Advances in Anthropology
2012. Vol.2, No.3, 139-156
Published Online August 2012 in SciRes (http://www.SciRP.org/journal/aa) http://dx.doi.org/10.4236/aa.2012.23017
Haplogroup R1a, Its Subclades and Branches in Europe during
the Last 9000 Years
Igor L. Rozhanskii, Anatole A. Klyosov*
The Academy of DNA Genealogy, Newton, USA
Received May 5th, 2012; revised June 11th, 2012; accepted June 25th, 2012
This study identifies and describes 38 branches of the haplogroup R1a STR haplotypes which currently
exist in Europe or which migrated from Europe to areas in the east, south, and southeast between 6000
and 4500 years before the present (ybp). The study is based on 2471 haplotypes which have been tested
for either 67- or 111-markers; it essentially creates a unified robust system, which assembles dozens of
R1a-SNPs and thousands of STRs and assigned haplotypes to branches, some of which do not have SNP
assignments as yet. The assembled system consists of base (deduced ancestral) haplotypes, one for each
STR branch and for each SNP-assigned subclade, each with its characteristic (ancestral) set of alleles, ar-
ranged in the chronological space from ~9000 ybp to 1300 ybp. We found that the most ancient R1a sub-
clades (R1a1-M198− and R1a1a-M198+/M417−) bearers of which currently live in Europe (the present
day haplotypes are scattered between England and the Balkans) appeared in Europe at least 7300 ybp, and
possibly 9000 ybp. R1a’s three principal downstream subclades, L664 (North-Western branch), Z93
(South-Eastern branch), and Z283 (Eurasian branch), split from their common European ancestor at about
the same time, around 7000 - 6000 ybp. L664 apparently stayed in North-Western Europe; its lineage re-
covered and began expanding ~4575 ybp. The Z93 subclade began to expand during the Aryan migra-
tions, on the Aryan’s journey to India and the Middle East in the 3rd-2nd millennia BC. The Z283 sub-
clade split ~5500 ybp into three branches. One of them, Z280 (the Central Eurasian branch) moved east to
the Russian Plain in 4800 - 4600 ybp, and formed at least 16 sub-branches there and in the course of the later
westward repopulation of Europe in the 1st millennium BC—1st millennium CE. Some of the older
branches, like the Russian Plain branch, largely stayed in the present Russia-Ukraine-Belarus-Poland-
Baltic countries region, and were described by early historians as the Scythians, Antes, Veneti, and a mul-
titude of different proto-Slavic tribes (though many of them belonged to haplogroups other than R1a,
primarily I1 and I2). Those R1a branches which are “older” than 3000 years, such as the Russian Plain
branch (4600 ybp), the Western Eurasian (4300 ybp), and the Balto-Carpathian (4300 ybp), did not move
en mass to Europe but stayed behind at the Russian Plain. In the middle of 1st millennium CE, the time of
the collapse of the Roman Empire, multiple migrations of R1a were taking place eastward and westward;
these migrations gradually formed the current landscape of R1a in Europe. All 38 branches and their dat-
ings are listed in the Appendix of this paper; current distribution maps are shown in the body of the paper.
Keywords: Y Chromosome; Mutations; Haplotypes; Haplogroups; TMRCA; STR; SNP; R1a1
Introduction
The preceding article in this journal (Klyosov & Rozhanskii,
2012) provided data in support of the theory that haplogroup
R1a arose in Central Asia around 20,000 years before the pre-
sent (ybp), and described the prior history of the haplogroup,
which is directly related to the appearance of Europeoids (Cau-
casoids) ~58,000 years ago.
At some point, apparently between 20 and 15 thousand ybp,
the bearers of R1a began a migration to the west, through Tibet
and over the Himalayas. They arrived in Hindustan no later
than 12,000 ybp. They apparently continued their way across
the Iranian Plateau, along East Anatolia and the rest of Asia
Minor between 10,000 and 9000 ybp. By ~9000 ybp they ar-
rived in the Balkans and spread westward over Europe and to
the British Isles. At that point, R1a still had DYS392 = 13 in
most haplotypes, as did its sister haplogroup R1b. This marker
is very slow to change, and mutates on average once in 3500
conditional generations (that is once in 87,500 years in one
DNA-lineage; the concept of conditional generations of 25
years in each is described in “Materials and Methods” section).
More than 99% of the R1a haplotypes known today bear the
mutation giving DYS392 = 11. Somewhere along this extended
timescale, bearers of R1a or its upstream haplogroups devel-
oped the Pre-proto-Indo-European (Pre-IE) language and car-
ried it along during their journey from Central Asia to Europe.
Linguists date the earliest signs of Pre-IE in Anatolia at 9600 ±
600 ybp (Gamkrelidze & Ivanov, 1995; Gray & Atkinson, 2004;
Renfrew, 2000), which coincides with the data of DNA gene-
alogy for the migration of R1a bearers across Asia Minor that is
described in Klyosov and Rozhanskii (2012).
Some known archaeological cultures in the Balkans and Cen-
tral/Eastern Europe, dated 8000 - 7000 ybp (Bug-Dniester,
Vinča, Starčevo, LBK, etc.), can be attributed, as least in part,
to bearers of R1a. Yet they also can be attributed to other an-
cient haplogroups, such as I, J, E, G.
As the bearers of haplogroup R1b1a2 began to populate
*Corresponding author.

I. L. ROZHANSKII, A. A. KLYOSOV
Europe after 4800 ybp (the Bell Beakers and other R1b1 mi-
gratory waves, including, perhaps the Kurgan people, though
their identification and haplogroup assignment remain unclear,
and likely represented both R1b and R1a bearers in different
time periods), haplogroup R1a had moved from Central and
North-Western Europe to the Russian Plain around 4800 - 4600
ybp (Klyosov, 2011). From there, R1a migrated to the south,
east, and south-east as the historic Aryans (Klyosov & Roz-
hanskii, 2012).
We have to emphasize that the definition of the Aryans in
DNA genealogy is broader than the definition of Aryans in
linguistics and archaeology. According to linguists and archae-
ologists, Aryans were the ancient people who spoke Indo-
European Aryan or “steppe” languages; this early group split at
some point into the Indo-Aryan and Iranian languages. When
and where that split occurred is a matter of debate; linguists
generally claim that the split occurred immediately before the
Aryans moved to India and Iran, while some archaeologists
believe that it happened a thousand years before that.
In DNA genealogy, with its emphasis on genetic heritage, the
Aryans, as the putative bearers of Indo-European dialects, also
clearly carried haplogroup R1a, which apparently was brought
to India and Iran coincident in time with the spread of Indo-
European (IE) dialects in those same regions. That time is gene-
rally recognized around 3500 ybp, since studies of Gordon
Childe (Childe, 1926) and even earlier. In other words, R1a
likely represented a substantial portion of the Aryan linguistic
and cultural migration, the roots of which we can trace back to
Europe and to earlier migrations, following R1a ancient move-
ments. As soon as we accept such a definition, the time of the
split of the Aryans into “Indo-Aryans” and “Iranians” becomes
evident: migration is swift but language change is slow. The
“language split” must have occurred before the Aryan entrance
into Anatolia and northern Syria (Mitanni). This reasoning
suggests that the split was around 4500 ybp—as the archaeolo-
gists predicted.
Today’s linguists find IE languages in Anatolia and Mitanni,
the South Urals, Iran, India, beyond the Ural Mountains, and in
South Siberia. All these languages have the same Aryan roots.
These peoples developed common horse-breeding terminology
and shared essentially the same vocabulary for household items,
and for gods and religion, although sometimes, apparently, the
shared words ended up with contrasting meaning, as in India
and Iran, where some ancient Aryan good and evil gods some-
times had opposite meanings.
Currently, most of descendants of the ancient peoples who
carry European R1a live in Eastern Europe, primarily in Russia
(up to 63% of the population) and Poland, Ukraine, Belarus (up
to 57% of the population in the last three countries) (Underhill
et al., 2009; Balanovsky et al., 2008; Kayser et al., 2005; Khar-
kov et al., 2004, 2005).
Haplogroup R1a and its regional distribution and history
were a subject of many studies, which have been reviewed in
the preceding paper (Klyosov & Rozhanskii, 2012). R1a was
identified in excavated DNA in Germany with the dating of
4600 ybp (Haak et al., 2008), in Siberia, with the dating of
3800 - 3400 ybp (Keyser et al., 2009), in Tarim Basin, Xinjiang,
China, with the dating of ~4000 ybp (Li et al., 2010). Its distri-
bution was described in various parts of Eurasia (e.g., Wells et
al., 2001; Behar et al., 2003; Cinnioglu et al., 2004; Kharkov et
al., 2004; Sengupta et al., 2006; Balanovsky et al., 2008;
Sharma et al., 2009; Underhill et al., 2009; Abu-Amero et al.,
2009). However, only in the last few years, thank mainly to
efforts of Dr. Peter Underhill’s laboratory at Stanford Univer-
sity, Dr. Thomas Krahn of Family Tree DNA’s Genomics Re-
search Center, and researchers of 1000 Genomes Project, many
new SNPs in haplogroup R1a have been discovered; as indi-
cated by beginning letters M, L, and Z, respectively, in the
SNPs below. Their base haplotypes, mutation patterns and his-
tory have not been described in the literature as yet. Apparently,
only one attempt was made regarding R1a-M458 subclade
(Underhill et al., 2009), however, the authors employed the
erroneous “evolutionary mutation rate” (see Klyosov & Roz-
hanskii, 2012, and references therein) and obtained a grossly
overestimated date for M458 appearance as 10,700 ybp in
Eastern Europe (Poland), about 250% higher than the figure
obtained in our studies. Commonly, 250% - 300% exaggeration
in historical datings is a “feature” of the “evolutionary mutation
rate” method (ibid.).
This study focuses on DNA lineages, or branches, of hap-
logroup R1a in Europe, from the Atlantic to the Ural Mountains,
and on the history of their appearance and migrations across
and from Europe in the last ~9000 years (Appendix 1). The
methodology of the study, based largely on considerations of
extended 67- and 111-marker haplotypes, and involving prin-
cipal methods of DNA genealogy, is described in detail in the
preceding papers in this journal (Rozhanskii & Klyosov, 2011;
Klyosov & Rozhanskii, 2012) and in the Materials and Meth-
ods section of this article. Available databases containing tens
of thousands of 67-marker haplotypes are listed in Rozhanskii
and Klyosov (2011) and in this paper (Appendix 2).
The Old European Branch (≥9000 ybp)
(R1a-SRY10831.2+, M417−)
The Old European Branch of R1a in Europe has a distinct
ancient DYS392 = 13, as does its sister haplogroup R1b (see
the diagram below). However, while in R1b this allele is by far
the predominant one, DYS392 = 13 occurs in R1a in only 0.4%
of all haplotypes. All others have DYS392 = 11. Actually, only
twelve DYS392 = 13 haplotypes have been identified thus far
in the 67-marker haplotype format, and nine of them form the
Old European branch (the haplotype tree of this branch is
shown in Figure 1).
The following diagram shows the upper part of the hap-
logroup tree with respect to R1a and its most ancient subclades
(R1b is shown for a comparison) (ISOGG-2012, a fragment,
http://www.isogg.org/tree/ISOGG_HapgrpR.html) in its current
classification:
R M207
R1 M173
R1b M343
R1a L62/M513, L146/M420
R1a1 L120/M516, SRY10831.2
R1a1a M17, M198, M512
R1a1a1 L457, M417
The two haplotypes at the bottom of the tree in Figure 1 are
from Belarus and the US, both have positive SNP SRY10831.2
and negative M198, which firmly places them into the R1a1
subclade on the haplotype tree above. They two have their
combined haplotype (here and throughout the paper haplotypes
are given in the FTDNA format):
140

I. L. ROZHANSKII, A. A. KLYOSOV
Figure 1.
The 67-marker 9 haplotype tree for the Old European
Branch in Europe. This and all other haplotype trees in
this paper are composed from haplotypes listed in the
IRAKAZ database (Rozhanskii et al., 2012). The upper
7-haplotype branch has SNP M417–, M198+ (R1a1a),
the lower 2-haplotype branch has M198–, SRY-
10831.2+ (R1a1).
13 25 15 10/11 12 12 12 13 12 13 13 31/33—16 9 9 11 11
24/25 16 20/21 29/34 12/13 16 16 16—10/11 11/12 19 23 15
15/16 17/20 17/18 35 38 11/12 11—11 8 15 17 8 11 10 8 11/12
10 12 22 22 17 10 12 12 15 8 12 22/23 22/23 13 12 11 13/14
10/11 11 12 12/13
The 25 mutations between them are translated to 25/.12 =
208 → 262 conditional generations (25 years each), or 6550
years. If they belong to the same DNA-lineage, their most re-
cent common ancestor lived ~3300 ybp. If they belong to dif-
ferent lineages, their common ancestor would have lived much
earlier.
The upper, rather tight series of haplotypes in Figure 1 all
have positive M198, but negative M417, which places them in
the R1a1a subclade. Their base haplotype is as follows:
13 25 15 11 13 14 12 12 10 14 13 31—16 9 10 10 11 25 14
19 31 12 15 15 15—10 11 19 23 16 16 17 19 36 38 11 11—11
8 17 17 8 12 10 8 12 10 12 22 22 15 10 12 12 13 8 13 23 22 12
12 11 13 11 11 12 12
All seven haplotypes have 78 mutations from the base hap-
lotype, which gives 78/7/.12 = 93 → 103 conditional genera-
tions, or 2575 ± 390 years to their common ancestor. However,
this date actually points to a time when the population ex-
panded. In reality, the base haplotype shown above differs from
the two M198– haplotypes by 40 and 41 mutations respectively;
if they belonged to the same subclade, that would sets them
apart by at least 12,300 and 12,800 years. Since they belong to
different subclades (R1a1 and R1a1a), their common ancestors
lived more than 6400 years before the present, and not less than
the age of the R1a1a-M17 subclade (at least 9000 ybp).
We do not know exactly where their common ancestor lived,
since members of this branch are scattered in Europe from the
descendants of those R1a bearers who came to the Balkans
Balkans to the British Isles (Map 1). It may be that they are
e a particular
su
Principal European R1a Subclades
As wahanskii
(2
nstream subclade in Europe was R1a1a1-M417,
w
R1a1a1 M417
~9000 ybp at the European end of their migration route from
Central Asia (Klyosov & Rozhanskii, 2012).
In this paper, unless we specifically describ
bclade with its particular SNP, we will use R1a as a common
overall designation of all downstream subclades of the hap-
logroup.
s described above and in Klyosov and Roz
012), R1a arose around 20,000 ybp in Central Asia, migrated
(in part) to Europe over the following 11,000 years, and arrived
in Europe, as R1a1 and R1a1a subclades, more than 7 thousand
years or, more probably, ~9000 ybp. There are only a few de-
scendants of these ancient subclades which are currently identi-
fied in Europe.
The next dow
hich arose ≥7600 ybp (see below). Its principal downstream
subclades are as follows:
R1a1a1i L664– (the Tenths, NW branch)
R1a1a1i L664+
R1a astern branch) 1a1h Z93 (South-E
R1a1a1g Z283 (Eurasian branch)
R1a1a1g1 M458 (European branch)
R1a1a1g2 Z280 (Central Eurasian)
R1a1a1g3 Z284 (Scandinavian)
The Tenths, or the North-Western branch, is the oldest sub-
cl
lotype tree is shown in Fig-
ur
ain purpose of presenting the overall tree in Figure 2
is
North-Western Branch (the Tenths), 4575 ybp
s
ade of R1a-M417, identified thus far. It arose at least 6300
ybp (see below) and has a distinct mutation DYS388 = 10 as
distinct from the DYS388 = 12 in majority of R1a haplotypes.
Subclade Z93 separated from the rest of R1a around 5700 ybp.
It migrated from Europe or from the Russian Plain no later than
4800 ybp. The migration was to the south (to the Middle East)
and south-east (Iran and India), where bearers of Z93 are the
predominant R1a bearers today. Subclade Z283 arose ~5500
ybp and populated Eastern Europe, part of Central Europe, and
further east to the Ural Mountains.
The overall 111-marker R1a hap
e 2. The purpose of the Figure is not in details of positions of
individual haplogroups, but to show the overall shape, configu-
ration of the tree, and relative positions of its subclades. A few
of M420 and M417 haplotypes, available in the 111-marker
format, sit at the very base/core of the tree, since they de-
scended from the most ancient European R1a ancestors. The
principal subclades of the Z283 Eurasian super-branch are lo-
cated around the tree—Z280, Z284, M458. Some of them con-
sist of two or several more sub-branches, which are described
below.
The m
to show that the SNP (single nucleotide polymorphism) sub-
clades and STR (short tandem repeat) branches form a robust
mutually supporting system. They are two sides of the prover-
bial coin; both are necessary for a justified, meaningful, and
advanced analysis of haplotype datasets.
This is one the most distinct of R1a branches. All haplotype

I. L. ROZHANSKII, A. A. KLYOSOV
142
Map 1.
European Branch, referred to birthplaces of the furthermost ancestors recorded in available data-
The Old
bases. Red pins denote a branch with SNP M198−, SRY10831.2+. Blue pins mark a branch with M417−,
M198+.
Figure 2.
rker haplotype tree of R1a haplogroup, composed of 251
f this branch have DYS388 = 10 (hence, the Tenths), and two-
live in the USA (with roots mainly in England, Ireland, Scan-
e is the old branch (with a common ances-
to
-branches as follows:
24 16 15 19 20 33 39 12 11—11
23 15 16 18 19 34 37 12 11—11
a common ancestor of 2125 ± 280
An 111-ma
haplotypes. Most of the tree is taken by the Eurasian super-branch Z283,
of which branches Z280 (Central Eurasian), Z284 (Scandinavian), and
M458 (European) are shown. Another two principal branches are Z93
(South-Eastern) and L664 (North-Western branch, the Tenths). The
L664 branch shown at 7 o’clock.
o
thirds of them are located in England, Ireland and Northern
Germany (Map 2), and almost the same number of their bearers
dinavia, Germany).
The haplotype tree (Figure 3) splits into four major lineages,
or sub-branches. On
r of ~4300 ybp), which in turn splits to subclades L664-
(4100 ybp; though, this SNP seems to be rather unstable) and
L664+ (4300 ybp), then the young branch, with L664+ (2250 ±
250 ybp), and two DYS448 null-mutated branches, of 1500 ±
325 ybp and its downstream of 260 ± 60 ybp, the last marks the
arrival of the Tenths to the USA (Klyosov, 2010a, 2010b,
2012).
The base 111-marker haplotype of the old branch splits into
two sub
13 25 16 10 11 14 12 10 10 13 11 30—15 9 10 11 11 24 14
19 31 12 15 15 16—11 11 19
8 17 17 8 12 10 8 11 10 12 22 22 16 10 12 12 14 8 14 23 22 12
13 11 13 11 11 12 13—31 15 9 16 12 26 27 18 13 12 1212 11
10 12 11 10 11 11 31 12 15 24 12 9 10 19 15 20 12 23 15 12 15
25 12 23 19 10 15 18 9 11 11
13 25 15 10 11 14 12 10 10 13 11 31—15 9 10 11 11 25 14
19 31 12 14 14 17—12 11 19
8 17 17 8 12 10 8 11 10 12 22 22 15 10 12 12 13 8 14 23 22 12
12 11 13 11 11 14 13—32 15 9 15 12 26 27 19 12 121212 10 9
12 11 10 11 11 31 12 14 24 13 9 10 19 15 20 12 24 15 12 16 24
12 24 19 10 15 19 9 11 11
which differ by as much as 33 mutations. One branch (shown
first) is likely L664−, with
ybp (.139 mutations per marker), another, likely L664+, is 2025
± 260 years “old” (.133 mutations per marker). The distance
between them sets these two lineages apart by 33/.198 = 167 →

I. L. ROZHANSKII, A. A. KLYOSOV
Map 2.
The map of the North-Western branch, depicting birthplaces of the furthermost ancestors recorded in available
Yellow pins denote an old branch. Deep-blue and light-blue pins mark younger branches. databases.
Figure 3.
A haplotype tree of the Tenth’s (DYS388 = 10), composed of 285 o
The dataset was provided by Martin Voorwin-
0 years. This placed their
common ancestor by (2125 + 2025 + 5000)/2 = 4575 ybp.
haplotype tree (ISOGG-2012)
f
67-marker haplotypes.
den and updated with the Prendergast family series in January 2012.
The younger branch on the left all has SNP L664+, where determined;
the branch on the 3 o’clock (the Prendergast family with a common
ancestor ~1000 ybp) has both L664+ and L664−, which raises a ques-
tion on technical (methodological) stability of the L664−. The old
branch has both L664+ and L664−.
An analysis of a more extended dataset of 67-marker haplo-
200 conditional generations, or 500
types of the Tenths gave 4300 years to their common ancestor,
which is the same value within the margins of error, and
showed that the divergence of L664+ and L664− branches oc-
curred around 4100 ybp, or almost from the common ancestor
of currently living Tenths (Klyosov, 2012).
The robustness of the calculations despite the small number
of extended haplotypes is clear. Twenty-one 67-marker haplo-
types show 191 mutations, which give 191/21/.12 = 76 → 83
generations, or 2075 years to a common ancestor. The same
haplotypes in the 111-marker format give 324 mutations, that is
324/21/.198 = 78 → 85 generations, or 2125 years to the com-
mon ancestor. Obviously, the margins of error of 2075 ± 260
and 2125 ± 240 ybp may be somewhat excessive for the calcu-
lations.
The combined 67-marker base haplotype of the Tenths dif-
fers from the Old European base haplotype (see above) by 29
mutations, which places their common ancestor at ≥7600 years
before the present. Since they belong to different subclades,
M417− and M417+, their common ancestor lived 7600 ybp
only if the “depth” of the M417+ subclade is no more than
7600 years. In that case, the Tenths actually initiated the
M417+ subclade.
South-Eastern Branch (Z93), ~5700 ybp
According to the

I. L. ROZHANSKII, A. A. KLYOSOV
R1a1a1h Z93
R1a1a1h1 Z94
R1a1a1h1 L342.2
R1a1a1h1a L657
subclade Z93 has tades, Z94, L342.2 and
657. All four subcnly the Middle East,
hree downstream subcl
lades populate maiL
India, and some locations in the eastern part of the Russian
Plain, hence, it is called the South-Eastern branch (Map 3). It
arose in Europe ~5700 ybp and migrated along with the future
Aryans across the Russian Plain to the south, east, and south-
east between 5000 and 3500 ybp.
The haplotype tree of the R1a-Z93 clade is shown in Figure
4. Its Jewish R1a1 branch includes more than half of the whole
tree (104 haplotypes out of 203 haplotypes); all the Jewish
haplotypes are related to the downstream subclade L342.2. The
Bashkir and the Kyrgyz branches also have the same L342.2. In
fact, L342.2 is the predominant subclade in the Z93 clade, em-
bracing 98% of its haplotypes. Only a few haplotypes on the
tree and one English lineage of closely related individuals are
L342.2 negative, which tells us the initial state of Z93.
Bearers of Z93 are spread from England through Armenia,
Oman, and Kuwait, all the way to Khakassia in South Siberia;
the latter (or his ancestors) apparently was a migrant from the
Russian Plain. Six currently known representatives have the
following base haplotype
13 24 16 11 11 15 12 12 10 13 11 30—15 9 9/10 11 11 24 14
20 31 12 15 15 15—11/12 11 19 22/23 16 16 18 17 35 40 13 11
—11 8 17 17 8 12 10 8 11 10 12 22 22 15 10 12 12 13 8 13/14
23 21 12 12 11 13 10 11 12 13
All six have 111 mutations from the base haplotype which
gives 111/6/.12 = 154 → 177 generations, or 4425 ± 630 years
to their common ancestor. In fact, they belong to different mi-
nor sub-branches, which split from the trunk at the very begin-
ning of the Z93 subclade, and their actual TMRCA is expected
to be 5000 years (or more) before the present.
Most of the haplotype tree is L657 negative. An example of
L657− base haplotype of an available 20-haplotype Asian
branch in the 111 marker format is given below:
13 25 16 11 11 14 12 12 10 13 11 30—15 9 10 11 11 24 14
20 32 12 15 15 16—11 12 19 23 16 16 18 19 34 38 13 11—11
8 17 17 8 11 10 8 11 10 12 22 22 15 10 12 12 13 8 14 23 21 13
12 11 13 11 11 12 13—32 15 9 15 12 26 27 19 12 12 13 12 10
9 12 11 10 11 11 30 12 13 24 13 9 10 19 15 19 11 23 15 12 15
24 12 23 19 10 15 17 9 11 11
An example of L657+ base haplotype of an available Asian
branch is as follows:
13 25 16 10 11 14 12 12 10 13 11 30—16 9 10 11 11 24 14
20 32 12 15 15 17—12 11 19 23 15 17 17 19 36 40 14 11—11
8 17 17 8 11 10 8 11 10 12 22 22 15 10 12 12 13 8 12 23 21 12
12 11 13 11 11 12 12
The first base haplotype is apparently ancestral to the branch,
consisting of haplotypes from Turkey, Saudi Arabia, Qatar, the
United Arab Emirates, Palestine, India, Kazakhstan, Iraq, Paki-
stan, Poland, Germany, and one from Tatar Russia. 20 of 67-
marker haplotypes in the branch contain 354 mutations, which
place their common ancestor at 354/20/0.12 = 148 → 174 gene-
rations, or 4350 ± 490 ybp. Fourteen 111-marker haplotypes in
the branch contain 381 mutations, which place their common
ancestor at 381/14/.198 = 139 → 162 generations, or 4050 ±
460 ybp.
The second sub-branch is non-symmetrical, and contains six
haplotypes from Saudi Arabia with a common ancestor of only
315 ± 110 ybp, and one very different haplotype from India,
with an apparent common ancestor for all L657+ haplotypes of
2700 ybp. There are 16 mutations between the two L657+ and
Map 3.
The map of the South-Eastern branch, depicting birthplaces of the furthermost ancestors recorded i
available databases. Magenta pins denote a parent Z93 branch, red and green pins correspond to Bash-
rgyz branches, respectively. The rest of the branch (L342.2 positive) is marked by deep-blue
n
kir and Ky
pins. The most represented Ashkenazy Jewish branch (not shown) covers densely an area of historical
Polish-Lithuanian Commonwealth.
144

I. L. ROZHANSKII, A. A. KLYOSOV
Figure 4.
A haplotype tree of the South-Eastern branch (Z93), composed of
203 of 67-marker haplotypes. Practically all haplotypes (excep
−) are assigned to L342.2+ subclade. There are two
L-
mIt
ight relate to the appearance of the L342.2 subclade.
0 12 12 13 8 14 23 21 13
12
42.2) to 4800 years before the pre-
se
4 with the 67 markers reported by FTDNA. Eight Kyrgyz
ha
xxx 10 12 12x 8 14 xx 12 12 11 xx 11 12 12
hich corre-
sp
111-marker haplotype has the following alleles:
4 9 11 11 11 24 14
0 35 38 14 11—11
8 17 17 8 12 10 8 11 10 12 22 22 15 10 12 12 14 8 14 23 21 12
0 15 20 11 23 15 12 15
extended one among R1a1 haplotypes, both in num-
ber of h of all
Europea estern
br
t
four L342.2
L657+ branches, of 7 and 17 haplotypes each. The tight, numerous
young branch on the left is the Jewish R1a1 branch. The Bashkir
(10 haplotypes) and Kyrgyz (7 haplotypes) branches are also shown.
657− branches in the 67-marker format, which places a com
on ancestor of the two base haplotypes at 5300 years ago.
m
The R1a-L342.2 Bashkir branch of 10 haplotypes points to
their common ancestor of about 1125 ± 190 ybp. Their 111-
marker base haplotype is as follows:
13 24 16 11 11 15 12 12 12 13 11 31—15 9 10 11 11 24 14
20 31 12 15 15 15—11 11 19 23 16 15 18 19 35 39 14 11—11
8 17 17 8 12 10 8 11 10 10 22 22 15 1
11 13 11 11 12 13—33 15 9 15 11 26 28 19 11 12 1212 10 9
12 11 10 11 12 30 13 14 24 13 9 10 19 15 19 12 24 14 13 15 24
12 23 19 10 15 17 9 11 11
It has 28 mutations compared with the 111-marker predomi-
nantly Middle Eastern L657− base haplotype, which places
their common ancestor (L3
nt.
The Kyrgyz branch is based mostly on the SMGF database
(www.smgf.org), which employs another format, and shares
only 3
plotypes are known in the 67- and 111-marker formats, but
six of them belong to a rather recent lineage (650 ± 150 ybp),
with possible origin in Altay. Since some alleles in the base
haplotype of the entire branch are uncertain, they are marked by
“x”:
13 25 16 1111 14 12 12 10 14 11 32—15 9 11 11 23 14 2131
12 1515 16—11 11 19 23 16xxxxx15 11—11 8 17 17 8 x 10 8
xx 12
A common ancestor of the Kyrgyz branch lived 2100 ± 250
ybp; this and the other branches of the L342.2 share a common
ancestor within the range of 4400 ± 500 ybp, w
onds to the ancestor of L342.2 itself.
Finally, the large Jewish R1a-L342.2 branch with its base
12 11 13 10 11 12 13—32 15 9 17 12 27 27 19 12 12 12 12 10
9 12 11 10 11 11 30 12 12 25 13 9 10 2
25
13 25 16 10 11 14 12 12 10 13 11 30—1
20 30 12 12 15 15—11 11 19 23 14 16 19 2
12 23 19 10 15 17 9 11 11
This group descends from a common ancestor who lived
only 1300 ± 150 ybp. It arose in Germany, but as part of a line-
age with a common ancestor living 4475 ± 400 ybp, or what
has been called the “Abraham times” (Klyosov & Rozhanskii,
2012).
It is of interest to compare which base haplotype is closer to
the Jewish base haplotype—the Middle Eastern or the Bashkir
base haplotype. It would seem that the Middle Eastern haplo-
type should be closer. Indeed, the mutational distances in the
111-marker format are 30 and 40, respectively. The common
ancestor of the Jewish base and the Middle Eastern base, and of
the Jewish base and the Bashkir base lived 4900 ± 500 and
4400 ± 500 ybp, respectively—or at the same time, within the
margin of error. It fits well with the 4800 ybp obtained above
for the L342.2 subclade. The common ancestor of all these
L342.2 branches lived at the time of the Aryan migrations from
Eastern Europe across the Russian Plain on their way to the east
and south.
Eurasian Branch (Z283), ~5500 ybp
The Eurasian R1a branch Z283 includes haplotypes from
Europe and from the Asian part of Russia which spreads to
Pacific Ocean on the east. It is also found in low frequencies in
Asia Minor and among ethnic Armenians. The Eurasian branch
is the most
aplotypes and in its territory. It embraces 86%
n R1a1 haplotypes, while the Tenths (North-W
anch) take only 12%. The rest are minor subclades, such as
the Old Europeans. The Z93 subclade, with its mainly Asian
haplotypes, encompasses 10% of the European R1a1 haplo-
types, and it was not included in the above figures. Three coun-
tries—Poland, Russia and Germany—embrace 55% of all the
Eurasian branch, with the rest mainly in Ukraine, Belarus, the
Czech Republic, Slovakia, Slovenia, Lithuania, Finland, Hun-
gary, and Italy.
R1a1a1g Z283 (Eurasian branch)
R1a1a1g2 Z280 (Central Eurasian branch)
R1a1a1g2a P278.2 (West Carpathian)
R1a1a1g2b L365 (North European)
R1a1a1g2c L366 (Central Eurasian-3)
R1a1a1g2d Z92 (North Eurasian)
R1a1a1g2d (North Carpathian)
R1a1a1g2e L784
R1a1a1g3 Z284 (Scandinavian branch)
R1a1a1g3a L448 (Young Scandinavian)
R1a1a1g3a1 L176.1 (Scottish Highland)
R1a1a1g3b Z287 (Old Scandinavian)
R1a1a1g1 M458 (European branch)
R1a1a1g1a Central European branch
R1a1a1g1a1 L1029
R1a1a1g1b L260 (West Slavic branch)
The aboveades of diagram shows the downstream subcl

I. L. ROZHANSKII, A. A. KLYOSOV
Eurasian is the Central
Eurasia which include al-
most half followed first
by the Scan and its subclades), with
one-third 458 (and its
ubclades)lo-
trees are shown here not for a detailed analysis but for
their overheir own
rather distbranches
which do not have ato Fig-
ur
Tpe
bye
timt
pl
1111 12 13
The CPlain
base hations
separate the common3 → 34 generations,
or 850 years, and their common ancestor lived (4600 + 3500 +
branch Z283. The most populous one
n branch (Z280) with its subclades,
of all Eurasian (Z283) haplotypes. It is
dinavian branch (Z284
of the Eurasian haplotypes; and then by M
with 21% of the Eurasian haplotypes. A few haps
types from Canada, Germany, Switzerland, England, Poland
and Turkey have been proven to belong to the parent Z283
branch.
Central Eurasian Z280 Subclade
(4900 ± 500 ybp)
Figure 5 shows the 67-marker 801-haplotype tree of the cen-
tral Eurasian branch (Z280); Figure 6 presents the 111-marker
tree of 96 haplotypes, currently available in this extended for-
mat. The
all shape. All four available SNPs have t
inct branches on the trees, along with other
n assigned SNP as yet (see legend
es 5 and 6).
Overall, there are 13 currently identified branches within
subclade Z280. A common ancestor of Z280 lived 4900 ± 500
ybp, and had a base (ancestral) 111-marker haplotype as fol-
lows:
13 25 16 11 11 14 12 12 10 13 11 30—15 9 10 11 11 24 14
20 32 12 15 15 16—11 12 19 23 16 16 18 19 35 38 14 11—11
8 17 17 8 12 10 8 11 10 12 22 22 15 10 12 12 13 8 14 23 21 12
12 11 13 11 11 12 13—32 15 9 15 12 26 27 19 12 12 12 12 10
9 12 11 10 11 11 30 12 13 24 13 9 10 19 15 20 11 23 15 12 15
24 12 23 19 10 15 17 9 11 11
Some of the Z280 subclade bearers can be assigned, at pre-
sent, to a rather loose parent branch, the Central Eurasian group
proper, descending directly from the above base haplotypes.
Base haplotypes of the most populous branches are shown be-
low, along with the timespans to their common ancestors.
The Russian Plain (RP) Branch, or Central
Eurasian-1 (CEA-1) Branch (4600 ± 500 ybp)
13 25 16 11 11 14 12 12 10 13 11 30—15 9 10 11 11 24 14
20 32 12 15 15 16—11 11 19 23 16 16 19 19 35 38 14
11—11 8 17 17 8 12 10 8 11 10 12 22 22 15 10 12 12 13 8
14 23 21 12 12 11 13 11 11 12 13
he RP base haplotype differs from the Z280 base haploty
only two mutations (425 “lateral” years, that is an averag
e required for these two mutations to occur), as marked; i
aces their common ancestor at (4900 + 4600 + 425)/2 = 4960
ybp, which is Z280 itself (4900 ± 500 ybp) within the margin of
error.
Central Eurasian-2 (CEA-2) Branch
(3500 ± 400 ybp)
13 25 16 11 11 14 12 12 10 13 11 30—15 9 10 11 11 24 14
20 31 12 15 15 16—11 1119 23 16 16 18 19 34 38 14 11—11 8
17 17 8 12 10 8 11 10 12 22 22 15 10 12 12 13 8 13 23 21 12
12 11 13
entral Eurasian-2 branch differs from the Russian
plotype by 4 mutations (marked above). Four muta
ancestors by 4/.12 = 3
Figure 5.
A haplotype tree of the Central Eurasian branch (Z280), composed of
801 of 67-marker haplotypes. The North European (L365), Central
Eurasian-3 (L366), West Carpathian (P278.2), and North Eurasian (Z92)
branches have their identified SNPs. The tree contains other branches
which do not have their SNPs identified as yet, and some of them are as
follows, clock-wise: North Carpathian (NC), Balto-Carpathian-1 (BC-
1), West Eurasian-3 (WEA-3), Central Eurasian-1 (CEA-1), WEA-1,
CEA-2, Eastern Carpathian (EC), BC-3, BC-2.
Figure 6.
A haplotype tree of the Central Eurasian branch (Z280), composed of
96 of 111-marker haplotypes. The North European (L365), West Car-
pathian (P278.2), and North Eurasian (Z92) branches are indicated, as
well as (clock-wise): Balto-Carpathian (BC), North Carpathian (NC),
Eastern Carpathian (EC).
850)/2 = 4475 ybp. This is the time of the Russian Plain com
mon ancestor (4600 ± 500 ybp) within the margin of error.
-
146

I. L. ROZHANSKII, A. A. KLYOSOV
entral Eurasian-3 (CEA-3) Branch
12 22 22 15 10 12 12 12 8 13 23 21 12
(RP
branch)stor at
~4825 th the
Russian Plain (RP) an (Z280) common
an
100 ± 500 ybp)
type by three mutations (650 years), hence, their
common ithin the
margin of-branch
(WEA-1) with a com0 ybp. It devi-
at
).
19 23 16 16 17 20 36 39 12 11—11
8 17 17 8 12
12 11 13
The base haplotyp280 by 8 mutations,
w
14 11—11
8 17 17 83 21 12
12 11 13
Twelve mutations base Z280 haplo-
ty
arpathian (BC) Branch (4300 ± 500 ybp)
Eastern Carpathian (EC) Branch
(2600 ± 300 ybp)
13 25 16 11 11 14 12 12 10 13 11 30—15 9 10 11 11 24 14
20 32 12 15 15 16—11 11
C
(2500 ± 350 ybp)
13 25 16 11 11 14 12 12 10 13 11 30—15 9 10 11 11 24 14
20 31 13 15 15 16—11 12 19 22 16 16 18 19 33 39 13 11—11
17 17 8 12 10 8 11 10 12 10 8 10 10 12 22 22 15 10 12 12 13 8 14 23 21
1111 12 13
e differs from that of Z
8
12 11 13 11 11 12 13
The Central Eurasian-3 is a minor branch, characterized by
SNP L366. It differs from the parent base haplotype hich corresponds to 1800 years, and translates to a common
ancestor at 4650 ybp. Again, it is well within margin of error
for TMRCA of the entire Central Eurasian branch.
by 11 mutations. This places the common ance
ybp, which is within the margin of error for bo
and the Central EurasiWestern Carpathian (WC) Branch
(2600 ± 300 ybp)
13 25 15 10 11 15 12 12 11 13 11 29—15 9 10 11 11 24 14
20 31 12 15 15 16—11 12 19 23 17 16 17 19 35 39
cestors.
Western Eurasian (WEA) Branch
(4
12 10 8 11 11 12 22 22 15 10 12 12 13 8 13 2
12 11 12 13
(2775 years) from the
13 25 16 11 11 14 12 12 11 13 11 30—15 9 10 11 11 24 14
20 33 12 1515 16—11 12 19 23 16 16 1819 34 38 14 11—11 8
17 17 8 12 10 8 11 10 12 22 22 15 10 12 12 13 8 14 23 21 12
12 11 13 1111 12 13
The Western Eurasian base haplotype differs from the Z280
base haplo
pe place the common ancestor of the Western Carpathian
branch at 5100 ybp. Its assignment is also confirmed by SNP
P278.2.
The above branches are spread almost uniformly over most
of
ancestor lived 4825 ybp. This is Z280 itself w
error. The WEA branch has a distinct sub
mon ancestor of 2300 ± 30
Central and Eastern Europe; however, they are virtually ab-
sent in Scandinavia (Map 4).
Balto-C
es by 10 mutations from the parent WEA base haplotype, or
by 2275 “lateral” years. This places their common ancestor
with the parent branch at 4300 ybp, i.e., the WEA parent branch
itself (4100 ± 500 ybp
13 25 16 10 11 14 12 12 10 13 11 30—15 9 10 11 11 24 14
20 32 13 15 15 16—11 12 19 23 16 16 18 18 34 37 13 11—11
Map 4.
The map of the parent Central Eurasian, Western Eurasian, Eastern and Western Carpathian branches, de-
picting birthplaces of the furthermost ancestors recorded in available databases. Yellow pins denote the par-
ent branch, pins of other colors correspond to junior branches.

I. L. ROZHANSKII, A. A. KLYOSOV
8 17 17 8 12 10 8 11 10 12 22 22 15 10 12 12 13 8 14 23 21 12
12 11 13 11 11 12 13
The Balto-Carpathian branch deviates by 6 mutations (1325
“lateral” years) from the Z280 base haplotype. The BC branch
has two distinct sub-branches (BC-1 and BC-2) with common
ancestors of 2550 ± 300 and 2200 ± 250 ybp, respectively.
They deviate by 4 and 9 mutations from the parent (BC) base
haplotype, or by 850 and 2025 “lateral” years respectively. This
places their common ancestor with the parent branch at 3850
and 4300 ybp, or the BC parent branch itself (4300 ± 500 ybp).
In contrast to the “core” branches of Z280, the Balto-Car-
pathian shows distinct geographical distribution, as reflected by
its name (Map 5). Its BC-2 sub-branch is characteristic for
Slovenians and Croatians, being their “trademark” genealogical
lineage.
Northern Eurasian (NEA) Branch
(4450 ± 450 ybp)
This branch is characterized by SNP Z92. It consists of two
major sub-branches. The older branch (NEA-1) (3600 ± 400
ybp) has the following base haplotype:
13 25 16 11 11 14 12 12 11 13 11 30—15 9 10 11 11 24 14
20 33 12 14 15 16—11 12 19 23 16 16 18 18 34 40 14 11—11
8 17 17 8 12 10 8 11 10 12 22 22 15 10 12 12 13 8 13 23 22 12
12 11 13 11 11 12 13
Its base haplotype deviates from that of Z280 by 9 mutations
(2025 years), which places their common ancestor at 5260 ybp,
within the margin
The base haplo
300 ybp) is:
13 25 16 11 11 15 12 12 10 13 11 30—15 9 10 11 10 25 14
20 32 12 14 14 16—12 12 19 23 15 16 18 20 34 38 13 11—12
8 17 17 8 12 10 8 11 10 12 22 22 15 10 12 12 13 8 13 23 22 12
12 11 13 11 11 12 13
It has 12 mutations (2775 years) from the base Z280 haplo-
type and 13 mutations (3025 years) from the NEA-1 base hap-
lotype. The common ancestor of these two sub-branches of Z92
lived 4450 ± 450 ybp.
Present-day bearers of both NEA branches populate Lithua-
nia, Belarus, and the north-eastern part of Poland (Map 6). It is
the most widespread genealogical lineage among ethnic Rus-
sians.
Northern European (NE) Branch
(2600 ± 350 ybp)
13 25 15 11 11 14 12 12 10 13 11 30—17 9 10 11 11 23 14
20 33 13 15 15 16—11 11 19 23 16 15 18 19 34 38 14 11—11
8 17 17 8 12 10 8 11 10 12 21 22 16 10 12 12 13 8 14 24 21 13
12 11 13 1111 12 13
The Northern European base haplotype differs by 13 muta-
tions (3025 “lateral” years) from that of Z280, and is inde-
pendently defined by SNP L365. It again places its common
ancestor at 5260 ybp, which is Z280 itself.
Northern Carpathian (NC) Branch
(2150 ± 300 ybp)
9 10 11 11 24 14
5 40 14 11—11
3 8 14 23 21 12
of error from 4900 ± 500 ybp for Z280 itself.
type of the younger branch (NEA-2) (2350 ±
13 25 17 11 11 14 12 12 10 13 11 31—14
20 32 13 15 15 16—11 12 19 23 16 16 18 19 3
9 17 17 8 12 10 8 11 10 12 22 22 15 10 12 12 1
Map 5.
The map of the Balto-Carpathian branch, depicting birthplaces of the furthermost ancestors recorded in
branch, deep-blue and green pins correspond to BC-1 available databases. Yellow pins denote the parent
and BC-2 sub-branches, respectively.
148

I. L. ROZHANSKII, A. A. KLYOSOV
Map 6.
The map of the Northern Eurasian branch, depicting birthplaces of the furthermost ancestors recorded in
available databases. Yellow pins denote the older branch (NEA-1), magenta pins correspond to NEA-2.
12 11 13 11 11 12 13
This base haplotype differs by 7 mutations (1550 years) from
the base Z280 haplotype.
There is also a minor branch, which is defined by subclade
L784 (2500 ± 450 ybp):
13 24 17 11 11 14 12 12 10 14 11 30—15 9 10 11 11 24 14
20 31 13 15 15 16—11 12 19 23 16 17 17 19 34 39 12 12—11
8 17 17 8 12 10 8 11 10 12 22 22 15 10 12 12 13 8 14 23 21 12
12 11 13 11 11 12 13
The latter branches seem to be geographically limited to pre-
sent day Poland and Eastern Germany, and—in lower frequent-
cies—the Baltic countries, Ukraine, and Russia (Map 7).
Overall Z28
The eight base
Plain, West Eurasian, Balto-Carpathian, North European, East
Carpathian, North Eurasian, North Carpathian, and West Car-
pathian) differ among themselves by 68 round-up or 64 full
mutations (some mutations are fractional), resulting in the Z280
base haplotype, listed above. It gives 64/8/.12 = 67 → 72 gene-
rations, or 1800 years down from their average “age” (3100
years). This results in 4900 years from their common ancestor,
which is Z280 itself (4900 ± 500 ybp, see above).
European Subclade M458 (4200 ± 450 ybp)
This subclade consists of two principal branches, the Central
European (CE) and West Slavic (WS). A division between
them is clearly seen on their haplotype tree (Figure 7).
The CE branch, in fact, consists of two sub-branches, both
arose 2900 ± 400 ybp. One of them (CE-1) has the following
base haplotype:
13 25 16 10 11 14 12 12 11 13 11 29—16 9 10 11 11 23 14
20 32 12 15 15 16—11 11 19 23 17 16 18 19 34 38 14 11—11
8 17 17 8 11 10 8 12 10 12 21 22 15 10 12 12 13 8 14 25 21 13
12 11 13 11 11 12 13
It differs by 14 mutations (3325 years) from the Z280 base
haplotype (see above), which places their common ancestors at
≥5600 ybp. The “equal or higher” sign here indicates that the
difference between them cannot be less that the “age” of sub-
clades Z280 and M458.
The second branch (CE-2) differs from CE-1 in only two loci,
having there three mutations: a multi-copyuplication 12-15-
15-16 → 12-12-15-15-15-16 in DYS464 (presumably a one-
in CDYb. These
E-1 and CE-2 at
3100 ybp, if both branches arose independently from their com-
mon ancestor. It might be thought that CE-2 is a CE-1 down-
stream branch, which arose immediately after CE-1, within the
indicated margin of error in their dating. This is likely, since
even the 67 marker haplotype tree cannot cleanly separate the
CE branches (see Figure 7). Their haplotypes are nearly iden-
tical statistically, except that they descended from slightly dif-
ferent ancestral (base) haplotypes. Also, there is a downstream
subclade L1029 within the CE branch which embraces the most
of bearers of both the sub-branches. Apparently, it arose at the
same time as the CE branch itself.
In contrast, the Western Slavic branch (L260) is quite distinct.
It does not include any CE haplotype on the tree (Figure 7). It
arose 2700 ± 300 ybp, and its base haplotype is as follows:
13 25 17 10 10 14 12 12 10 13 11 30—16 9 10 11 11 23 14
20 31 12 15 16 16—11 11 19 23 16 16 18 19 34 39 13 11—11
0 Base Haplotype and Its “Age”
haplotypes of the main branches (Russian
time event), and a two-step mutation 38 → 40
three mutations place a common ancestor of C
d

I. L. ROZHANSKII, A. A. KLYOSOV
Map 7.
The map of the Northern European (green pins) and Northern Carpathian (magenta pins) branches, depict-
ing birthplaces of the furthermost ancestors recorded in available databases. Blue pins denote the minor
branch, which is characterized by SNP L784.
Figure 7.
A 67-marker haplotype tree of the R1a-M458 subclade, composed of
352 haplotypes. The Central European (CE) branch takes the right-hand
side and the upper left-hand side. The distinct Western Slavic (WS)
branch is on the left.
8 1
12 1
2 muta-
tio
Slavic branch, which makes the M458 subclade 4200 years
the Z280 base haplotypes
(1
7 17 8 12 10 8 12 10 12 22 22 15 10 12 12 13 8 14 25 21 12
1 13 11 11 12 13
It differs by 14 mutations (3325 years) from the Z280 base
haplotypes, which places their common ancestor at the same
≥5600 years (see the explanation above). There are 1
ns between the CE-1 base haplotype and that of the Western
ed on available haplotypes. “old” at least bas
The European subclade M458 shows nearly the same geo-
graphical distribution as the Central Eurasian Z280, but statis-
tically it is more densely represented on its western edge (see
also Underhill et al., 2009), reaching up to 70 % of all R1a1
bearers in the Czech Republic (Map 8). On the other hand, it is
relatively rare among ethnic Lithuanians and Slovenians, who
belong predominantly to Z280 subclade.
Scandinavian Subclade Z284 (4300 ± 500 ybp)
This subclade includes four principal branches, the Old Scan-
dinavian (OS, Z287/Z288, which arose 3700 ± 400 ybp),
Young Scandinavian (YS, L448, which arose 2700 ± 350 ybp),
the Scottish Highland (SH, L176.1, which arose 1850 ± 290
ybp), and the parent branch, which is negative to the known
downstream SNPs (which arose 4300 ± 500 ybp).
Additionally, the R1a nomenclature also includes the L175
subclade as downstream of L176.1, however, it is most proba-
bly a private SNP. The haplotype tree is shown in Figure 8.
The lower part of the tree is represented by the Old Scandi-
navian branch with its base haplotype as follows:
13 25 16 10 11 14 12 12 10 13 11 30—15 9 10 11 11 24 14
20 31 12 15 15 16—11 12 19 23 16 16 18 19 35 39 12 11—12
8 17 17 8 12 10 8 11 10 12 22 22 16 11 12 12 13 8 14 23 21 12
12 11 13 11 11 12 13
It differs by 8 mutations from
800 years), which places their common ancestor at ≥5200
years (see the explanation above regarding the “equal or higher”
sign).
Since there are 17 identical (base) 12-marker haplotypes
150

I. L. ROZHANSKII, A. A. KLYOSOV
Map 8.
The map of the M458 subclade, depicting birthplaces of the furthermost ancestors recorded in available da-
tabases. Blue pins denote the Central European branch (CE-1 deep-blue, CE-2 light-blue), green pins corre-
spond to the West Slavic branch.
Figure 8.
A 67-marker haplotype tree of the R1a-Z284 (Scandinavian) subclade,
composed of 540 haplotypes. Three principal subclades/branches are
shown: the Old Scandinavian (Z284), the Young Scandinavian (L448),
and the Scottish Highland (L176.1).
among 178 OS haplotypes in the dataset, the logarithmic me-
thod gives [ln(178/17)]/.12 = 117 → 133 generations, or ~3325
ye
15 9 10 11 11 23 14
20 32 12 15 15 16—11 12 19 21 16 16 17 18 34 38 12 11—11
8
12 11 13
11—12
8
s to their common ancestor. The com-
pa
ars to the common ancestor; this is within the above 3500 ±
400 ybp, as determined by the linear method. The good fit
(within 5%) means that practically all 178 OS haplotypes actu-
ally descended from one common ancestor (Klyosov, 2009a).
The Young Scandinavian branch (L448+) has the following
base haplotype:
13 25 15 11 11 14 12 12 10 14 11 31—
17 17 8 12 10 8 11 10 12 22 22 15 11 12 12 13 8 14 23 21 12
11 11 12 12
Since the YS base haplotype is younger than the OS base, it
is more distant from the Z280 base haplotype on the same hap-
logroup tree; indeed, they deviate by 12 mutations (2775 years);
this places their common ancestor at ≥5300 ybp, which is prac-
tically identical with the date obtained above with the OS base
haplotype.
The same figure is reproduced with a rather loose set of hap-
lotypes, positive at Z284 and negative at L448 and Z287/Z288
(4200 ± 500 ybp), with the base haplotype as follows:
13 25 15 10 11 14 12 12 10 13 11 30—15 9 10 11 11 24 14
20 32 12 15 15 16—11 11 19 23 16 16 17 18 35 39 12
17 17 8 12 10 8 11 10 12 22 22 15 11 12 12 13 8 14 23 21 12
12 11 13 11 11 12 13
Its 10-mutation difference (2275 years) with the base Z280
results in ≥5700 year
rison of the above base haplotype with those for OS and YS
suggests that it is identical or nearly identical to the base hap-
lotype of the entire Z284 subclade, with its estimated “age” of

I. L. ROZHANSKII, A. A. KLYOSOV
~4300 years.
13 25 16 11 11 14 12 12 10 14 11 31—15 8 10 11 11 23 14
20 29 12 15 15 15—10 12 19 21 16 16 17 19 34 38 12 11—11
8 17 17 8 12 10 8 11 10 12 22 22 15 11 12 12 13 8 14 23 21 12
12 11 13 11 11 12 12
Seven mutations (1550 years) with the YS base haplotype
confirms their relative position of the subclade tree.
The known sub-branches of Z284 were found mostly in
Scandinavian countries and on the Isles (Map 9). Norwegians
and Swedish-speaking Finns seem to be the typical bearers of
this genealogical DNA-lineage.
Chronology of the Main M417+ Subclades
The combined data of SNP mutations and extended STR
haplotypes allowed us to make a step-wise reconstruction of the
base haplotypes of the major subclades of the R1a1 haplogroup.
A consensus tree was calculated from these data (Figure 9). Its
topology follows closely that of SNP phylogeny, while the
dates of its principal nodes are quite robust, as it is shown by
the examples above.
Conclusion and Some Historical Considerations
It follows from the data presented in this study, that all three
principal downstream R1a subclades, L664, Z283 and Z93,
split from their common European ancestor at about the same
time, around 7000 - 6000 ybp, and that all three went through a
population bottleneck, and then expanded 1000 - 1500 years
L664 apparently stayed in North-Western Europe; its
lineage recovered and began expanding ~4575 ybp.
The Z280 subclade (descendants of Z283) moved east to the
Russian Plain ~4800 - 4600 ybp when the Arbins, bearers of
haplogroup R1b, were arriving in Europe from different direc-
tions—from North Africa to the Pyrenees (4800 ybp); then—as
the Bell Beakers—to the continent; and finally from Asia Mi-
nor and the Middle East to the Apennines and the Balkans
(4800 - 4500 ybp). The Z280 subclade and its Russian Plain
branch formed at least 16 sub-branches from the middle of the
3rd millennium BC and later in the course of the westward re-
population of Europe in the 1st millennium BC-1st millennium
CE.
The Z93 subclade began to expand during the Aryan migra-
tions, on their way to India and the Middle East in the 3rd-2nd
millennia BC. The eastward migration of the Aryans continued
to the east (the Ural Mountains and beyond, to the Altay), south
east (Iran and Hindustan), south (Anatolia, Mitanni and further
south to the Middle East) between 4500 and 3500 ybp. The
Jewish R1a common ancestor lived 4525 ± 400 ybp; his de-
scendants also almost vanished, but their long population bot-
tleneck ended around the 8th century CE, in the Diaspora, ap-
parently in Germany.
During the 1st millennium BC bearers of R1a started to re-
populate Europe west of the Carpathian Mountains. This led to
the appearance of the Northern European branch (L365, 2600
ybp), Western Carpathian (P278.2, 2600 ybp), Eastern Carpa-
thian (2600 ybp, SNP is unknown), Northernpathian (2150
ybp, SNP is unknown), two Balto-Carpathian branches (2550
The haplotype of the Scottish Highland branch (L176.1) is
later.
Car
Map 9.
The map of the Scandinavian Z284 subclade, depicting
available databases. Magenta pins denote the parent br
ond
birthplaces of the furthermost ancestors recorded in
anch, green pins mark the Old Scandinavian one
to the young Scandinavi(Z287/Z288), deep-blue and light-blue pins corresp
Scottish Highland sub-branch (L176.1), respectively.
an branch (L448) and its
152

I. L. ROZHANSKII, A. A. KLYOSOV
extended haplotypes from available databases.
f present-day bearers of particular branches. Big
k the assignments by SNP, wherever possible.
in the text.
Figure 9.
a on
A o
mar
as
estern Eurasian-1 (2300 ybp). The same sequence of popula-
tion events can be traced in two subclades which are down-
stream from Z283: M458 (European) and Z284 (Scandinavian).
The former arose 4200 ybp in a yet unknown region, and ex-
panded as two Central European branches of subclade M458
(both arose 2900 ybp) and the Western Slavic branch (L260,
2700 ybp). Z284 arose probably in 4300 ybp, somewhere on
the Scandinavian peninsula, and produced three principal Scan-
dinavian subclades—Old Scandinavian (Z287, 3700 ybp),
Young Scandinavian (L448, 2700 ybp), and Scottish Highland
(L176.1, 1850 ybp).
Some of the older branches, e.g. the Russian Plain branch,
stayed in the present Russia-Ukraine-Belarus-Poland-Baltic
region; ancient historians described these peoples as Scythians,
Antes, Veneti, and various other proto-Slavic tribes (though
many of them belonged to other haplogroups, primarily I1 and
I2). R1a branches which are “older” than 3000 years, such as
the Russian Plain branch (4600 ybp), the Western Eurasian
branch (4300 ybp), the Balto-Carpathian (4300 ybp), did not
move en mass to Europe and stayed behind at the Russian
Plain.
In the middle of the 1st millennium CE, at the times of the
collapse of the Roman Empire, many populations, including
R1a, were migrating westward and eastward; these migrations
gradually formed the current landscape of R1a in Europe. All
38 branches of R1a and their datings are listed in the Appendix
of this paper; current distribution maps are shown in the body
of this paper.
Presently, the largest R1a populations live in Russia (up to
63% of the total population in the South of historical Russia), in
Poland, Ukraine and Belarus (up to 57% of the population).
Outside of Eastern and Central Europe, R1a accounts for to
15%-20% of the population of India (some estimates place it up
to 30% in North India), up to 9% of the population in the Mid-
dle East belongs to R1a (Aby-Amero et al., 2009); in the rest of
the world, the presence of R1a is minor (Klyosov & Rozhanskii,
2012, and references therein).
Materials and Methods
Two thousand four hundred and seventy-one (2471) of 67-
and 111-marker R1a1 haplotypes were considered in this study.
Together, they constitute the IRAKAZ R1a database (Rozhan-
skii et al., 2012), which was collected from FTDNA, YSearch,
and private lists, kindly provided for public usage by Martin
Voorwinden, Andrew MacEachern and others, to whom we are
deeply indebted.
The methodology of haplotype dataset analysis was des-
cribed in Rozhanskii and Klyosov (2011). In this study the
linear and the logarithmic method were used, the latter when
mutation counting was not required, as described in Klyosov
(2009a). The mutation rate constants are listed in Klyosov
(2009a); Rozhanskii and Klyosov (2011), and for a number of
cases are given in the text of this paper.
The concept of a “conditional generation” is an important
part of DNA genealogy, and it is not related to a common
“generation”. This is because a generation length is a “floating”
figure which depends on a society, culture, traditions, and par-
ticular circumstances (wars, famine, cataclysms, epidemics,
The R1a1 haplogroup tree according to the dat
The lengths of colored bars correspond to TMRC
circles denote the principal nodes. Red circles
Designation of branches follows the same notation
and 2200 ybp, SNPs are unknown), two Northern Eurasian
branches of Z92 subclade (3600 and 2350 ybp; the Z92 sub-
lade itself arose 4450 ybp, apparently on the Russian Plain), c
W

I. L. ROZHANSKII, A. A. KLYOSOV
etc.). It means thions of “generations” into years
should be done b
The calibration wa
in the preceding p
skii & Klyosov, 2011).
As it follows from basic rules of chemical kinetics, the ex-
perimentally determined ratio n/N in the equation
n/N = kt
ultimately determines a product kt.
In this equation:
n = a number of mutations (from the base haplotype) in a se-
ries of N haplotypes,
N = a number of haplotypes, descending from one common
ancestor (having the base haplotype) in the series,
k = the average mutation rate constant for the haplotypes (per
generation) of the given format,
t = a number of generations (TMRCA).
Therefore, if we set the length of the generation, for example
as 25 years (making it “the “conditional generation”, which is a
mathematical value rather that a floating “generation”), then the
value of k can be calibrated with respect to a certain timespan
during which n mutations occurred per N haplotypes. For ex-
ample, in 84 of 67 marker haplotypes of the Clan Donald (the
common ancestor of which, John, Lord of the Isles, died in
1386, with the TMRCA equaling to 650 years before present,
which is 26 “conditional generations” of 25 years each), there
were 262 mutations. This gives 262/84 = 26 k, and hence k
= .12 mutations per the conditional generation (25 years) per
the 67 marker haplotype.
As it is shown inii & Klyosov, 2011), the ob-
tained mutation rate
ducible for many ser
haplogroups from ma
ct, it is reproducib
lective number of mutations in the dataset having one common
ogy is the loga-
ation counting in
haplotypes. It is based on the assumption (which largely or
practically always holds true) that a transition of the base hap-
lotypes into mutated ones (or, rather, belonging to descendants
inherited mutated haplotypes) is described by the first-order
kinetics:
N = Aekt
that is
ln(N/A) = kt
where:
N = a total number of haplotypes in a set,
A = a number of unchanged (identical, not mutated) base
haplotypes in the set,
k = an average mutation rate,
t = a number of generations to a common ancestor.
One can see the logarithmic method does not consider a
number of mutations in haplotypes; only mutated and non-
mutated (base) haplotypes are considered.
The above equation is the fundamental one in chemical ki-
netics. Radioactive decay, for example, is described by the
same equation, though it does not involve an analogy with
“back mutations”, since it is truly irreversible. Back mutations
(or reverse mutations), though, are quite common in haplotypes,
and their analysis and corrections for back mutations are given
in (Klyosov, 2009a; Rozhanskii & Klyosov, 2011).
Haplotype trees were composed using HYLIP, the Phylog-
eny Inference Package program (Felsenstein, J., 2004; for mul-
ee Klyosov, 2009a,
for back mutations
2009a); Rozhanskii
011). Margins of error were calculated as de-
scribed in Klyosov (2009a).
at convers
y calibrating using known historical events.
s performed on many examples, as described
aper in Advances in Anthropology (Rozhan-
ancestor.
Another principal method in DNA geneal
rithmic method, which does not involve mut
(Rozhansk
constant of .12 is rather accurately repro-
ies of 67 marker haplotypes from many
ny tested populations around the world. In
le for ANY tested population in the world
tiple examples of usage and calculations s
2009b and references therein). Corrections
were introduced as described in Klyosov (
and Klyosov (2
fa
for which 67 marker haplotypes are available (Rozhanskii &
Klyosov, 2011). This “calibration” connects a number of muta-
tions in a series of haplotypes (in a given format and length,
from the 6 marker haplotypes to the 111 marker haplotypes) to
a known number of years to a known historical event or a
known genealogy, setting a length of generation as 25 years,
and verifying the mutation rate constant related to this “condi-
tional generation”.
It should be noted that we could have set the timespan to the
common ancestor of the Clan Donald as 13 generations of 50
years each and it would not have changed a thing. There would
still be 650 years to the common ancestor. The mutation rate
constant would simply be twice as high.
The mutation rate constant for the 111-marker haplotypes,
also employed in this study, equals to .198 mutation per haplo-
type per conditional generation (Rozhanskii & Klyosov, 2011).
The above equation is essentially the basis for the “linear”
method in DNA genealogy. It is based on mutation counting in
haplotypes in the dataset compared with the “base” haplotype,
which is (or the best available approximation to) the ancestral
haplotype for the dataset. The “base” haplotype is the ancestral
haplotype in an ideal case. However, since those base haplo-
types often are deduced ones, it would be inaccurate to call
them “ancestral” when that might not necessarily be true.
Hence, we call them “base” haplotypes. Base haplotypes in the
dataset were determined by minimization of mutations; by de-
finition, the base haplotype is one which has the minimum col-
Acknowledgements
The authors are indebted to Dr. Judith Remy Leder for her
va
P
luable help with the preparation of the manuscript.
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I. L. ROZHANSKII, A. A. KLYOSOV
Appendix 1
list of R1a branches in Europe (dates in ybp). A
1 R1a1-M198− (or SRY10831.2*) ≥9000
2 R1a1a-M198+ M417− (or M17*) ≥9000
3 The Tenths, NW branch, ~4575
s, DYS448 null mutation
1125 ± 200
3700 ± 400
27 (WC) 2600 ± 300
28 Z280-Z92, North Eurasian (NEA) 4450 ± 450
29 Z280, NEA-1 3600 ± 400
30 Z280, NEA-2 2350 ± 300
31 Z280, Western Eurasian (WEA) 4100 ± 500
32 Z280, WEA-1 2300 ± 300
33 Z280, Eastern Carpathian (EC) 2600 ± 300
34 Z280, Balto-Carpathian (BC) 4300 ± 500
35 Z280, BC-1 2550 ± 300
36 Z280, BC-2 2200 ± 250
37 Z280, Northern Carpathian (NC) 2150 ± 300
38 Z280, L784 2500 ± 450
4 The Tenths, L664+ ~4575
5 The Tenth1500 ± 325 and 260 ± 60
(USA)
6 Z93+ ~5700,
started to expand ~4050
7 Z93-L342.2+ 4900 ± 500
8 Z93-L342.2-L657+ 4050 ± 460
9 Z93-L342.2, Jewish branch 4475 ± 400,
started to expand 1300 ± 150
10 Z93-L342.2, Bashkir branch
11 Z93-L342.2, Kyrgyz branch 2100 ± 250
12 Z283 (Eurasian branch) ~5500
13 M458 (European branch) ~4200
14 M458 CE parent ~3100
15 M458 CE-1 2900 ± 400
16 M458 CE-2 2900 ± 400
17 M458-L260 WS 2700 ± 300
18 Z284 (Scandinavian branch) 4300 ± 500
19 Z284-Z287, Old Scandinavian
20 Z284-L448, Young Scandinavian 2700 ± 350
21 Z284-L448-L176.1,
Scottish Highland 1850 ± 290
22 Z280, Central Eurasian (CEA) 4900 ± 500
23 Z280, CEA-1, Russian Plain 4600 ± 500
24 Z280, CEA-2 3500 ± 400
25 Z280-L366, CEA-3 2500 ± 350
26 Z280-L365, North European (NE) 2600 ± 350
Z280-P278.2, Western Carpathian
Appendix 2
lo
as
h
io s
N
h
o
h
h
io
h
h
olts
u
h
ct
p
The following DNA projects were selected as primary hap-
type databases:
http://www.familytreedna.com/public/R1aY-Haploroup/default.
px?vgroup=R1aY-Haplogroup&section=yresults
http://www.familytreedna.com/public/R1a/default.aspx? section
=yresults
ttp://www.familytreedna.com/public/r-arabi a/default.aspx?sect
n=yresult
http://www.familytreedna.com/public/Hungarian_Magyar_Y-D
A_Project/default.aspx?section=yresults
ttp://www.familytreedna.com/public/India/default.aspx?sectio
n=yresults
http://www.familytreedna.com/public/Turkic/default.aspx? secti
n=yresults
ttp://www.familytreedna.com/publicwebsite.aspx?vgroup=ger
many&section=yresults
ttp://www.familytreedna.com/public/Finland/default.aspx?sect
n=yresults
http://www.familytreedna.com/public/Czech/default.aspx?secti
on=yresults
ttp://www.familytreedna.com/public/slovakia/default.aspx?sec
tion=yresults
ttp://www.familytreedna.com/public/polish/default.aspx? secti
n=yresu
http://www.familytreedna.com/public/russian_impire.aspx/defa
lt.aspx?section=yresults
ttp://www.familytreedna.com/public/russiadna/default.aspx? se
ion=yresults
Reference data were selected according to SNP assignment
from YSearch database: (http://www.ysearch.org) and public
rojects of FTDNA (http://www.familytreedna.com).
156