - 10. This is a typical albeit slightly mu-

tated L23 haplotype with its characteristic first allele DYS393 =

12. 26 haplotypes of those 29 were identical, as shown above,

and the whole Bashkir L23 branch has a common ancestor who

lived only 575 ± 175 ybp. However, this base haplotype differs

from the European R-L23 base haplotype 12 24 14 10 X X X

12 12 13 14 29 - 11 by three mutations, which sets these two

base haplotypes apart by 3/.018 = 167 → 200 conditional gen-

erations—5000 years, and places their common ancestor at

5500 ybp. Other Bashkir haplotypes belonged to R-M269 (one

haplotype), R-M73 (10 haplotypes), and R-U152 (8 haplotypes).

The last series of haplotypes were all identical to each other,

and thus are derived from a very recent common ancestor, who

certainly had a European origin [U152 arose in Europe 4125 ±

450 ybp (Klyosov, 2011b)].

We therefore see today’s reflection of ancient migrations of

the Arbins westward from C

uth Siberian region, across the South Urals and further to the

Russian Plain and then the Caucasus.

Almost all R1b1a2 haplotypes in the Caucasus region belong

to the subclade L23 (with its characteri

cent paper (Balanovsky et al., 2012) 90 Caucasian haplotypes

of R1b haplogroup were listed, and with exception of five R1b*

haplotypes and a relatively “young” Abkhazian branch (Figure

7) 79 of 81 haplotypes (97.5%) in the dataset were of the L23

subclade (Note: the cited paper did not consider haplotype trees

nor has analyzed the haplotypes in the manner presented here).

The same pattern is observed with Armenian R1b haplotypes,

and with most of Anatolian R1b haplotypes (Klyosov, 2010c,

11c). The 67 marker base R-L23 haplotype, obtained from an

extended haplotype dataset from the world over (tree Figure 8)

is as follows:

12 24 14 11 11 14 12 12 12 13 13 29 - 16 9 10 11 11 25

15 19 29 15

12 - 11 9 15 16 8 10 10 8 10 11 12 23 23 16 10 12 12 15 8

12 22 20 13 12 11 13 11 11 12 12 (L23)

The L23 base short Caucasian haplotype in Figure 7 fits ex-

ly to the above haplotype (the matching alleles are marked in

ld). A common ancestor of the L23 subclade lived ~6200 ybp

(Klyosov, 2010d, 2011a). 81 Caucasian L23 haplotypes con-

taining 425 mutations from their base haplotype, give 425/

81/.035 = 150 → 176 conditional generations, or 4400 ± 490

years to their common ancestor. The “younger” date (compared

with the “age” of L23 of about 6200 ybp) can be explained by a

detailed consideration of an extended series of 107 of R-L23

haplotypes listed in the FTDNA Project (see the legend in Fig-

ure 8). The tree in Figure 8 splits into two parts. On the left are

38 haplotypes, with the base

12 24 14 11 12 15 12 12 12 13 13 29 - 16 9 10 11 11 25

15 19 30 15 15 16 18 - 11

12 - 11 9 15 16 8 10 10 8 11 11 12 23 23 15 10 12 12 16 8

12 22 20 13 12 11 13 11 11 12 12 (L23, branch)

A. A. KLYOSOV

Figure 7.

The 19-marker 90 haplotype tree for R1b haplotypes in the Caucasus (Lezghins, Osset

Circassians, Chechens, Dargins, Kaitaks). Haplotypes 1 - 5 (above on

On the right are 69 haplo

12 24 14 11 11 14 12 12 12 13 13 29 - 16 9 10 11 11 25

17 36 38 12

fro

9 30 15 15 16 18 - 11 11 19 23 16 16 18 17 36 37 12

10 30 12 13 24 13 10 10

18 10 14 17 9 11 11

br d

th/2 =

5525 ± 700 years—similar to 5400 ± 800 ybpe,

Avars, Ankhaz,

the left) represent R1b subclade, and haplotypes 49, 57 - 59 (above on the left) repre-

sent a “young” Abkhazian branch. Haplotypes were listed in (Balanovsky et al., 2012)

types, with the base 19 12 11 13 12 10 9 12 12 10 11

19 15 19 13 24 17 12 15 24 12 23

15 19 29 15 15 16 17 - 11 11 19 23 15 16 18

12 - 11 9 15 16 8 10 10 8 10 11 12 23 23 16 10 12 12 15 8

12 22 20 13 12 11 13 11 11 12 12 (L23, branch)

More than 70% of haplotypes of the Armenians and Turk

m the dataset belong to the second and larger branch as well

all eight Iraqis and all five Iranians in the dataset. Presence

of R-L23 of the Iranians might be a result of diffusion of the

subclade from Anatolia eastward, or the migration of the Arbins

might have been southward from the Russian Plain east of the

Caspian Sea.

Both branches descended from their ancestral R-L23 base

haplotype, and are parted by 9 mutations (marked in bold).

These 9 mutations are accumulated over 9/.12 = 75 → 81 con-

ditional generations, or 2025 “lateral” years. The first branch

split 4600 ± 490 ybp; the second, 4200 ± 440 ybp. Therefore,

their common ancestor lived (4600 + 4200 + 2025)/2 = 5400 ±

800 ybp. This fits within margin of error to the time when a

common ancestor of the subclade R-L23 lived (~6200 ybp).

Extended, 111 marker haplotypes available for the same data-

set, and for the smaller branch the base haplotype is as fol-

lows:

12 24 14 11 12 15 12 12 12 13 13 29 - 16 9 10 11 11 25

15 1

12 - 11 9 15 16 8 10 10 8 11 11 12 23 23 15 10 12 12 16 8

12 22 20 13 12 11 13 11 11 12 12 - 36 15 9 15 12 25 26

(L23, branch)

It differs by 16 mutations from that of the larger branch (

utations added by the 68 - 111 extension), which separates th

anches by 16/.198 = 82 → 90 generations, or 2250 years, an

e R-L23 common ancestor lived (4600 + 4200 + 2250)

, obtained abov

d illustrates the consistency of calculations.

It seems that the Caucasian R-L23 haplotypes with their

common ancestor of 4400 ± 490 years belong to one of the

branches in the tree in Figure 8. A much smaller Caucasian

R1b dataset, analyzed earlier (Klyosov, 2008a) resulted with

similar times—4650 ± 700 ybp, as the recent, larger dataset of

short haplotypes (Balanovsky et al., 2012). The Caucasian

R-L23 haplotypes may have experienced a population bottle-

neck around 5000 ybp.

120 of 17 marker Armenian haplotypes were published re-

cently in (Herrera et al., 2011). A haplotype tree, composed

from those haplotypes, is shown in Figure 9. It consists of three

approximately equal (by number, or by “weight”) branches.

Five haplotypes of the M269 mini-branch are nearly equal to

each other, evidencing only two mutations among their 85 al-

leles. Their common ancestor was 300 ± 210 ybp. The balance

of 115 haplotypes of L23 subclade, have 784 mutations from

their base haplotype 12 24 14 11 11 14 X X 13 13 13 29 - 17 15

19 12 15 12 23 (the right-hand side of the haplotype corre-

sponds to DYS 458, 437, 448, GATA H4, DYS 456, 438, 635).

A. A. KLYOSOV

Figure 8.

The 67-marker tree composed with 107 haplotypes of the subclade R-L23. 40% of the haplotypes

belong to Armenians and Turks, followed by Iraqis and Italians (5% each). The haplotypes were

roject http://www.familytreedna.com/public/ht35new/default.aspx

It gives 784/115/.034

6250 ± 660 years fro

typical timespan to a common ancestor of R-L23 subclade, and

ped for

subclades when tested several years ago as the typing was not

2/.09 = 164 → 196 generations, or 4900 ± 530 years

from u-

firy

ce with 11% and

others are in singular percentages (Lashgary et al., 2011). Such

collected in the FTDNA P

= 201 → 250 conditional generations, or

m the common ancestor. This again is a

al., 2008].

Among Jewish R1b haplotypes (most were not ty

is in an agreement with the “age” of its upstream R-M269 sub-

clade of ~7000 ybp.

The same base haplotype, as shown in the preceding para-

graph, was found in a dataset of 238 Armenian six-marker R1b

haplotypes published earlier (Weale et al., 2001) and analyzed

in (Klyosov, 2008a). It included haplotypes from six regions of

Armenia, Karabakh, Iran, and other areas of the Armenian di-

aspora. It can be presented as 12 24 14 11 X X X 12 X X 13 X.

An average “age” of the common ancestor of R1b haplotypes in

all the six regions was 5750 ± 1500 years, which is in line with

other estimates for the R-L23 subclade.

Considering other populations, containing a rather high share

of R-L23 haplotypes, which in turn could be a tracer of ancient

migration of the Arbins, one can mention Russia with their 37%

of available R1b haplotypes having DYS393 = 12 (Roewer et

al., 2008; Klyosov, 2010c and references therein) and the base

haplotype 12 24 14 11 11 14 X X 12 13 13 29/30 - 16 15 19

(the last three alleles are of DYS 458, 437, 448), the same as

R-L23, shown above, and with a common ancestor of the Rus-

sian R1b of 6775 ± 830 ybp (Klyosov, 2009d); Anatolia and

Lebanon with their 12 24 14 11 X X X 12 13 13 29 and 12 24

14 10 X X X 12 13 13 29 (mutated DYS391 1110 is marked)

and with the respective common ancestors of 6000 ± 820 ybp

(Anatolia) and 5200 ± 670 ybp (Lebanon) [Klyosov, 2010c;

haplotypes were listed in Cinnioglu et al., 2004, and Zalloua et

available), one branch remarkably resembled R-L23 haplotypes,

and its 37 marker base haplotypes was as follows (Klyosov,

2008b):

12 24 14 10 11 14 12 12 12 13 14 29 - 17 9 10 11 11 25

15 19 29 15 15 16 17 - 11 11 19 23 16 16 18 17 37 38 12

12 (L23, Jewish)

In this particular instance the dataset contained 42 of 37

marker haplotypes, which collectively had 620 mutations; this

gives 620/4

a common ancestor. The base haplotype differs by 9 m

tions from the “young” Jewish base haplotype above in its

st 37 markers, which makes the 4900 ybp haplotype likel

cestral. The oldest identified Jewish R1b base haplotype, is

5400 ± 500 ybp (Klyosov, 2008b). It has certainly appeared

in the Middle East, during the Sumerian era.

Sumers, the likely Bearers of R1b1a2 Haplotypes,

and Their Descendant Assyrians

Assyrians are one of the oldest surviving groups descended,

as it is believed, from the historic Sumers. Among Assyrians,

R1b is the major haplogroup, reaching 40% of the studied

population; haplogroup J takes second pla

A. A. KLYOSOV

Figure 9.

The 17-marker tree composed with 120 haplotypes of the R-L23 haplogroup, except

five haplotypes of the M269 subclade (numbers 1 - 5). The tree is composed based on

data in (Herrera et al., 2011).

a high percentage of R1b

Mesopotamia. According t

Appendix) half of their R1 subclade

with DYS393 = 12, and all nine 12 marker haplotypes have 35

een 6000

from the Pontic

dle East Westward

In the descriptions above we left bearers of R1b haplogroup

ndants

serve

lished in (Barac et al., 2ericic et al., 2005, and

haplogroup is very unusual in South

o the Assyrian FTDNA Project (see

b bearers belong to R-L23

Migrations of the Arbins

Steppes, Asia Minor and Mid

to Europe

mutations, which gives 35/9/.02 = 194 → 240 conditional gen-

erations, or 6000 ± 1200 years to their common ancestor. Only

6 haplotypes were available in the 25 marker format, and they

have 48 mutations; they yield 48/6/.046 = 174 → 211 genera-

tions, or 5275 ± 930 years. Those are indeed Sumerian times

(e.g., Kramer, 1971). Most of the bearers of those haplotypes

now live in Iraq and Turkey. Another half of the Assyrian hap-

lotypes in the Project, mostly from Iran, have a slightly mutated

“classical” European R1b1a2 base haplotype (with DYS464 =

15 15 17 17), and a common ancestor of 850 ± 360 ybp calcu-

lated from the first 12 markers, and 1100 ± 280 ybp from the

first 37 markers. This R1b evidence is clearly a relatively recent

event in Assyrian population, brought from Europe.

The R-L23 subclade is clearly traced from the Russian Plain

south via the Caucasus, where it prevails among R1b haplo-

types, and via Anatolia, where it is very pronounced; down to

South Mesopotamia, where Sumers had lived betw

d 4500 ybp (Kramer, 1971). Since timespans to common

ancestors of those R-L23 haplotypes are around 6000 - 5000

ybp, it is quite likely that those common ancestors lived among

the Sumers and their ancestors (in the Caucasus and Anatolia).

It might be an additional feature for linguists, some of whom

consider Sumerian as a remnant of a subgroup of the Dene-

Caucasian language superfamily (e.g., Bengtson, 1997).

in the Caucasus/Russian Plain/Pontic steppes, Anatolia, and

Lebanon/Southern Mesopotamia around 6000 - 5000 ybp. All

three areas were the relay regions for the Arbins to move fur-

ther westward. Haplotypes of their present day desce

as the tracers of those ancient migrations.

A dataset of R1b1a2 haplotypes on the Balkans was pub-

003a, 2003b; P

ivate communications with M. Pericic). It contains a series of

obviously R-L23 haplotypes with a base 12 24 14 11 11 15 X X

X 13 13 29, and a common ancestor of 4050 ± 890 ybp

(Klyosov, 2010c). Another series of the Balkan R1b1a2 has the

base 13 24 14 11 11 11 X X X 14 13 29, with a common an-

cestor of 4975 ± 1300 ybp (ibid). Four mutations between them

(including multi-copy mutations counted differently) place their

common ancestor at ~7000 ybp to the time of ancient R-M269

haplotypes of the Russian Plain’s (or further to the east). Mi-

grations westward from the Pontic steppes are supported also

by multiple archaeological evidences, which include those dat-

ing to the 3rd millennium BC (e.g., Mallory, 1989).

The haplotype tree shown in Figure 8 provides some clues

regarding possible directions of those ancient migrations. 40%

of all 107 haplotypes of the tree belong to the Armenians and

Turks; of those, 70% are haplotypes on the right-hand (larger)

part of the tree. The upper left branch in the tree contains hap-

lotypes from Russia, Lithuania, Poland, Croatia and Ireland.

A. A. KLYOSOV

They all have the same pattern of mutations, with the base hap-

lotype

12 24 14 11 11 14 12 12 12 12 13 29 - 16 9 10 11 11 25

15 19 31 14 15 16 18 (L23, mainly Eastern Europe)

which differs from the base haplotype of the left-hand side

38-haplotype branch by three mutations (marked; some alleles

are fractional). This places their common ancestor with the

entire branch at 4600 ybp. This indicates that the above largely

East-European branch split from the sizeable branch at the Rus-

sian Plain in the middle of the 3rd millennium BC and appar-

enf

se -

peared

→ 122 generations, or by 3050 “lateral” years

(the

25w

a fork,

only one mutation 12 → 13 in the first allele

ese.

In fact, the entire right-hand large branch in Figure 8 consists

as R-P312, R-L21, R-U152, R-L2, R-L20, etc. The

semon ancestor of 3550 ±

79a common ancestor who

livestor of all of them lived

5025 ± 630 ybp, which fits the time and direction of ancient

ed by the linear method, and

tly migrated westward. That is one migration route (out o

veral described) which explains how R-L23 haplotypes ap

in Europe during those times.

Another migration route took place from the Russian Plain

southward, as indicated by another quite distinct branch on the

opposite side (at 5 o’clock) of the haplotype tree. The branch

includes haplotypes from Russia, Lithuania, Armenia (two hap-

lotypes), Turkey (three haplotypes), Syria. The base haplotype

is as follows:

12 24 14 11 11 14 12 12 12 13 13 30 - 16 9 10 11 11 25

15 19 28 15 15 16 17 (L23, mainly South)

There are five full mutations between these two base haplo-

types of the branches, both which includes Russian and Lithua-

nian haplotypes, albeit, with a principally different history. One

group of L23 bearers went west to Europe; another went south

to the Caucasus and the Middle East. They are separated by

5/.046 = 109

is time is required on average to make five mutations in th

marker haplotype). In other words, these two branches sho

in migration routes of R-L23 from the Russian Plain

estward and southward.

One additional branch of R-L23 includes haplotypes from

Armenia, Lebanon, Bulgaria, Italy, France, Spain, Germany. It

is located at 7 o’clock on the haplotype tree in Figure 8. Their

base haplotype

12 24 14 11 11 14 12 12 12 13 13 29 - 16 9 10 11 11 25

15 19 29 15 16 16 18 (L23, South and South-West)

differs from the preceding discussed branch by only 2.6 muta-

tions or by 1550 ”lateral” years. This points to R-L23 move-

ment from the Middle East westward to Europe. Its neighboring

branch, including haplotypes from Turkey (three haplotypes),

Greece and Albania evidences nearly the same base haplotype

as the above, with

YS393) and 18 → 17 (17.3 on average) in the last allele. It is

sentially the same migration route from Asia Minor to Europ

of sub-branches with predominantly Armenian and Turkish

(some Iraqi and Iranian) R-L23 haplotypes with singular inclu-

sions of haplotypes from Greece, Germany, Netherlands, Eng-

land, Scotland. This reflects migration routes of the Arbins

from Asia Minor and the Middle East to Europe mainly be-

tween 4500 and 3500 ybp.

It may be expected that some migrations from the Middle

East to Europe are associated with this mutation of DYS393 12

→ 13 in the subclade R-L23. However, other, later migrations,

could have occurred from the European continent eastward, and

belong to more recent subclades, such as L51, L11, P312, U106,

L21, U152, etc. (Klyosov, 2011b). Indeed, most Sardinian hap-

lotypes have DYS393=13, such as the following base haplo-

types on Sardinia (calculated from data provided in [Contu et

al., 2008]):

13 24 14 11 11 14 X X X 13 X 29

13 24 14 10 11 14 X X X 13 X 29

13 24 14 10 11 15 X X X 13 X 29

The first one was presented on the tree by a series of identi-

cal haplotypes which are obviously derived from the very re-

cent common ancestor (Klyosov, 2008a, 2010c). They are in-

distinguishable from common European R1b1a2- M269 sub-

clades, such

cond one has a timespan to its com

0 ybp, the third descended from

ed 2900 ± 620 ybp. A common anc

igration of the Arbins from the Middle East to Europe (how-

ever, DYS393 = 13 would be unusual for them), and from

Iberia up north to the continent, and subsequently populating

Europe in all possible directions.

One of the most common R1b base haplotypes in Sicily is as

follows (Di Gaetano et al., 2009): 13 24 14 11 11 14 X X 12 13

13 29, which is identical to one of the Sardinian base R1b hap-

lotypes above, and indistinguishable from common R1b sub-

clades in Europe. The timespan to a common ancestor for that

haplotype in Sicily calculated from the cited study is 4550 ±

1020 years. The same base haplotype is the most common in

Italy (Capelli et al., 2007), with a timespan to the common an-

cestor of 4125 ± 500 years calculat

00 ± 1160 years calculated by the logarithmic method (Kl-

yosov, 2008a).

There is one more, but very important migration route of the

Arbins which does not practically include R-L23 haplotypes

(only 5% of DYS393 = 12 among Iberian R1b haplotypes, it

corresponds to a random mutation from the parent DYS393 =

13). It is a route from the Middle East westward along North

African Mediterranean seacoast. R-L23 was either not repre-

sented or did not survive along this route. It seems that bearers

of R-V88 were part of the journey, however, split and went

southward and settled in Central Africa (mainly Cameroon and

Chad), mentioned previously. The migration of the Arbins from

the Middle East to the Atlantic, then across the Strait of Gibral-

tar to the Pyrenees took place from ~ 5500 - 5200 ybp to 4800

ybp when the Arbins landed in Iberia (see supportive references

to archaeological data below). Part of the way from Egypt to

Iberia could have been made by sea, details are not known as

yet. There are some historical reports of arrival of the Egyptian

military fleet to Iberia some 5000 ybp; there are some allegedly

Egyptian mummies and fragments of Egyptian tombs exhibited

in the Royal Academy of History in Madrid and in the Tar-

ragona City Museum. Their status, however, is rather vague. It

seems, nevertheless, that bearers of R1b subclades, mainly

R-M269, and newly formed L51 and L11 (see below) had ar-

rived in Iberia and this was the beginning of the archaeological

Bell Beaker culture.

Previous to discussing the Bell Beakers and history of the

Arbins in Europe, it is worth mention of one more rather vague

evidence of the R1b journey via ancient Egypt between 5500

and 5200 ybp. It concerns the alleged R1b haplotype of Phar-

aoh Tutankhamun.

A. A. KLYOSOV

Alleged Pharaoh Tutankhamun R1b1a1-M269*

Haplotype and Its Possible History

Recently the Swiss company iGENEA has published the al-

leged 16 marker haplotype of the Pharaoh: 13 24 14 11 11 14 X

X 10 13 13 30 - 16 14 19 10 15 12

(www.igenea.com/en/ index.php?c=62).

Here the first 12 markers are shown in the FTDNA format,

and the rest are DYS 458, 437, 448, GATA H4, DYS 456, 438.

It is obviously not a typical European R1b1a2 haplotype, since

it h

The mostype is

17 9 10 11 11 25

tor of the two lived (6850 + 6200 + 3300)/2 =

81 -

ce e

m Altay, South Siberia,

whe

it is-

doun

R1b haplotypes ybp.

Sea to Iberia, up north to the European continent. The

first two principal routes were associated with bearers of the

wh -

media very

clos

as DYS439 = 10, and not a common European 12. There are

nly about 0.5% R1b haplotypes in Europe with DYS439 = 10.

likely and the most closely related base haplot

at of R1b1a2-M269, shown in Figure 5, in which the same

markers as those available in the Pharaoh haplotype are noted

in bold:

12 25 14 11 11 14 11 12 12 13 13 29 -

15 19 29 15 15 16 17 - 10 11 19 23 15 15 19 17 35 38 12

12 - 11 9 15 16 8 10 10 8 11 10 12 23 23 16 10 12 12 15 8

13 22 20 13 12 11 13 11 11 12 12 (M269)

There are 6.8 mutations between the two 16 marker haplo-

types (some alleles in M269 haplotype are fractional), which

translates to 6.8/.0315 = 216 → 274 conditional generations, or

6850 years between them. Since Tutankhamun lived 3300 ybp,

and the R-M269 base haplotype is 6200 ± 900 years “old”, then

a common ances

75 ybp. This might have been either an ancient R-M269 an

stor of the Tutankhamun from Central Asia, or the respectiv

stream subclade.

The main point here is that the Pharaoh haplotype is not

me erroneously (or intentionally) picked European R1b1a2

haplotype; it is an archaic haplotype of R1b haplogroup, likely

of the R-M269 ancient subclade.

It should be mentioned here that the founder of the Egyptian

pharaohs was Narmer, the origin of whom is not firmly known,

and was the founder of the First Dynasty, living ca. 32nd cen-

tury BC—around 5200 ybp. This date fits the migration times

for the Arbins along the Mediterranean coast in North Africa. It

cannot be excluded that the bearers of R1b haplogroup may

have actually established the Royal lineage in ancient Egypt. It

does not mean that the lineage was not interrupted through the

Dynasties; however, it kept returning into the Dynasties, if the

R1b origin of some Pharaohs is right.

The Overview of the Migratory Path of the

Arbins, Bearers of R1b Haplogroup, from

Central Asia to Europe

An overall map of R1b migration routes from the very “be-

ginning”, where “beginning” is undefined in detail thus far and

can be estimated between 16,000 and 12,000 ybp, and until

their arrival to the Pyrenees by 4800 ybp as future Bell Beakers,

is shown in Figure 10. Their migratory path was slowly taking

place from Central Asia, apparently fro

n some very different R1b haplotypes were discovered. As

described in the first sections of this paper, their tremen

s mutational difference with European and Middle Easter

places their common ancestors at ~16,000

The legend to Figure 10 describes those Central Asian/Si-

berian populations. There are many Neolithic, Chalcolithic and

Eneolithic archaeological cultures in the area, such as Tersek,

Ural, Surtandi, Mahandzhar, Iman-Burluk, Botai, Atbasar, Kel-

teminar, and other archaeological Central Asian cultures in

present-day Russia (e.g, Zakharov, 2010), which might be as-

signed to the Arbins; however, it would be premature to assign

any of them to R1b or to any other haplogroup. Such a task is

quite new for archaeologists. It is tempting to point at Serogla-

zovo, Khvalyn, Samaran, Middle-Volga and adjacent archaeo-

logical cultures of 12,000 - 5000 ybp of the European Russian

east as the most likely R1b cultures. We cannot, however for

the same reason of prematurity and it would be irresponsible to

suggest such at this time. The same may be said for Yamnaya,

Catacomb and neighboring archaeological cultures of Central

and South Russia, which apparently were shared by both R1b

and R1a bearers, albeit in different time periods. R1b before

5000 ybp, R1a after 4500 ybp have confused archaeologists

who have observed “different roots” of those cultures, spread-

ing in different directions and in different times.

The map shows that current bearers of R1b spread over Cen-

tral Russia up to Arkhangelsk on the White Sea. Very likely it

was a relatively recent relocation although it remains to be de-

termined. Currently there are only about 5% of R1b bearers in

the European part of Russia.

As it was described above in this study, the Arbins went

South through the Caucasus to Mesopotamia and the Middle

East around 6000 - 5000 ybp; established the Sumer civilization;

went westward via Egypt to the Atlantic, and across the Gibral-

tar Strait to the Pyrenees. On their way some R1b-V88 bearers

split; went deep into Africa, and currently populate Cameroon

and Chad in appreciable amounts (see legend to Figure 10 and

description and references above). By 4800 ybp the Arbins

have reached Iberia to become the first Bell Beakers. This date

was obtained for a common ancestor of haplotypes of P312 and

U106 subclades (see the next Section), and is supported by

archaeology data (Cardoso & Soares, 1990; Martinez et al.,

1996; Cardoso, 2001; Muller & Willigen, 2001; Nocete,

2006).

Several Entry Routes of R1b1a2 Haplotypes to

Europe: From the Russian Plain, from Asia

Minor/Middle East, from the Pyrenees

To sum up the preceding section, the Arbins were entering

Europe from the east by several routes: from the Russian Plain

(between the Pontic Steppes and the Baltic Sea); from Asia

Minor and the Middle East; and after a long way around Medi-

terranean

-L23 subclade. The Iberian route was made by mainly M269

ich at the time of entering the Pyrenees split R-L51 and im

tely after R-L11. Both are very similar and have

e timespans to their common ancestors, as it is shown in the

next section. In 4850 ybp L11 promptly split off two “brother”

subclades, P312 and U106 (Klyosov, 2011b) which after a long

“population bottleneck” on the edge of extinction, eventually

survived and expanded around 4000 - 3700 ybp, and actively

populated Europe, first as Bell Beakers, between 4000 and

3000 ybp, and then up to the era of Ancient Rome, Gauls and

Celts, mentioning only those names which present certain

“milestones” in history. In fact, there were dozens if not hun-

dreds of ancient R1b tribes in Europe.

A. A. KLYOSOV

Figure 10.

The map of R1b haplotype locations considered in this study, until they arrived to Iberia ~4800 ybp. The overall migration path of the Arbins had

apparently started in South Siberia, where present day R1b haplotypes of the Shors, Teleuts, Altayans, Uighurs, Khakass, Tuva people (the right-hand

side pins) are tremendously different from those in Europe and the Middle East. The migration route continued through the regions in Middle Asia

and Russia, where Bashkirs, Kazakhs, Uzbeks, Mari, Tatars, Kalmyks presently live. From the Central Russia region the Arbins moved to the Cauca-

sus and further South to Anatolia, Middle East, present day Iraq, Lebanon, and then westward along the Mediterranean Sea onto Northern Africa t

the Atlantic. On that way R1b-V88 bearers split and went deep into Africa, where their descendants currently live in Cameroon and Chad (Cruciani e

eir migratory path into Africa is reproduced here from the above reference. The map shows R1b locations previous to arrival to the

strea he

ea.

Cu-

rope, ed a

al., 2010). Th

Pyrenees ~4800 ybp, from where they moved North as Bell Beakers and populated Europe between 4800 - 3000 ybp and later.

Main R1b1a2 Subclades on the European

Continent: Entering Europe 4800 - 4500 ybp

from the East and from the South West

Nearly all haplotypes of the subclade R-M269 6000 - 5000

ybp belonged either to R-M269* para-group, or to its down-

R1b1a2a1a1b4 L21/M529

R1b1a2a1a1b4b M222

R1b1a2a1a1b4g1 L226

R1b1a2a1a1b4h1 P314

The above diagram shows that the immediate downstream

subclades of RSOGG-2012, in

an abbreviateddes is much

more understood ra-

tion of the Arbins is

The question is—wdes could

have arisen? If they are d have

split in Asia Minor, thePlain,

and enter Europe from ther, that

is only L51 or only L11 tion

bottlenecks, hence, look “you be

(in terms of mutations and th,

m subclade R-L23. The Arbins who had migrated from t

st to Europe during those times mostly carried DYS393 = 12

rrently we see only those R1b1a2 (xL23) haplotypes in Eu

which are derived from common ancestors who liv

-L23 were L51 and then L11 (I

form). The dynamics of these subcla

via Iberia into the continent where the mig

beakers. ing identified with the Bell Be

aximum ~4500 ybp. Either descendants of more ancient L23

ancestors have not survived into the present time or there were

none from the eastern Russian Plain, Asia Minor, and the Mid-

dle east.

R1b M343

R1b1 L278, M415, P25_1, P25_2, P25_3

R1b1a P297, L320

R1b1a1 M73, M478

R1b1a2 L265, M269, M520, S3, S10

here those L51 and L11 subcla

6000-5000 years “old”, they coul

Middle East or on the Russian

ese. The “intraclade” haplotyp

subclade, might reflect popula

nger” than they in fact should

e respective TMRCA). However

eir “interclade” comparison could reveal their lost (due to

bottlenecks) timespans to more ancient common ancestors. To

analyze those subclades, a combined L51-L11 haplotype tree is

shown in Figure 11.

One-third of haplotypes on the tree belong to the L51 su

R1b1a2a L23/S141, L49.1

R1b1a2a1a L51/M412/S167

R1b1a2a1a1 L11, P310S129, P311

ade, and they occupy the right side. Another two-thirds are

L11 haplotypes, which are on the left and make some “insert”

branches on the right. The 67 marker haplotypes of L51 and

R1b1a2a1a1a U106/S21

R1b1a2a1a1b P312/S116

R1b1a2a1a1b3 U152/S28

A. A. KLYOSOV

Figure 11.

The 67-marker tree composed with 42 haplotypes of the R-L51 subclade (15 haplotypes on the

right, numbers between 1 and 23) and R-L11 (27 haplotypes on the left, numbers between 26 and

61). Haplotypes are listed in the FTDNA public Project

http://www.familytreedna.com/public/ht35new/default.aspx.

L11 subclades are very similar, therefore the tree could not

distinguish them in a ce, the mix of the

branches. The tree als

clades is also very sim

indicates the “age”) of

same. The base haploty

y of

th r

~2n

28 2

= ir

fits well with the arrival time of the Arbins to Iberia (4800 ybp),

to the distribution pattern of the current bearers of L51 in

yrenees and immedi-

, see [Myres et al.,

. It is practically absent in European South-East, East, and

st (ibid.).

Analysis of the L11 subclade is more complicated since its

nd the tree by at least four branches,

etween them (marked in bold) present in

number of cases; hen

o shows that the “age” of the two sub-

ilar, since the “height” (which generally

the branches around the tree is about the

pe of R-L51 subclade is as follows

Europe (the highest frequency is in the P

ately up north in France and on the Isles

2010]

North-Ea

13 24 14 11 11 14 13 12 12 13 13 29 - 17 9 10 11 11 25

15 19 30 15 16 17 18 - 11 11 19 23 16 15 18 17 36 38 12

12 - 11 9 15 16 8 10 10 8 10 10 12 23 23 16 10 12 12 16 8

12 22 20 14 12 11 13 11 11 12 12 (L51)

It deviates from the base R-L23 haplotype (5400 ± 800 ybp)

by 9 mutations (12 mutations are marked above, but man

em are fractional), that is by 9/.12 = 75 → 81 generations, o

025 years of the “lateral” distance. All 15 haplotypes contai

0 mutations from the base haplotype, which gives 280/15/.1

156 → 184 generations, or 4600 ± 535 years from the

mmon ancestor, L51. In turn, it gives (2025 + 5400 + 4600)/2

6000 years, which is an “age” of a common ancestor of both

L23 and L51 subclades. Obviously, it is the common ancestor

of L23 subclade himself.

In other words, the “age” of the L51 subclade as 4600 ± 535

years is obtained from mutations on the L51 branch on the tree

(Figure 11) and confirmed 1) by a mutational distance from the

base haplotype of the parent L23 subclade, 2) by the “age” of

the L23 subclade, determined earlier, and 3) by the general

phylogeny of the R1b subclade. The “age” of the L51 subclade

haplotypes are spread arou

ch with its common ancestor. They differ from each other by

32 mutations which gives 32/4/.12 = 67 → 72 generations, or

1800 years below their average “age” ~2500 years. Therefore

the “age” of the R-L11 subclade is ~4300 years, which is in-

deed consistent with 4600 ± 535 years for the L51 subclade.

The base haplotype for the L11 subclade is as follows:

13 24 14 11 11 14 12 12 12 13 13 29 - 17 9 10 11 11 25

15 19 30 15 15 17 17 - 11 11 19 23 16 15 18 17 36 38 12

12 - 11 9 15 16 8 10 10 8 10 10 12 23 23 16 10 12 12 15 8

12 22 20 13 12 11 13 11 11 12 12 (L11)

Five deviations b

ct 3.7 mutations (since most of them are fractional), which

gives 3.7/.12 = 31 → 32 generations; that is only 800 “lateral”

years between them. This confirms that L51 and L11 subclades

are very close to each other in time. Their common ancestor

(which presumably should be L51) lived (800 + 4600 + 4300)/2

= 4850 ybp. It seems that L11 split off only ~250 years later.

All of them are likely to have established the archaeological

100

A. A. KLYOSOV

Bell Beaker culture. The oldest artifacts of the Bell Beakers

were found in

13 29 - 17 9 10 11 11 25

There are 5.5 (maations between them

d 4175 ± 430 ybp

(K+

41 -

clt

ths,

dee

is -

an with seafaring carriers of L11* bearers from

d of R1b (with sub-

se haplotype differs from the

“b

span to its com

da s

exs

sh -

nie

me time. It only confirms that their

Portugal, dated 4800 - 4600 ybp (Cardoso &

ares, 1990; Martinez et al., 1996; Cardoso, 2001; Muller &

illigen, 2001; Nocete, 2006).

Main R1b1a2 Subclades on the European

ontinent: Population of Europe after 4800 ybp,

Main Subclades on the Isles and Their

Likely Origin

The phylogeny of R1b (see the diagram above) shows that

the further downstream subclades of R-L11 are P312 and U106.

Their 67 marker haplotypes are as follows:

13 24 14 11 11 14 12 12 12 13 13 29 - 17 9 10 11 11 25

15 19 29 15 15 17 17 - 11 11 19 23 15 15 18 17 36 38 12

12 - 11 9 15 16 8 10 10 8 10 10 12 23 23 16 10 12 12 15 8

12 22 20 13 12 11 13 11 11 12 12 (P312)

13 23 14 11 11 14 12 12 12 13

5 19 29 15 15 17 17 - 11 11 19 23 16 15 17 17 37 39 12

12 - 11 9 15 16 8 10 10 8 10 10 12 23 23 16 10 12 12 15 8

12 22 20 13 12 11 13 11 11 13 12 (U106)

rked in bold) mut

YS456 in P312 fluctuates between 15 and 16 in various

datasets), which separates the two base haplotypes by 5.5/.12 =

46 → 48 generations, or by 1200 years. Since common ances-

tors of both subclades lived 4350 ± 700 an

lyosov, 2011b), their common ancestor lived (1200 + 4350

75)/2 = 4850 ± 700 ybp. Indeed, it is the “age” of the sub

ade L11 within margin of error. Generally, it is safe to accep

e “age” of both P312 and U106 as ~4200 ybp, since estimate

pending on a dataset, are fluctuating around this value. Ther

1200 years between them (see above) which defines a time

an to their common ancestor, L11, as (4200 + 4200 + 1200)/2

4800 ybp.

As it was stated above, the four subclades, L51, L11, P31

d U106 historically represent the first wave of Bell Beaker

movement from Iberia to the European continent. The highest

frequency of P312* subclade in Europe is currently observed in

Iberia (Myres et al, 2010), however, it is naive to judge ancient

locations of the subclades based on their current distribution.

The mixing of them on the Continent has been too intense in

the past millennia to make any unambiguous extrapolations.

The current regional distribution of the R-M269’s downstream

subclades which moved from the Pyrenees nearly five millennia

ago is classically non-informative, except in instances of some

local subclades. One of them is R-M222.

It seems that the history of R-M222 appearance on the Isles

may have beg

eria to the Isles some 4500 - 4000 ybp. However, it is just one

theory. Other involve migrations of L21 bearers to the Isles by

land. Then, P312, a downstream of L11, is also intensely rep-

resented on the Isles along with its downstream subclades, as

well as its “brother” subclade U106 and its downstreams (ibid).

L21, a downstream of P312, is abundant on the Isles, unlike its

“brother” subclade U152 (ibid). This ancient migration of the

Arbins from Iberia to the Isles explains some genetics data on

the “Spanish origin of the Irish”.

M222, a downstream of L21, is among the most represented

R1b subclades on the Isles. A massive Ireland Heritage FTDNA

project (see Appendix) lists several thousan

ades) haplotypes in the Isles, of which M222 is the largest,

and includes about 25% of all R1b (Klyosov, 2010e). The sub-

clade is expressed the most in Ireland. In Scotland it is observed

mainly in the lowlands and in central region, and poorly pre-

sented in England. The base haplotype of the M222 subclade is:

13 25 14 11 11 13 12 12 12 13 14 29 - 17 9 10 11 11 25

15 18 30 15 16 16 17 - 11 11 19 23 17 16 18 17 38 39 12

12 - 11 9 15 16 8 10 10 8 10 10 12 21 23 16 10 12 12 16 8

12 25 20 13 12 11 13 11 11 12 12 (M222)

It is of interest that the above ba

rother” base haplotypes of P312 and U106 by 18 mutations

in each case (in fact, 15 mutations, since some mutations are

fractional). It shows that P312 and U106 are indeed very close

(in time and in heritage) base haplotypes, and that their com-

mon ancestor, a bearer of L11 base haplotype, lived very close

in time to both of them. 15 mutations place their common an-

cestor at 4600 ± 500 ybp.

Multiple analyses of M222 haplotype datasets gave a time-

mon ancestor of currently living M222 descen-

nts of 1450 ± 160 years (Klyosov, 2010e). However, cros

amination of a number of lineages in the M222 subclade ha

own that the M222 lineage arose not later than in the begin

ng of the Common Era, some 400 years earlier than the abov

timate. It points to a possible population bottleneck between

the time when M222 arose, and the expansion of the subclade

in the middle of the 1st millennium CE. One cannot possibly

say, at least yet, where specifically the M222 subclade first

appeared (that is, the G → A single nucleotide mutation oc-

curred in certain nucleotide of Y chromosome of a certain indi-

vidual, the bearer of the upstream L21 subclade), whether in the

Isles, in North Western Europe, or elsewhere. We also cannot

determine specifically against what R-M222 direct descendants

have been struggling for survival for several centuries. Was it a

long climatic event? The Roman invasions? Epidemics? Fam-

ine? or a combination? We know, however, that it was the Isles

where this mutation on the Y chromosome was eventually car-

ried on and flourished.

The phylogenic diagram shows that the M222 mutation arose

concurrently with its “brother” SNP mutations which now de-

fine subclades L226 and P314. The first one has a common

ancestor who lived 1500 ± 170 ybp, which is at the same time

with that of M222, albeit in L226 the subclade-defined muta-

tion was C  T. Its base haplotype is as follows:

13 24 14 11 11 14 12 12 11 13 13 29 - 17 8 9 11 11 25 15

19 29 13 13 15 17 - 11 11 19 23 15 15 18 17 36 38 12 12

- 11 9 15 16 8 10 10 8 10 10 12 23 23 15 10 12 12 15 8 12

22 20 13 12 11 13 11 11 12 12 (L226)

It differs by as many as 21 mutations from its “brother”

M222 subclade, despite the observation that their common

ancestors lived at the sa

mmon ancestor, the founder of subclade L21, lived around

4100 ybp. It also shows that L21 formed almost immediately

from P312 (see the phylogenic diagram) since P312 arose 4350

- 4200 ybp and L21 around 4100 ybp. Their 67 marker haplo-

types are identical, within some fractional mutational differ-

ence:

13 24 14 11 11 14 12 12 12 13 13 29 - 17 9 10 11 11 25

15 19 29 15 15 17 17 - 11 11 19 23 15 15 18 17 36 38 12

12 - 11 9 15 16 8 10 10 8 10 10 12 23 23 16 10 12 12 15

A. A. KLYOSOV

8 12 22 20 13 12 11 13 11 11 12 12 (P312)

13 24 14 11 11 14 12 12 12 13 13 29 - 17 9 10 11 11 25

15 19 29 15 15 17 17 - 11 11 19 23 16 15 18 17 36 38 12

12 - 11 9 15 16 8 10 10 8 10 10 12 23 23 16 10 12 12 15

8 12 22 20 13 12 11 13 11 11 12 12 (L21)

The only apparent difference, in DYS456, is also fractional,

since this allele fluctuates in P312 between the values of 15 and

16.

Despite their similar chronology, subclades M222 and L226

have quite different histories for the preceding 2600 years. Yet

an ,

sin n

hap;

ho 2

mutations, respectively. It is a third, completely differ-

a rather robust system of

this paper,

are as follows (iplotype per the

conditional genees show exam-

ht x?id=9

using software PHYLIP,

Phm (see Klyosov, 2009c,

20ons for back mutations

wc). Margins of

erlyosov, 2009c).

rmined by minimi-

za plotype is one

wr of mutations in the

dacestor. The base haplo-

ty losest approximation to

two base haplotypes

is determined mutations

ed num-

other “brother” subclade, P314 (more accurately, P314.2

ce the same P314 SNP mutation, T → C, has occurred i

plogroup H2a), is somewhat “older”, with its 2225 ± 300 yb

wever, it could be within the same time frame when M22

tually arose. Its base haplotype

13 23 14 11 11 14 12 13 13 13 13 29 - 17 9 10 11 11 26

15 19 29 15 15 16 17 - 10 11 19 23 16 15 18 18 37 38 12

12 - 11 9 16 16 8 10 10 8 11 10 12 23 23 17 10 12 12 15

8 12 22 20 13 12 11 13 11 11 12 12 (P314.2)

is again remote from those of M222 and L226 subclades, by 24

and 17

t lineage in the L21 family. The 10 full mutations from its

L21 parent base haplotype gives 10/.12 = 83 → 91 generations,

or 2275 years of the mutational difference, and places the be-

ginning of the L21 subclade at (2275 + 4100 + 2225)/2 = 4300

years, which is within margin of error from 4100 years esti-

mated above. All these base haplotypes along with the intra-

clade and interclade datings form

NA genealogy of the Arbins in Europe and Asia.

Materials and Methods

Four thousand four hundred eight (4408) of R1b haplotypes

(with subclades) were collected in databases from FTDNA,

YSearch, and in peer review publications.

The methodology of haplotype datasets analysis was de-

scribed in the preceding publication in this journal (Rozhanskii

& Klyosov, 2011). In this study the linear and the logarithmic

method were employed; the latter when the base haplotype in

the dataset was easily identifiable, as described in (Klyosov,

2009c). The mutation rate constants are listed in (Klyosov,

2009c; Rozhanskii & Klyosov, 2011), and for a number of

cases are given in the text of this paper. The most widely used

mutation rate constants, including those employed in

n a number of mutations per ha

ration of 25 years); the referenc

es of the haplotypes in the format indicated:

.013, the 8 marker haplotype (Zhong et al.)

.017, the 8 marker (Contu et al.)

.018, the 10 marker (Myres et al.)

.020, the 11 marker (Cruciani et al.)

.019, the 11 marker (Zalloua et al.)

.020, the 12 marker FTDNA format

.024, the 12 marker (Malyarchuk et al.)

.034, the 17 marker Y-filer format (Herrera)

.035, the 19 marker (Balanovsky et al.)

.046, the 25 marker FTDNA format

.090, the 37 marker FTDNA format

.120, the 67 marker FTDNA format

.198, the 111 marker FTDNA format

FTDNA haplotype formats are given in

tp://www.familytreedna.com/faq/answers.asp

Haplotype trees were composed

ylogeny Inference Package progra

09d and references therein). Correcti

ere introduced as described in (Klyosov, 2009

ror were calculated as described in (K

Base haplotypes in the dataset were dete

tion of mutations; by definition, the base ha

hich has the minimum collective numbe

taset, derived from one common an

pe is the ancestral haplotype or the c

e latter.

A timespan to the common ancestor of

as follows: 1) count the number of

between the two base haplotypes, 2) divide the obtain

ber by the mutation rate constant, 3) introduce a correction for

back mutations, calculated using the following formula (Klyo-

sov, 2009c):



obs

λ1expλ

 (1)

where:

obs

λ = observed average number of mutations per marker in

a dataset (or in a branch, if the dataset contains several branches/

lineages).

λ = average (actual) number of mutations per marker cor-

rected for back mutations, 4) add the obtained value, multiplied

by 25 (years), which represent the “lateral” timespan between

times of appearance of the two base haplotypes, to TMRCAs

for the both base haplotypes and divide by 2. The result repre-

sents the TMRCA (time for the most recent common ancestor)

for the two base haplotypes under study. The following exam-

ples are based on specific cases in this study.

Example 1: Calculation of a timespan to

of the branch (the most remote on the hap

) which contains 12 of eight marker haplotypes collec-

a common ancestor

lotype tree shown in

Figure 1

tively having 65 mutations from their base haplotype. Since the

mutation rate constant for these haplotypes equal to 0.013 per

haplotype per a conditional generation (25 years), we have

65/12 .013 =/ 417 → 619 generations, that is 619 × 25 = 15,475

years to a common ancestor of the branch. The arrow shows a

correction for back mutations. This correction can be calculated

using a formula (1) as follows. Since the observed number of

mutations per marker is 65/12/8 = 0.677, we employ formula (1)

and obtain



obs

11exp 0.677λ

2

The obtained number of 1.484 is the coefficient of the cor-

rection for back mutations. Therefore, by multiplying 417 ×

1.484, we obtain that the corrected number of generations is

619, that is 619 × 25 = 15,475 years. This is usually designates

as 417 → 618 (generations). Since for 65 mutations in the

dataset the margin of error is 15.93% (calculated as explained

in Klyosov, 2009c), we at last obtain the timespan to a common

ancestor of the haplotypes to be equal to 15,475 ± 2500 years.

Example 2: Two 67 marker base haplotypes of subclades

R-P312 and R-U106 differ by 5.5 mutations (see the text

above). Applying the same rule explained in the preceding ex-

102

A. A. KLYOSOV

ample, we get 5.5/.12 = 46 → 48 generations, that is 1200 years

from a common ancestor of the two base haplotypes (that is, of

the two subclades). Should the two base aplotypes have the

same “age” (the same TMRCA), their common ancestor lived

ybp

(the timeframe between of R1a and R1a1 is

uncertain) and R1b ~16,

roto-Kugran” and/or

“K

lture and the state and migrated westward to Europe

guages were

. M., Janicijevic, B., Parik, J., Rootsi, S.

et al. (2003a). Y chr Forensic Science

International, 138,

Barac, L

uman Genetics, 11, 535-

542. doi:10.1038/sj.ejhg.5200992

0 years “deeper” in time from either one of them. However,

in this particular case the two TMRCA are equal to 4350 and

4175 years, respectively. Therefore their common ancestor

lived (1200 + 4350 + 4175)/2 = 4850 years ago.

Conclusion

The results of this study lend a support to the theory that

haplogroup R1b arose in Central Asia, apparently in South

Siberia or the neighboring regions, around 16,000 years before

present. The preceding history of the haplogroup is directly

related to the appearance of Europeoids (Caucasoids) ~58,000

ybp, likely in the vast triangle that stretched from Western

Europe through the Russian Plain to the east and to Levant to

the south, as it was suggested in (Klyosov, 2011d). A subse-

quent sequence of SNP mutations in Y chromosome, with the

appearance of haplogroups NOP ~48,000 ybp and P ~38,000

ybp in the course of their migration eastward to South Siberia,

eventually gave rise to haplogroup R ~30,000 ybp and R1

~26,000 ybp, and then to haplogroup R1a/R1a1 ~20,000

the appearance

000 ybp (ibid.).

Based on the respective syllables, we call bearers of R1a the

Aryans, and those of R1b the Arbins. In the first case that name

is justified since the bearers of the R1a haplogroup became the

legendary Aryans who arrived in the Hindustan and Iranian

Plateau ~3500 ybp. In other words, those Aryans belonged to

R1a haplogroup, hence, the double meaning (albeit coinciding)

of the term the Aryans. The Arbins is a convenient common

term in avoidance of the repetition: “The bearers of haplogroup

R1b”.

At some point in time, the Arbins began migration to the

west, across Central Asia, North Kazakhstan, South Urals, to

the Russian Plain where they have established a number of

archaeological cultures between 12,000 and 4500 ybp (include-

ing apparently Seroglazovo, Khvalyn, Samaran, Middle Volga,

Drevneyamnay, Catacomb, and also “P

urgan” cultures which are largely considered as controversial

and not accepted by many historians; it should be emphasized

that all those above suggestions regarding the archaeological

cultures can be viewed at present only as very tentative ones).

They migrated southward, in part (leaving their R1b haplo-

group and the respective haplotypes behind), over the Caucasus

to Anatolia around 6000 ybp; to the rest of Asia Minor, and to

the Middle East. The Arbins have apparently established the

Sumer cu

several routes, carrying mainly the R-M269 subclade and its

downstream L23 subclade. One route is the northern route,

from the Russian Plain to the west, ~4600 - 4400 ybp; another,

concurrently along Asia Minor and the Middle East westward

with the same two subclades; and yet another which will popu-

late Europe the most, migrating along North Africa-Mediter-

ranean Sea via ancient Egypt to the Pyrenees, to arrive ~4800

ybp. On this route the R1b-V88 tribe split off and went south,

eventually to Central Africa (mainly Cameroon and Chad judg-

ing by their present-day distribution), where a common ances-

tor of the current R1b-V88 haplotype lived ~4400 ybp.

At the arrival time to Iberia ~4800 ybp, the M269 subclade

split off M51 and soon thereafter L11 and its downstream sub-

clades. They became the Bell Beakers and moved north along

with newly arisen subclades P312 and L21, the latter within a

few centuries after P312. Those subclades and their downstream

clades have effectively without major interruptions populated

Europe from the Atlantic to the Balkans, Carpathian Mountains,

present day Poland, the western border of the Russian Plain, to

the Baltic Sea evidenced by the smooth haplotype trees as wit-

ness to non-stop proliferation of R1b haplotypes.

The Isles had a different history of their R1b haplotypes and

lineages. The bearers of L11, P312 and L21 moved to the Isles

by land and sea concurrently with those Arbins who were

populating Europe between 4000 and 2500 ybp, and formed the

respective “local” subclades, such as P314, M222, L226, which

rgely populated the Isles. As a result, a significant part of the

Isles is populated almost exclusively by the Arbins whose fre-

quency reaches 92% - 96% among the population. In general,

the frequency of the Arbins in Western and central Europe

reaches—albeit not uniformly—some 60% of the population.

This study essentially presents an example of the application

of DNA Genealogy for studying the history of mankind. This

example is a complex and challenging endeavor which also

touches upon some mysterious puzzles of history and linguis-

tics. One of those puzzles is what language or lan

oken by the Arbins from 16,000 to 3000 ybp and it almost

certainly was a continuing, in its dynamics, a non-Indo-Euro-

pean language. Assumptively, these languages are considered

by linguists as assorted and disconnected “dead” and not-so

dead languages, such as proto-Turkic, Sumer, North-Caucasian,

Dene-Caucasian, Basque, and many pre-IndoEuropean lan-

guages in Europe of 5000 - 2000 ybp, some later. The language

of the Arbins may have been originally one language easily

flowing through millennia and across Eurasia. However, this is

a subject of another study.

Acknowledgements

The author is indebted to Susan Hedeen for her valuable help

with the preparation of the manuscript.

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PPENDIX

The following DNA projects were selected as primary hap-

type databases:

http://www.familytreedna.com/public/R1b1b1/default.aspx?s

ction=yresults

http://www.familytreedna.com/public/ht35new/default.aspx

treedna.com/public/AssyrianHeritageDNA

px?section=yresults

http://www.familytreedna.c om/public/IrelandHeritage/defaul

t.aspx?section=yresults

http://www.familytreedna.com/public/R-L21/de fault.aspx? se

ction=yresults

Reference data were selected according to SNP assignment

from YSearch database: (http://www.ysearch.org) and public

projects of FTDNA (http://www.familytreedna.com)

http://www.family

Project/default.as

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