Advances in Anthropology
2012. Vol.2, No.4, 198-213
Published Online November 2012 in SciRes (http://www.SciRP.org/journal/aa) http://dx.doi.org/10.4236/aa.2012.24022
198
Re-Examining the Out-of-Africa Theory and the
Origin of Europeoids (Caucasoids).
Part 2. SNPs, Haplogroups and Haplotypes
in the Y-Chromosome of Chimpanzee and Humans
Anatole A. Klyosov, Igor L. Rozhanskii, Lyudmila E. Ryabchenko
The Academy of DNA Genealogy, Newton, USA
Received July 12th, 2012; revised August 22nd, 2012; accepted September 12th, 2012
Our consideration of human haplogroups, and our analysis of the dynamics of the Y-chromosome nucleo-
tide flow from primates to humans during the evolution of genus Homo has shown that a common ances-
tor of the majority of present day human males, both African and non-African, lived approximately
160,000 years ago. The haplogroup of this common ancestor has been identified as the α-haplogroup,
which is equivalent or close to haplogroups A1/A1b in the current phylogeny. The archaic lineages (cur-
rently summarily designated A0) descend from an ancestor who lived no later than 180,000 years ago,
and probably much earlier. The α-haplogroup and the A0 lineages have significantly different nucleotide
patterns, and they certainly did not descend one from another. Furthermore, our research points up the ar-
eas of mutations in Y-chromosome in H. sapiens, which allows us to use chimpanzee MSY (the male-
specific region of the Y-chromosome) as a proxy for genus Homo’s common α-haplogroup ancestor.
When we studied slow mutating 16-marker haplotypes, we discovered that chimpanzees and present day
humans had a common ancestor 5.5 ± 0.9 million years before the present. It is clear that, when they are
compared to loci in other primates, such as gorillas, orangutans, and macaques, many human
Y-chromosome loci have been conserved from our common ancestor. Results of our analysis of haplo-
types, conserved (ancestral) nucleotides, and SNPs suggest that there is no reason to believe that ancestors
of non-Africans (β-haplogroup, i.e. haplogroup BT and its downstream haplogroups) descended from
haplogroups A0, A1a, or any other African haplogroup. The data are adequately described by a model
which shows that the African lineages and non-African lineages diverged from the α-haplogroup ap-
proximately 160,000 years before the present and that the Y-chromosomes of the two groups have
evolved independently (in terms of Y-chromosome) since then. We have no indication of where the
common ancestor of the α-haplogroup lived; he could just as easily have lived in Europe, in Asia, or in
the Middle East, as in (less likely) Africa. We believe that all the presuppositions posited in support of the
Out-of-Africa hypothesis fail to hold up under simple scrutiny. This study shows that the Out-of-Africa
hypothesis has not been adequately substantiated. The common assertion that “anatomically modern hu-
mans came out of Africa some 70,000 years ago” has never been convincingly calculated or determined
otherwise; our research suggests that it is incorrect.
Keywords: Y-Chromosome; Mutations; Haplotypes; Haplogroups; Primates; Chimpanzees, SNP;
Out-of-Africa
Introduction
In the first part of our two-part study (Re-examining the “Out
of Africa” theory and the origin of Europeoids (Caucasoids) in
light of DNA genealogy [Klyosov & Rozhanskii, 2012a]) we
considered the origin of anatomically modern humans (AMH),
who presumably belong to Y-chromosomal haplogroups A
through T according to the classification developed in human
genetics and in the DNA phylogeny of man. Our first article
also set forth a time frame for the origin of haplogroups A and
B—the first (partly) African and the second the origin of seem-
ingly non-African, which later migrated to Africa.
In our earlier paper we identified the relative position of the
haplogroups known today, and suggested a re-examination the
validity of the Out-of-Africa hypothesis. Looking at slow
changing 22-marker haplotypes, we discovered that African
haplogroup A originated 132,000 ± 12,000 years before the
present, and is remote from all other haplogroups, which origin-
nated (or, rather, came through a population bottleneck) about
64,000 ± 6000 ybp. These other haplogroups—BT (names BT
and BR are both used in the nomenclature and are considered
practically equivalent in general context) and those downstream
from it--include the Europeoid (Caucasoid) haplogroups from F
through T that originated 58,000 ± 5000 ybp. About 160,000 ±
12,000 ybp haplogroups A1, A1a, A1b (in the current classify-
cation) and BT had a common ancestor in the α-haplogroup.
We showed that BT did not descend from haplogroup A; but
rather that the lineages were two sides of a bifurcation in the α-
haplogroup. The two lineages evolved distinctly with respect to
the Y-chromosome.

A. A. KLYOSOV ET AL.
The Essence of the Out-of-Africa Hypothesis
The Out-of-Africa hypothesis arose in the middle of the
1980s, under heavy influence of DNA-related interpretations.
One of the first articles (Stringer & Andrews, 1988), which
initiated the idea, stated that “Genetic data on present human
population relationships and data from the Pleistocene fossil
hominid record are used to compare two contrasting models for
the origin of modern humans”. We now know that early genetic
data were inaccurate and that the hominid fossil record was
(and still is) controversial. In the same year, by studying 42
world populations Cavalli-Sforza et al. (1988) found that the
phylogenetic tree separates Africans from non-Africans. They
took this as an indication that “the origin of modern humans
was in Africa”, even though the opposite, that is “the origin of
modern humans was not in Africa” would have been equally
valid. Later, the Out-of-Africa hypothesis acquired five essen-
tial presuppositions:
1) All haplogroups, from B to T, descended from haplogroup
A.
2) The first anatomically modern human (AMH) came Out-
of-Africa around 70,000 years ago.
3) Africa has the highest genetic diversity. Therefore, the
AMH descended from African populations.
4) Genetic diversity decreases as the distance from Africa
increases. Therefore, Africa is the homeland of mankind.
5) The earliest AMH was found in Africa. He presumably is
the direct ancestor of all the human beings living today.
We concluded in the first article of this set (Klyosov & Roz-
hanskii, 2012a) that the Out-of-Africa hypothesis needed
re-examination. The mutations in the Y-chromosome haplo-
types of Africans and non-Africans clearly indicate that
1) non-Africans could not have descend from Africans;
2) haplogroup B which is about 120,000 years remote from
haplogroup A, is not-initially-an African haplogroup;
3) African haplogroup A and all non-African haplogroups
(BT) diverged from a common α-haplogroup ancestor who
lived 160,000 ± 12,000 ybp.
There is no indication that non-African haplogroups resided
in Africa. Furthermore, it was recently recognized that there is
no such thing as a single haplogroup A. Instead, haplogroup A
is an umbrella for a variety of haplogroups, some of which
originated more than 160,000 ybp.
The Five Presuppositions of the
Out-of-Africa Hypothesis
Let us briefly consider the five presuppositions listed above,
which are the underpinnings of the Out-of-Africa hypothesis.
Presupposition 1 was maintained for many years by the In-
ternational Society of Genetic Genealogy (ISOGG) with the
following description: “The BR haplogroup split off from hap-
logroup A 55,000 years before present (bp). It probably ap-
peared in North East Africa” Similarly, the introduction to the
FTDNA haplogroup A Project (see “Materials and Methods”)
begins “Haplogroup A is unique in that all other human hap-
logroups spring from its primal branches”. Another similar
quotation—“Haplogroup A, first appearing 55,000 years ago,
is the oldest of all Y haplogroups and is considered a direct ge-
netic link to early man” (Chromosomal Laboratories, 2005).
Since we know that the common ancestor of haplogroup A and
all other haplogroups belonged initially to the α-haplogroup
(which arose 160,000 ybp), and haplogroup B is remote from
haplogroup A by at least 120,000 years, it is clear that hap-
logroup BT (which arose 64,000 ybp) could not possibly have
“split off from haplogroup A 55,000 ybp”. This chronological
construction provides inadequate support to the Out-of-Africa
hypothesis.
Even if some researchers did not consider haplogroup
A-M91 to be ancestral to the BR haplogroup (e.g. Cruciani et
al., 2002), the literature abounds in interpretations which create
an impression that African haplogroups were ancestral to
non-African haplogroups. For example, Chiaroni, Underhill and
Cavalli-Sforza (2009) state: “haplogroups A and B are the
deepest branches in the phylogeny and are essentially restricted
to Africa, bolstering the evidence that modern humans first
arose there”. Actually, haplogroup B is not among the deepest
branches, (cf. Figure 1). Second, Chiaroni, et al., do not indi-
cate from which particular haplogroup “modern humans first
arose [in Africa]”. From haplogroup A? No, haplogroup A (see
Figure 1) is distinct from BT, as Cruciani, et al., pointed out in
2002. From haplogroup B? No, BT was initially one hap-
logroup and haplogroup B split from it. From some chromoso-
mal Adam 160,000 years ago evolving directly into non-Afri-
cans? This cannot be serious. It seems that such Out-of-Africa
statements have not been thought through in terms of Y-chro-
mosome phylogeny.
It is interesting to note that SNP M91 long considered an Af-
rican SNP, has recently been re-assigned by ISOGG to
non-African haplogroups (ISOGG-2012) (cf. Cruciani et al.,
2002; ISOGG, 2006-2011). This move may have been an at-
tempt to keep the nomenclature in line with the Out-of-Africa
hypothesis. However, the respective nucleotide sequences in
chimpanzee MSY and the BT haplogroup are identical (9T),
while the African subclade A1a has 8T; therefore, M91 should
belong to the African lineages. On the other hand, M91 is 8T in
A1b, which is a part of the α-haplogroup. This means, that after
its transition from the α-haplogroup to the β-haplogroup (BT)
8T9T, the sequence returned to its ancestral state, and the
SNP in BT should be designated differently than M91. All of
these data indicate that BT haplogroups did not descend from
the “African haplogroup A”; additionally, the slow 22-marker
haplotypes quite convincingly show this (Klyosov & Roz-
hanskii, 2012a). More genome-based data will be provided
below in this paper.
Presupposition 2 became a cliché in almost every article in
which the Out-of-Africa hypothesis has been mentioned. A few
examples, specifying when AMH allegedly came Out-of-Africa,
are:
 “50 thousand years ago” (Jobling & Tyler-Smith, 2003)
 “50 - 60 thousand years ago” (Shi et al., 2010; Mellars,
2011)
 “50 - 70 thousand years ago” (Hydjasov et al., 2007;
Stoneking & Delfin, 2010)
 “60 thousand years ago” (Li & Durbin, 2011; Henn et al.,
2011)
 “60 - 70 thousand years” (Ottoni et al., 2010)
 “60 thousand years ago” (Stewart & Stringer, 2012)
 “45 - 50 thousand years ago” (Fernandes et al., 2012)
 “50 - 65 thousand years before present” (Behar et al., 2008)
 “60 thousand years ago” (Chiaroni et al., 2009)
 “50 - 75 thousand years ago” (Patin et al., 2009)
 “50 thousand years ago” (Edmonds et al., 2004)
 “45 thousand years ago” (Moorjani et al., 2011)
 “50 - 70 thousand years ago” (Xue et al., 2005)

“70 - 80 thousand years ago” (Majumder, 2010)

A. A. KLYOSOV ET AL.
200
Figure 1.
Haplogroup tree of the H. sapiens Y-chromosome derived from haplotypes and subclades. This diagram
shows the most recent common ancestors (TMRCAs) of H. sapiens as described in this study. To prepare this
tree, we analyzed 7415 haplotypes from 46 subclades of 17 major haplogroups. The timescale on the vertical
axis shows thousands of years from the common ancestors of the haplogroups and subclades. The tree shows
the α-haplogroup, which is ancestral to both the African and non-African haplogroups, and the β-haplogroup,
which is equivalent to haplogroup BT in the current classification. The left branch haplogroup A arose
~132,000 ybp. Haplogroups F through R (which includes T) represent Europeoids (Caucasoids) who arose
~58,000 years before the present. The Mongoloid and Austronesian haplogroup C split ~36,000 ybp; the Mid-
dle Eastern haplogroups D and E split ~42,000 ybp (Klyosov and Rozhanskii, 2012a).
logroup A or B. So, from whom might they have descended in
Africa, and when?
 “40 - 80 thousand years ago” (Campbell and Tishkoff,
2010)
Presupposition 3 is phrased variously: Hellenthal et al.
(2008) say, “the haplotype diversity is highest in Africans”,
(Campbell & Tishkoff, 2010), say that “Africa not only has the
highest levels of human genetic variation in the world but also
contains a considerable amount of linguistic, environmental
and cultural diversity”. In reality, highest diversity does not by
any means represent a homeland, unless the homeland has been
totally isolated and has not been affected by populations from
outside. New York City has a higher diversity than Boston, but
that doesn’t mean that New York City is a homeland for Boston.
Diversity quite commonly is a result of mixing, and Africa is a
home of mixed populations, including quite ancient populations
known presently as A0, A1, etc., plus haplogroup B, which did
not descend from haplogroup A. When it arrived in Africa, it
greatly increased the diversity there. The same is with a popula-
tion of R1b-V88, which arrived in Africa on its migratory way
along the Mediterranean Sea (Cruciani et al., 2010) and cur-
rently resides in Cameroon and Chad, also contributing to di-
versity in Africa. As it is shown below, lineages A0 do not only
predate the α-haplogroup (A1/A1b), but evolved (in terms of
Y-chromosome) independently on the latter. It is likely that A0
represent the African lineages, while the α-haplogroup with its
downstream haplogroups represent the non-African ones.
 “55 - 70 thousand years ago” (Soares et al., 2009)
 “between 40 and 70 thousand years ago” (Sahoo et al.,
2006)
 “between 35 and 89 thousand years ago” (Underhill et al.,
2000)
 “between 80 and 50 thousand years ago” (Yotova et al.,
2011)
 “between 50 and 100 thousand years ago” (Hublin, 2011)
 “between 27 - 53 and 58 - 112 thousand years ago” (Carri-
gan and Hammer, 2006)
 “around 70 - 60 thousand years ago” (Curnoe et al., 2012).
In fact, however, none of these authors seem actually to have
calculated or otherwise obtained the numbers that they cite.
Besides, at this point, there is no methodology which can cal-
culate those numbers. The numbers are taken from thin air, and
cross-cited in dozens of articles. It is astonishing to us that no
scholar has ever questioned the figures. The Out-of-Africa hy-
pothesis needed the figures, and they seem to have been pro-
vided without justification.
In fact, neither “haplogroup A” (there are now a number of
them), nor any other haplogroup could possibly have come out
of Africa 70,000 years ago, since the temporal distance between
A and all other haplogroups is at least 120,000 years.
Non-African haplogroups clearly did not descend from hap-
In short, Africa is home to truly ancient (compared to BT)
populations of various haplogroups designated presently as A,

A. A. KLYOSOV ET AL.
and other haplogroups which arrived in Africa at different times.
So, diversity by itself does not support the idea of a common
homeland in this context.
Presupposition 4 is stated clearly by Atkinson (2011) in a
recent article in which he asserts that “human genetic and phe-
notypic diversity declines with distance from Africa ··· under-
pinning support for an African origin of modern humans ··· loca l
language diversity ··· points to parallel mechanisms shaping
genetic and linguistic diversity and supports an African origin
of modern human languages”. This effort to shore up the
Out-of-Africa hypothesis also fails. Since haplogroups A and
BT diverged from the same α-haplogroup (Figure 1), this does
not mean that the younger lineage descended from the older.
Our research shows that they parallel each other such as two
teeth in the proverbial fork. They descend not from each other,
but from a common ancestor who was the α-haplogroup.
Haplogroup R1a (20,000 ybp) and haplogroup R1b (16,000
ybp) (Klyosov & Rozhanskii, 2012b; Klyosov, 2012), arose in
Central Asia (ibid.). Both are younger than haplogroup A
(120,000 ybp) but they are not descended from haplogroup A.
A boy is not a descendant of his older brother. The fact that the
Indo-European (IE) languages are younger than the click lan-
guages does not prove that IE languages descended from click
languages or that Africa is a homeland of the languages of
man.
Presupposition 5, that the earliest AMH was found in Africa,
is unconvincing for the following reasons:
1) We do not have the DNA of most fossils. Without identi-
fying the haplotypes of fossil DNA, any statement about the
relationship of fossils to present-day H. sapiens is irrelevant.
2) The oldest AMH skeletal fragments found in Africa carry
many archaic features. They were not exactly AMH, and many
clearly do not have modern H. sapiens morphology. Many
skeletal fragments are poorly dated. None of the early fossils
meet the stringent morphological criteria applicable to living H.
sapiens (Tattersall, 2009). A number of authors have argued
that the earliest hominids found in Sub-Saharan Africa had
African regional morphological features (e.g. Habgood, 1989).
Subsequent papers gradually began to claim that sub-Saharan
hominids had AMH features. This issue is controversial and
complicated; it is the subject of many articles (e.g. Rogers &
Jorde, 1995; Hanihara et al., 2003; Tattersall, 2009; Pinhasi et
al., 2011; Rightmire, 2009; Prat et al., 2011; Mellars & French,
2011) and by no means solved.
3) The AMH could have migrated to Africa from elsewhere.
If our ancestors could have migrated out of Africa, they could
equally well have migrated into Africa.
4) Few modern human skulls older than 20,000 years have
been discovered in Africa south of Ethiopia. One of the oldest
is from Hofmeyr, South Africa, dated to 36 ± 3 thousand years
ago. Still, it displays some archaic features (Grine et al., 2007,
2010), although it is the age or even slightly younger than the
bones of an AMH of 38 - 45 thousand ybp (e.g. Pinhasi et al.,
2011; Prat et al., 2011; Hoffecker, 2011; Benazzi et al., 2011;
Higham et al., 2011). Actually, the Hofmeyr skull demonstrates
that humans in Africa 36,000 years ago resembled those in
Eurasia, which leaves the question of who migrated where
unsolved.
While the Out-of-Africa hypothesis continues to be presented
as truth, it is based on questionable presumptions. It should be
noted that the earliest authors were careful in their descriptions
of the origin of Africans and non-Africans; they did not claim
that one group descended from the other. For example, Nei and
Takezaki (1996) wrote: “[Our] results indicate that Africans
are the first group of people that split from the rest of human
populations”, and “the root of the human population tree still
remains unresolved”.
The above is only a brief account of the controversies in the
anthropology of man and about the alleged African origin of H.
sapiens. In this study, we focus mainly on the DNA evidence
propounded to support the Out-of-Africa hypothesis. We have
found that none of the evidence is convincing. As the principal
methodology, we consider base (ancestral) haplotypes of hap-
logroups A0, A, and B, determine timespans to their common
ancestors as it was initiated in the preceding paper (Klyosov &
Rozhanskii, 2012a), compare their haplotypes with that of
chimpanzee MSY and show that the methodology of DNA
genealogy works even on timespans of millions of years, com-
pare ancestral nucleotides of Y-chromosomes of chimpanzee
(and the DNA fragments of some other primates) to those in
humans, based of that comparison differentiate lineages of
haplogroups A0, A/A1b, A1a and their downstream subclades,
assign the nucleotides to the α- and β-haplogroups, and finally
show that the African and non-African lineages indeed split
apart from the α-haplogroup; therefore, non-African lineages
did not descend from the African ones. Both African and
non-African lineages have evolved-in terms of their Y-chro-
mosomes—concurrently (in parallel) from their common an-
cestor who lived around 160,000 years before the present. One
more set of the African lineages, collectively called A0 hap-
logroups, has evolved independently (again, in terms of their
Y-chromosomes) on the α- and β-haplogroups as well as on
other African lineages. It seems that A0 are the most archaic
African lineages, which arose at least 180,000 ybp and possibly
much earlier.
The Current Structure of Haplogroup A
Currently, there is no unified haplogroup A. It is a loose set
of lineages in which each has its own set of SNPs and the list of
SNPs is growing. Some of them are shown below, thank to the
researchers and Institutions named in the “Materials and Meth-
ods” section.
Figure 2 shows a haplotype tree of haplogroup A. It is based
on tests collected by the FTDNA Haplogroup A Project. The
tree consists of four principal parts. The most remote branch in
the tree is the doublet of two relatives (1 and 2). They do not
have SNPs that occur in other haplogroups (ISOGG designated
their haplogroup as A0*), and have a base 22-marker haplotype
as follows
13 11 12 - 10 11 - 16 - 10 9 14 14 8 8 8 9 12 11 12 8 12 12
11 11 (A0*)
This is remote from all known haplotypes in the world; for
example, A0* has DYS438 = 16—a slow marker in which a
mutation happens about once in 45,000 years. All other hap-
logroups have DYS438 = 9-11. In other words, A0* differs
from other haplotypes by 5 to 7 mutations. This newly design-
nated haplogroup A0* differs from the deduced ancestral hap-
lotype of the composite haplogroup A described by Klyosov
and Rozhanskii (2012a) by 27 mutations:
12 11 11 - 9 11 - 10 - 10 8 14 15 7 10 8 12 13 11 16 8 13 9
11 12 (A)
The differences translate to 250,000 years, and place their
common ancestor at 190,000 ybp. With respect to time, this

A. A. KLYOSOV ET AL.
Figure 2.
A 22 marker tree of 39 haplotypes of haplogroup A. These haplotypes
are part of the FTDNA Haplogroup A Project. The upper right branch is
haplogroup A1a (presumably M31); it consists of haplotypes from
Finland, England, Switzerland, Cape Verde. Number 3—at about 4
o’clock on the figure—is from an African-American. His DNA has
close matches in Cameroon and presumably belongs to A0, which is
positive at SNP V152 (Simms et al., 2011), Four and 5 (both Afri-
can-American) are positive for L896; the former has close matches in
Ghana. One and 2 belong to close relatives who have been assigned to
haplogroup A0*. The remaining 28 haplotypes belong to haplogroup
A1b1b2b-M13/M32.
common ancestor is older than the α-haplogroup (160,000 ±
12,000 ybp). As we will see later, A0* has a different pattern of
nucleotides than α-haplogroup, which actually might coincide
with the A1/A1b haplogroup in the current nomenclature.
The A0* haplotype shown above differs from the base hap-
logroup B haplotype described in Part 1 of this study (Klyosov
& Rozhanskii, 2012a) by 29 mutations:
11 12 11 - 11 11 - 10 - 11 8 16 16 8 10 8 12 10 11 15 8 12 11
12 11 (B)
There are 286,000 years between haplogroup A0* and hap-
logroup B. Clearly, A0* could not be ancestral to haplogroup B,
or to haplogroup BT.
The upper right branch in the tree (haplotypes 6 - 11) belongs
to haplogroup A1a; it is composed mainly of European haplo-
types, and has the following base haplotype:
12 10 11 - 7 13 - 8 - 10 8 15 17 6 10 9 12 13 11 16 8 13 11
11 12 (A1a)
It differs from the ancestral base haplotype of haplogroup A
by 14 mutations (84,300 years). Using calculations based on 25
and 67 marker haplotypes, we have determined the age of the
A1a branch of about 8500 ybp. Since the age of haplogroup A
is 132,000 ± 20,000 years, and the A1a base haplotype is 14
mutations apart, their common ancestor lived 122,000 ± 15,000
ybp. This is haplogroup A within the margin of error, hence,
A1a is a downstream subclade of haplogroup A. Generally, it is
known from the phylogeny, and the haplotypes confirm it.
Haplotypes 3, 4 and 5 on Figure 2 belong to three different
lineages of A0. They differ by 12, 7 and 11 mutations, respec-
tively, from the presumed base haplotype:
12 12 11 - 10 11 - 12 - 11 8 15 15 8 10 8 0 12 11 14 8 13 8
12 12
These differences correspond to conditional
generations, or about 52 thousand years before the present.
Another way of arriving at the most recent common ancestor
(TMRCA) is using the permutation method described in
Klyosov (2009). The permutation method does not require the
base haplotype, and it is based on the sum of the squared muta-
tion differences between each allele in all the loci (22 loci in
this particular case) divided by the square of a number of hap-
lotypes (3 haplotypes in this case) and by the mutations rate
constant (0.00027 mutations per marker per 25 years). Since the
number of squared permutations in these three haplotypes was
122, we obtain 122/22/9 = 0.616 mutations per marker, and the
common ancestor lived 25 × 0.616/0.00027 = 57,000 years be-
fore the present, which is close to the above value.
1685 2167
In the 28 haplotypes which form haplogroup A1b1b2b-
M13/M32 on the left side of the tree, there is a young Saudi
Arabia branch of four haplotypes with a base haplotype as fol-
lows:
12 11 11 - 9 11 - 10 - 10 9 12 12 7 12 8 0 13 11 16 9 14 9 11 11
(A1b1b2b)
It differs from the base haplotype of the rest of the hap-
logroup by 40 mutations in the 67 marker format, and by 3
mutations in the 22 marker format. This translates to 12,300
years between their common ancestors.
Most of the haplotypes of haplogroup A in the dataset are
represented by people who report European or Middle Eastern
backgrounds. Therefore, the premise that haplogroup A almost
exclusively resides in Africa, is not accurate, at least based on
data on hand, even if they are clearly not representative. We
simply do not know the current distribution of haplogroup A
carriers around the world, and we know even less that distribu-
tion some hundred thousand years ago.
How Old Is Haplogroup A0?
Only four A0 haplotypes are currently known in the 22
marker format (numbers 1/2, 3, 4 and 5 on the tree in Figure
2):
13 11 12 - 10 11 - 16 - 10 9 14 14 8 8 8 9 12 11 12 8 12 12
11 11 (haplotype 1/2)
12 12 14 - 11 12 - 14 - 11 8 15 15 7 10 8 0 13 13 14 8 13 8
12 11 (haplotype 3)
12 12 11 - 10 11 - 11 - 11 8 15 15 8 10 8 0 11 13 14 7 13 9
12 13 (haplotype 4)
12 13 10 - 10 11 - 10 - 11 8 15 15 8 9 8 0 10 9 14 8 12 8 11
12 (haplotype 5)
Since their ancestral haplotype is unknown, we need again to
apply the permutation method (Klyosov, 2009) to determine
how deep in time their common ancestor might have lived. The
sum of the squared mutation differences between each of them
is 310, and we arrive at 310/16/22 = .881 mutations per marker
in the four A0 haplotypes. The common ancestor of the five A0
individuals lived 0.881/0.00027 × 25 = 81,550 years before the
present. This value makes sense only if all four haplotypes
belong to the same haplogroup, but we know that they don’t,
therefore, the actual figure is >81,550 ybp.
Since the haplotype of the doublet (1/2) is quite distant
from the others, we can use another way to estimate a time span
between the contemporary haplotype 1/2 and the base haplo-
type of 3, 4 and 5. They differ by 23 mutations, which corre-
spond to 186,000 years between them. Therefore, the common
ancestor for entire A0/A0* lineages lived (186,000 + 52,000)/2
202

A. A. KLYOSOV ET AL.
≈ 120,000 years ago, again assuming that these haplotypes
belong to the same haplogroup. This means that the actual time
to their common ancestor is significantly higher than 81,550
ybp. The common ancestor of A0 and the other African haplo-
types lived more than 180,000 years before the present
(Klyosov & Rozhanskii, 2011).
Let us move to the main subject of this study, namely, what
the Y-chromosome nucleotides and SNPs show about the con-
nection between haplogroup BT and haplogroup A. As we have
shown above, haplogroup A cannot be ancestral to haplogroup
BT. The two lineages diverged independently from the α-hap-
logroup. Additionally, one of several lineages (currently de-
signated as A0 and A0*) evolved independently of the hap-
logroup A.
Nucleotides and SNPs in the Y-Chromosome of
H. Sapiens and the Y-Chromosome and/or DNA
Fragments of Some Primates
As the principal methodology of this study we consider nu-
cleotides and SNPs in
1) some living primates of the family Hominidae (e.g. chim-
panzee, gorilla, orangutan) and the macaque (of the family
Cercopithecidae, tribe Papionini)
2) newly discovered lineages in a variety of haplogroups A,
3) haplogroup BT and its descendant haplogroups R1a and
R1b (as specific examples of Europeoids [Caucasoids]).
First, we should consider whether or not we are correct when
we choose to compare nucleotides and SNPs in living primates
and living humans. We can assume that human and chimpanzee
genomes diverged 6.5 million years ago (cit. Green et al., 2010)
or 6 million years ago (Hughes et al., 2012; Scally et al., 2012),
or 5 to 7 million years ago (Prüfer et al., 2012). The tribe Papi-
onini emerged 6 - 8 million years ago and the Hominidae fam-
ily emerged 13 - 18 million years ago (Perelman et al., 2011).
Following Cruciani et al. (2011) and Scally et al. (2012), and
references therein, we can take 1.0 × 10−9 single-nucleotide
substitutions per base per year as an estimate of the mutation
rate in the Y-chromosome. This tells us that in 6.5 million years
a nucleotide in the Y-chromosome would mutate 0.013 - 0.015,
or between 1.3% and 1.5%. Whatever the exact figure, it is
clear that it is safe to use primate nucleotides as proxies for
nucleotides in present day humans. We know that chimpanzee
Y-chromosomes and human Y-chromosomes are remarkably
divergent in structure and gene content (Hughes et al., 2010);
however, nucleotides in the Y-chromosome positions we have
considered (see below) appear to have retained their structure
quite remarkably (see Figure 3).
Figure 3 shows a comparison of a 97-nucleotide human
DNA fragment with the respective MSY fragment in chimpan-
zee and the DNA fragments from the whole genome shotgus
sequence of gorilla, pongo, and macaque. Eighty seven percent
(84 of the 97 nucleotides) are identical in humans and in these
primates (see Figure 3). The fragments contain nucleotide C
(position 51) which becomes SNP P82 (mutation CA) con-
verting haplogroup A1 into A1a. Other SNPs which mark this
conversion are M31, V4, V14, and V25.
One can see that the ancestral nucleotides in the DNA of
these primates have been stable for 15 million years or so; thus,
they can be employed for comparison with the respective nu-
cleotides in human Y-chromosomes.
Since the chimpanzee genome is by many accounts the
closest to humans and the most thoroughly studied, and since
the chimpanzee genome sequence is available (see Materials
and Methods), it was employed in our further studies.
How Long Ago Did the Split between the
Common Ancestor of Chimpanzees and
Humans Occur?
To answer this question we have employed slow markers
available for the 22-marker panel (Klyosov & Rozhanskii,
2012a); the respective alleles were determined for chimpanzee
and human MSYs (see Materials and Methods). Overall, 16
markers out of the 22 were recovered in the chimpanzee
Y-chromosome, as follows: DYS 426, 388, 392, 455, 438, 578,
641, 472, 425, 594, 436, 490, 617, 568, 640, 492. Six other
markers in chimpanzee MSY did not have homology with the
human MSY as a result of a rearrangements of genome that
occurred during the past~6.5 million years. Therefore, the 16-
marker haplotype of present-day chimpanzee MSY was ana-
lyzed (see Materials and Methods) as follows:
8 15 10 4 5 9 10 5 10 4 4 7 4 4 8 9
In present-day H. sapiens males the respective haplotype is
as follows. Only the predominant alleles in all haplogroups,
particularly in the oldest ones, were included (see Materials and
Methods):
11 12 11 11 10 8 10 8 12 10 12 12 12 11 11 11
The mutation rate constant for those 16 markers equals .00410-
Figure 3.
Comparison of the MSY fragments (1 - 97) from Humans and Chimpanzee, and the DNA fragments in the genome of Gorilla, Pongo and
Macaque (whole genome shotgun sequence), along with a consensus tree fitting mutation patterns (inset).

A. A. KLYOSOV ET AL.
AK mutations per haplotype (Klyosov, 2011). The two haplo-
types differ by 64 mutations, or, by 4 mutations per marker on
average. The correction for back mutations is determined as (1
+ e4)/2 = 28 (ibid.). This means that the calculated time to the
common ancestor of H. sapiens and Pan troglodytes must be
multiplied by 28. The sixty-four mutation difference between the
MSY human and chimpanzee haplotypes computes to a common
ancestor equal to 64 0.004115,610437,080 conditional
generations (the arrow indicates a correction for back muta-
tions), or 10.9 ± 1.7 million years distance between the present
day Y-chromosomes of chimpanzees and humans. In other
words, the split occurred 5.5 ± 0.9 million years ago (the ob-
tained timespan should be divided by two, since it separates the
present day MSYs). This fits the numbers which are commonly
cited by geneticists, e.g. 5.6 - 8.3 million years ago (Green et al.,
2010) or 3.9 - 5.9 million years ago (Sun et al., 2011), or the
figures of 6 and 5 - 7 million years ago, given in the preceding
section of this paper. This result also shows that calculations
using slow mutated markers can be employed even for millions
of years back.
Nucleotides and SNPs in the Y-Chromosomes of
Chimpanzee and Humans
Let us follow some mutations in the Y-chromosome from
chimpanzee to several principal human lineages: A0, A1, A1a,
A1b, and the β-haplogroup (which is close to or identical with
BT). This might give us a clue to the direction of mutational
flow and, therefore, to which lineage was derived from which.
Here we have to adhere to one condition: we cannot call anyone
of lineages African unless we know that it actually originated in
Africa. We know only where the majority of bearers of the
subclades presently live. For example, all 11 individuals as-
signed to the A0 haplogroup live in the United States, with
“unknown origin” marked on the list in the Haplogroup A Pro-
ject (June 2012); one is marked as African American. In other
words, we simply do not know where their ancestors lived, say,
100,000 years ago; we assume that they lived in Africa, how-
ever, it is not enough to determine that “their common ancestor
arose in Africa”.
Seventeen individuals assigned to the A1a subclade and its
four sub-lineages, currently live in England, Switzerland,
Finland, Cape Verde, and the US. Most of them marked un-
known origin; however, some of them indicated European
countries as origin of their recent ancestors. Forty-six bearers of
A1b1 and its downstream subclades currently live in England,
Ireland, Scotland, Italy, Algeria, Turkey, Kuwait, Saudi Arabia,
Tunisia (one), Chad (one), South Africa (one), the US. None of
the individuals in the project belongs to upstream A1 or A1b
subclades. Therefore, SNPs V168, P108 and V221, which are
actually observed in BT (β-haplogroup) and down to hap-
logroups R1a and R1b, as well as in parallel downstream sub-
clades of A1b, could have been inherited as mutations from a
common α-haplogroup ancestor of both sides of the bifurca-
tion shown in Figure 1.
In the description below, one should take into account that
haplogroup A is a collective, a tentative grouping, which in-
cludes all that is not in haplogroup BT. Indexes, which are
(tentatively) assigned to haplogroup A subclades, change every
few months or even weeks. There is nothing African in the
indexes of the subclades. We can only assume, for the sake of
explanation, that 1) the α-haplogroup in Figure 1 corresponds
to A1/A1b haplogroups in the current classification; 2) one side
of the bifurcation represented in Figure 1 leads to the
β-haplogroup (currently BT or sometimes BR in the classifica-
tion); and 3) another leads to the “African” subclades, hap-
logroups A1a and downstream subclades of A1b. We can as-
sume that the latter are “African”, since their SNP mutations
are not observed in the β-haplogroup, including Europeoids.
The word “African” is in the quotations marks since we actu-
ally do not know where their common ancestors lived some
100,000 ybp or before (or after) that.
A0 lineages. There are two principal types of A0 lineages: 1)
those which retain the nucleotides found in MSY of living
chimpanzee (Table 1), and 2) those which contain specific
SNPs compared with the ancestral nucleotide in chimpanzees (Ta-
ble 2). These two types make haplogroup A0 very distinct from
those (Africans and non-Africans) ancestors of which descended
from the α-haplogroup. For the most cases A1 A1bBT
bearers, for example, have those 1)—type nucleotides mutated
into the respective SNPs; and 2)—type nucleotides retained as
ancestral in chimpanzee. It does not mean that the A0 bearers
are closer to chimpanzees than to other humans. It means that
the Y-chromosome in A0 bearers evolved (mutated) independ-
ently compared to other “African” and “non-African” lineages.
Besides, as shown above, A0 is significantly older than the α-
haplogroup.
As a result, H. sapiens living today have three identified thus
far principal MSY lineages: 1) “Common African” (part of
subclades of haplogroup A, branched from the α-haplogroup
and formed the left-hand bifurcation in Figure 1); 2) “non-
African” (descendants of the β-haplogroup, branched from the
α-haplogroup and formed the right-hand bifurcation in Figure
1); and 3) bearers of A0 lineages. Those three principal MSY
lineages are very different in terms of their Y-chromosomes,
however, they are brought much closer together by maternal
part of their DNA, so their resulting genome is adjusted with
every new generation.
In addition to those listed in Table 1, the following SNPs
have been assigned to the BT haplogroup in the current classi-
fication, and may be directly related to the α-haplogroup: L413,
L418, L438, L440, L604, L957, L962, L969, L970, L971, L977,
L1060, L1061, L1062, SRY10831.1, M42, M94, M139, M299,
P97, PK1.
The α-haplogroup. Although no individuals bearing A1 or
A1b SNPs are listed in the FTDNA Haplogroup A Project or in
the Walk Through YDNA Project, it can be safely assumed,
based on the SNPs patterns, that the α-haplogroup includes A1
and A1b subclades. Haplogroup A (in its entirety) is a hap-
logroup which split from the α-haplogroup some time after
160,000 ± 12,000 ybp; haplogroup A contains many mutated
nucleotides (SNPs) which are not mutated in A0 lineages. The
mutated SNPs in the α-haplogroup (A1/A1b) are inherited not
only by the African subclades but also by the non-African sub-
clades.
All 77 mutations shown in Table 1 represent SNPs for the α-
and β-haplogroups. The SNPs assigned to A1b are P108 and
V221; the SNPs assigned to BT are V29, V59, V64, V235 (and
those immediately below Table 1), total 27 SNP’s. The re-
maining 50 mutations in Table 1 are the SNPs assigned to A1,
according to the current classification. SNPs P305 and L986 are
mistakenly assigned to the A0 subclades in the current version
of ISOGG-2012 (June 2012); mistakenly, because they do not
204

A. A. KLYOSOV ET AL.
Table 1.
Ancestral nucleotides in chimpanzee MSY which are retained in A0
lineages, but are mutated in the α-haplogroups (i.e. in A1/A1b/BT).
Nucleotides SNP
Chimp A0 α-haplogroup
C C T P108
A A G P305
A A C L985
G G A L986
T T A L989
A A G L990
A A T L1002
G G del L1003
T T C L1004
A A G L1009
C C A L1053
T T C L1085
G G C L1089
G G C L1090
A A G L1093
C C A L1098
A A G L1099
T T C L1101
C C T L1105
C C A L1013
G G A L1114
A A C L1116
T T C L1118
G G T L1120
C C T L1123
T T C L1124
A A G L1125
A A G L1127
C C T L1128
T T G L1130
A A G L1132
C C A L1135
A A G L1136
C C T L1137
Continued
C C T L1142
A A G L1143
C C T L1145
A A G L1150
TAGG TAGG del L1153
G G C L1155
C C T V21
A A G V29
G G A V41
C C T V54
C C T V59
T T A V64
A A G V161
G G A V168
C C G V171
A A G V174
T T G V203
G G T V221
A A G V235
G G T V238
C C T V241
A A G V250
have the respective SNPs in A0, those SNPs belong to the
α-haplogroup (see Table 1).
Table 2 lists SNPs which belong to the A0 lineages; those
nucleotides are retained ancestral in the α-haplogroup. It again
shows the principal difference between A0 and BT.
In addition to the mutated nucleotides listed in Table 2, the
following ancestral nucleotides are mutated in A0 lineages but
retained in the α-haplogroup:
A—L1086, L1094, L1102, L1109, L1112, L1113, L1117,
V79, V81, V141, V223, V242,
C—L1088, L1097, L1100, L1103, L1104, L1107, L1108,
L1111, V164,
G—P114, L1087, L1091, L1096, L1106, L1115, L1119,
V139,
del—V152.
A total 62 SNPs in A0 lineages are listed above plus 30 more
(L981, L983, L988, L994, L1007, L1014, L1070, L1072,
L1073, L1075, L1076, L1078, L1079, L1080, L1081, L1082,
V150, V151, V153, V157, V158, V159, V161.1, V162, V164,
V169, V170, V181, V195, V203) which are currently (June
2012) identified and listed in public sources (see Materials and
Methods). The difference between the listed 52 mutations in
A1/A1b (α-haplogroup) and the 92 mutations in A0 lineages
compared to the ancestral nucleotides in chimpanzee MSY

A. A. KLYOSOV ET AL.
Table 2.
Ancestral nucleotides in chimpanzees which 1) are retained in the α-
haplogroup 
A1 A1bBT and 2) show a mutation in A0 lineages.
Nucleotides SNP
Chimp A0 α-haplogroup
A G A L529
A T A L896
C A C L982
G T G L984
C A C L991
A C A L993
T C T L995
TTA del TTA L997
T G T L998
C T C L999
C A C L1000
G T G L1001
A G A L1005
A T A L1006
C A C L1008
T C T L1010
C T C L1012
T C T L1016
T C T L1018
G A G V148
T G T V149
A G A V154
G A G V166
G A G V172
T G T V173
A T A V177
G T G V196
A G A V225
G A G V229
C T C V233
G A G V239
shows that A0 lineages and the α-haplogroup are remote from
each other; however, these data should not be used for calcula-
tions of age, because not all SNPs have been identified or as-
signed as yet.
A1a subclade. The A1a subclade is downstream from the A1
subclade; thus far, it has five SNPs identified (M31, P82, V4,
V14, V25) which were not inherited by the β-haplogroup. It
seems that A1a split from the α-haplogroup between 160,000
and 60,000 ybp, and eventually ended up in Africa, Europe,
Asia (see above). Most identified bearers of A1a’s currently
live in Europe or the US; needless to say, this does not tell us
Subclades downstream from haplogrou
much about where the subclade originated.
p A1b. The SNPs
of
tly, only one bearer of the A1b1 subclade is listed in
th
y on nucleotides in
ch
nstream SNP-mutations occurred in the
bo
tides and the
fo
these subclades (e.g. A1b1, A1b1a and A1b1b2-M13) are not
observed in the BT haplogroups, therefore, it appears that bear-
ers of these subclades ended up in Africa some time after their
ancestors split from the α-haplogroup (A1/A1b). However,
even this suggestion cannot be certain since many A1b1b2-
M13a bearers currently live in England, Ireland, Scotland, Italy,
Turkey, Algeria, Kuwait, Saudi Arabia, as well as in Tunisia
and Chad (cf. the FTDNA Haplogroup A Project). We do not
know when or where the subclades downstream from A1b
arose.
Curren
e FTDNA Haplogroup A Project (as origin unknown). The
majority of identified bearers of the haplogroup A subclades
belong to A1b1b2-M13 (60%); 23% belong to the A1a sub-
clade, 15% belong to the A0 subclades).
Once we have established that we can rel
impanzee MSY which are either inherited or generally intact
in the human Y-chromosome, or which mutate into the respect-
tive SNPs in certain lineages of human male haplogroups, we
can try to interpret the diagram in Figure 1 in terms of the an-
cestral nucleotide and acquired SNPs flow. Figure 1 shows that
incoming nucleotides of chimpanzee (from below on the dia-
gram) to the α-haplogroup should either flow as unchanged,
ancestral, non-mutated, or as mutated into the respective SNPs
into the α-haplogroup and then, mutated, to the both sides of the
tree. Therefore, those ancestral (from chimpanzee) and the
SNP-mutated (in the α-haplogroup) nucleotides are expected to
be present in both “common Africans” (not A0 bearers) and all
non-Africans. It is exactly what is observed. It does not mean
that non-Africans descended from an African common ancestor.
Those African and non-African branches of the tree are parallel
ones (see Figure 1).
Additionally, dow
th branches after they split from the α-haplogroup. Those
include 1) SNPs of the A1a subclade and its (future) down-
stream subclades, as well as downstream subclades of A1b,
which are expected to be observed in Africa. However, only
few of them have been identified there thus far due to a poor
representation of African haplotypes. Those SNPs have not
been found in bearers of haplogroups B through T, including
most of non-Africans. Those also include 2) relatively recent
SNPs in the descendents of the β-haplogroup, which (SNPs)
have resulted in the 19 principal haplogroups (B to T) down-
stream of the β-haplogroup. Those SNPs, of course, cannot be
found among bearers of haplogroups A and A0.
What is the pattern of intact (ancestral) nucleo
llow-up SNPs? First, we will look at two A0 lineages, actu-
ally identified in the WTY Project, and placed on the ISOGG-
2012 list as four subclades (A0, A0a, A0a1, and A0b). They are
not associated with the α-haplogroup. When we compare them
at 346 nucleotide sites, they differ from each other at 36 loci.
The two A0 lineages differ from each other in three blocks of
SNPs (between L92.2 and L348.2, between L979 and L1017,
and between L1036 and L1058), and in 153 sites—almost half
of the 346 nucleotides—they differ from the BT lineage. A pat-
tern of STR-mutations show that A0 lineages are much more
ancient than the α-haplogroup (Rozhanskii & Klyosov, 2011),
and did not descend from it. The data shown above suggest that
the α-haplogroup was the handle of the MSY fork from which
the “common African” and non-African lineages diverged and
evolved in parallel (see Figure 1); this pattern adequately de-
206

A. A. KLYOSOV ET AL.
scribes the evolution of the Y-chromosome of H. sapiens for
the last 160,000 years. This pattern leaves no room for the
Out-of-Africa hypothesis.
The deduced ancestral (base) haplotypes of haplogroups A
an
hat ana-
to
Conclusion
Our consideration o through T, and our
an
an
haplotypes, we
di
nucleo-
tid
hy
Materials and Methods
The SNPs cite1 and 2 and the
te
of the DNA with certain SNPs or STRs
in
/gb2/gbrowse/hschrY/ as follows:
e.ftdna.com/index.php?name=Draft&parent=root and
k
d B are shown above; the estimated time to their common
ancestors is between 132,000 ± 20,000 and 46,000 ± 5,000 years,
respectively (Klyosov & Rozhanskii, 2012a). These two hap-
lotypes differ from each other by as many as 18 mutations,
which translates to 123,000 years between them. Their common
ancestor lived approximately 160,000 ybp (Klyosov & Roz-
hanskii, 2012a). One can see that haplogroup B could not possi-
bly have descended from haplogroup A. However, the problem
would be resolved if the two had a common ancestor who lived
160,000 ybp. We do not know where this common ancestor
lived; there certainly is no indication that he lived in Africa. Ne-
anderthals did not live in Africa and, as it is known, did not have
a melanin-rich skin. Thus, it is not necessary to presume that all
lineages of the genus Homo had their origin in Africa.
Based on palaeoarchaeological evidence, it appears t
mically modern humans are likely to have originated some-
where in the vast territory from West Europe across the Russian
Plain in the east, and to the Levant in the south. Each of these
regions is renowned for the age of the skeletal remains of mod-
ern humans dating back to 45,000 - 40,000 ybp (for the latest
references see Highham et al., 2011; Benazzi et al., 2011).
f haplogroups A
alysis of the dynamics of the Y-chromosome nucleotide flow
from primates to humans during the evolution of genus Homo
has shown that a common ancestor of the majority of present
day human males, both Africans and non-Africans, lived
160,000 ± 12,000 years ago. This common ancestor has been
identified as belonging to the α-haplogroup, which is equiva-
lent or close to haplogroups A1/A1b in the current phylogeny.
The archaic lineages (currently designated A0) descend from
ancestor who lived at least 180,000 years ago (or much ear-
lier). The α-haplogroup and the A0 lineages have nucleotide
patterns that are distinct from each other, and both partly retain
ancestral MSY chimpanzee nucleotides and partly have them
mutated into the respective SNPs. Our research has shown that
at least 90% of chimpanzee MSY nucleotides are the same as
the nucleotides of living H. sapiens males. This comparison
points up the areas of change in H. sapiens, which allows us to
use chimpanzee MSY as a proxy for genus Homo’s common α-
haplogroup ancestor. It is clear that when they are compared to
loci in the DNA of other primates, such as gorillas, orangutans,
and macaques, many human Y-chromosome loci have been
conserved from our common ancestor.
When we studied slow mutating 16-marker
scovered that chimpanzees and present day humans had a
common ancestor 5.5 ± 0.9 million years before the present.
This date is in agreement with dates obtained in genome studies.
It is important, since it shows that the DNA genealogy ap-
proach can be extended to millions of years in depth.
The results of our analysis of haplotypes, conserved
es, and SNPs suggest that there is no reason to believe that
non-Africans (haplogroup BT and its downstream haplogroups)
descended from haplogroups A0, A1a, or other African hap-
logroups. These data are adequately described by a model
which shows that both the African lineages and non-African
ones diverged from the α-haplogroup approximately 160,000
years before the present and that the Y-chromosomes of the two
groups have evolved independently since then. Of course, the
Y-chromosome constitutes only a small part of the whole ge-
nome, and the term “evolve” here is related to the human MSY
only. It can serve, however, a very useful “probe” to trace the
human evolution with all reservations concerning the genome.
We have no indication of where the common ancestor of the α-
haplogroup lived; he could just as easily have been from
Europe, Asia, or the Middle East, as from Africa. We believe,
however, that only the A0 lineages represent the “truly African”
autochthonous inhabitants (along with the respective female
lineages), and interbreeding with them of incoming lineages of
haplogroups A and B (as well as some lineages of haplogroups
E, R1b, etc.), again, with the respective female lineages, created
the very diverse current African population. Only further stud-
ies will show whether this hypothesis is valid.
We believe that all the presuppositions of the Out-of-Africa
pothesis fail to hold up under simple scrutiny. This study
shows that the Out-of-Africa hypothesis has not been ade-
quately substantiated. The common assertion that “anatomically
modern humans came out of Africa some 70,000 years ago” has
never been convincingly calculated or obtained otherwise. The
Out-of-Africa hypothesis has never been proved; our research
suggests that it is incorrect with respect to anatomically modern
humans.
d in this study (see, e.g. Tables
xt) were identified by Thomas Krahn of FTDNA Genomics
Research Center (indicated by beginning letter L, such as L985),
Rozaria Scozzari and Fulvio Cruciani of Universita La Sapi-
enza, Rome, Italy (letter V), Peter Underhill of Stanford Uni-
versity (letter M) and Michael Hammer of University of Ari-
zona (letter P) and publicly available in various databases (see
below). Since sequences of extended fragments of Y-chro-
mosome are published in databases only for humans, chimpan-
zee, and macaque, search for homologous fragments for gorilla
and orangutan has been conducted in genomes of the primates,
listed in the European Nucleotide Archive (ENA) database,
employing Whole Genome Shotgun (WGS). Human and pri-
mate gene sequences were aligned using the National Center
for Biotechnology Information (NCBI) GenBank and ENA (the
links are listed below).
Search for fragments
human Y-chromosome has been conducted using the FTD-
NAY-chromosome Browser
http://ymap.ftdna.com/cgi-bin
(1) introduce the target SNP or STR into the field Landmark or
Region, click on Search, chose a size of the target fragment
(100 bp is recommended for an SNP), copy that fragment and
find a homologous one in GenBank or ENA, using option
BLAST. SNPs in particular haplogroups were those identified
at sites
http://ytre
http://www.isogg.org/tree/. Haplotypes of haplogroup A to
compose the tree in Figure 2 are listed in the FTDNA Project.
The base haplotypes of haplogroup A and B were determined in
(Klyosov & Rozhanskii, 2012a). Repeat motifs for the STRs
(http://www.smgf.org/ychromoso me/marker_details.jspx? mar
er and http://www.genebase.com/in/dnaMarkerDetail.php) are

A. A. KLYOSOV ET AL.
listed below:
DYS426 = 11
TGAAAGCATGACCACTTCATT
n repeats (GTT) in the greatest number of haplogroups:
am
Chimpanzee (Y-Chromosome)
ght repeats (GTT)]
e (GTT)7 on the 14th chromosome,
or
DYS388 = 12
TGATTAGCCGTTTAGCGATAT
test number of haplogrou-
ps
Chimpanzee (Y-Chromosome)
DYS392 = 11
GTCATCGCAGTGGCCCAAGTGA
repeats (TAT) in the greatest number of haplogroups:
am
CCCAAGTAA
GTTGCTCCA
[T
GAGAATGATACTGCCTAAGCC
AAGGCTGCAGTGAGCTGTGAT
f hap-
lo ups world-
w
GCCTAAGCC
AGTGTGATC
D
CACATTGTTGTTAAGCCCAGC
CAATCTCCCTGTGGTCGGGGC
r of hap-
lo s in haplogroups world-
w
osome has no DYS454
GTGGGGAATAGTTGAACGGTA
TTTTCTTTTCTTTTCTTTTCT
CACCACAACCTCCACTTCCCAGGTTCAAGC
CTCAAAGTA
TATTGTGTTGTTGTTGTTGTTGTTGTTGTT
GTTGTTGTTGACACAAAGTCTCGTCTTGTC
ACC
[Eleve
ong 41 base haplotypes in haplogroups world-wide there are
33 with the allele 11 (including the beta-haplogroup), and eight
with 12 (mainly haplogroup P and its downstream haplogroups)
(Klyosov & Rozhanskii, 2012)]
CTCAAAGTATGAAAGCATGACCACTTCATT
TAGTTGTTTTTTTGTTGTTGTTGTTGTTGT
TGTTGTTGACACAAAGTCTCGTCTTGTCAC
C
[Ei
NOTE: Macaques hav
angutans have (GTT)7 on the 17th chromosome
GAATTCATG
ATACATATTATGAAACATTATTATTATTAT
TATTATTATTATTATTATTATTTGAGACGG
ACTCTCGCTCTGTCGCCCAG
[Twelve repeats (ATT) in the grea
: among 41 base haplotypes in haplogroups world-wide there
are 28 with the allele 12 (including the beta-haplogroup), seven
with 13, two with 14 (I and I1), one 16, one 15 (J1 and J2, re-
spectively), one 11 (A), and one with 10 (B2a2) (Klyosov &
Rozhanskii, 2012)].
GAATTCATGTGAGTTAGCCGTTTAGCGATA
TATACATATTATGAAACATTATTATTATAT
ATTATTATTATTATTATTATTATTATTATT
GAGACGGACTCTCGCTCTGTCGCCCAG
[15 repeats (ATT)]
TAGAGGCA
TCTTGCAACATCTCCATCCATGTTGCTCCA
AAGGACCCAATTTTACTGTAAATGGTTGTA
TAGTATTTTATGGTCTACATAGACCATATT
TACCATATGTTCATCCATATTTTCTTCATT
AATCTAGCTTTTAAAAACAACTAATTTGAT
TTCAAGTGTTTGTTATTTAAAAGCCAAGAA
GGAAAACAAATTTTTTTCTTGTATCACCAT
TTATTTATTATTATTATTATTATTATTATT
ATTATTATTTACTAAGGAATGGGATTGGTA
GGTC
[Eleven
ong 41 base haplotypes in haplogroups world-wide there are
27 with the allele 11 (including the beta-haplogroup), six with
13, three with 12, three with 14, one with 10 (R2) and one with
7 (D3a) (Klyosov & Rozhanskii, 2012)].
Chimpanzee (Y-Chromosome)
TAGAGGCAGTCATTGCAGTGG
TCTTGCAACATCTCCATCCAT
AAGGACCCAATTTTACTGTAAATGGTTGTA
TAGTATTTTATGGTCTACATAGACCATATT
TACCATATGTTCGTCCATATTTTCTTCATT
AATCTAGCTTTTAAAAACAACTAATTTGAT
TTCAAGTGTTTGTTATTTAAAAGCCAAGAA
GGAAAACAATTTTTTTCTTGTATCACCACT
TATTTATTATTATTATTATTATTATTATTA
TTATTTACTAAGGAATGGGATTGGTAGGT
en repeats (TAT)]
DYS455 = 11
CTGAGCCGA
CACAAGGTC
CACCCGAGGGCACTCCAGCCTGGGCAACAC
TGTGAGACCATATATCTAAAATAAATAAAT
AAATAAATAAATAAATAAATAAAT AAATAA
ATAACGGAAGAACACTCGTTTCCACCCC
[Eleven repeats (AAAT) in the greatest number o
groups: among 41 base haplotypes in haplogro
ide there are 33 with the allele 11, four with 10, two with 9,
and one with 8 (I1) (Klyosov & Rozhanskii, 2012)].
Chimpanzee (Y-Chromosome)
CTGAGCCGAGAGAATGATACT
CACAAGGTCAAGGCTGCAGTG
ACCCGAGGGCACTCCAGCCTGGGCAACACT
GTGAGAGCATATATCTAAAATAAATAAATA
AATAACGGAAGAA
[4 repeats (AAAT]
YS454 = 11
GACTGACCT
AACATATCA
ACAGGCAAAAGCAAAATA AATAAATAAATA
AATAAATAAATAAATAAATAAATA AATAAC
CTAGGTGCTAATCCAAGTGATATGTTACAA
TGTTTCCTGTTGACACAACCCAACCTGGGT
GAAGTGAAGAGCTACATGTC
[Eleven repeats (AAAT) in the greatest numbe
groups: among 41 base haplotype
ide there are 32 with the allele 11, eight with 12, and one with
13 (T1a) (Klyosov & Rozhanskii, 2012)].
Chimpanzee (Y-Chromosome)
NOTE: The chimpanzee Y-Chrom
homologous fragments.
DYS438 = 10
CCAAAATTA
AACAGTATA
TTTCTTTTCTTTTCTTTTCTTTTCTTTTCT
ATTTGAAATGGAGTTTCACTCTTGTTGCCC
AGGCTGAAATGCAATGGTGTGATCTCGACT
208

A. A. KLYOSOV ET AL.
GATTCTCCTGCATCAGCCTCCCAGGTAGCT
GGGATTATAGGCGTCTGCCACCACGCCCAG
CTAATTTTTTGTGTTTTTAGTAGAGACAGG
GTTTCACCATGTTGGTGAGGCTGGTCTCGA
ACTCCAGACCCTGGGTGATC
[Ten repeats (TTTTC) in the greatest number of haplogroups:
am there are
27 ta-haplogroup), eight with
11
A
TTCTTTTGT
CACTCTTGT
-
m
D
CATTCAAATCCTCTCCCACT
TAAATAAATAAATAAATAAATA
AATAAATAAATAAAAAGCCTTT
f hap-
lo world-
wncluding the beta-hap-
lo
osome has no DYS531
ho
D
ACTCTAGCCTGGATGACACAAC
ATCTCAAATAAATAAATAAATA
pes out
of; 10 in F;
anskii, 2012)].
TGACAGAAC
AAATAAATA
GACAACAATTGTGTGTAAGTGACAAAATGT
D
AACTGAAAATCAT
TTTTGTTTTGTTTTG
TGTTTTGTTTTGTTTTGTTTTT
TTCTTGCTCTGTCACCC
plotypes
ou & Roz-
ha
C
GAGACTCTTCCATCTAAATAA
ATAAATAAATAAATAAATAAAT
types out
of Rozhan-
sk
C
GCCCGCTGCTCTCCAGCCTGG
TCTAAATAA
ATAAATAAAT
D
CACTCCAGCCTGGCG
AAT
TAATAATAATAAT GCCTCTTTG
AGTGCCTC
groups]
C
TAATAGTAAT
TTGCTGAAC
llow the
hu
-chromo-
some, however, shows (TAA)4 on the 3rd chromosome.
ong 41 base haplotypes in haplogroups world-wide
with the allele 10 (including the be
, and six with 9 (Klyosov & Rozhanskii, 2012)].
Chimpanzee (Y-Chromosome)
CCAAAATTAGTGGGGAATAGTTGAATGGT
AACAGTATATTTTCTTTTCTT
TTTCTATTTGAAATGGAGTTT
TGCCCAGGCTGAAATGCAATGGTGTGATCT
CGACTCACCACAACCTCCACTTCCCAGGTT
CAAGCGATTCTCCTGCATCAGCCTCCCAGG
TAGCTGGGATTATAGGCGTCTGCCACCATG
CCCAGCTAATTTTTTGTGTTTTTAGTAGAG
ACAGGTTTTCACCATGTTGGTGAGGCTGGT
CTTGAACTCCAGACCCTGGGTGATC
[Repeats (TTTTC)3(TTTTG)1(TTTTC)1] [described in (Gus
ao et al., 2002)]
YS531 = 11
GACCCACTGG
GCAAAAAA
AATAAATA
CGTCTACAAAGAAAGGGAGCA
[Eleven repeats (AAAT) in the greatest number o
groups: among 41 base haplotypes in haplogroups
ide there are 30 with the allele 11 (i
group), 10 with 10, and one with 12 (I2*) (Klyosov & Roz-
hanskii, 2012)].
Chimpanzee (Y-Chromosome)
GACCCACTGGCATCCAAATAACCTCCCTTG
GCAAAAAAAAAA…
NOTE: The chimpanzee Y-Chrom
mologous fragments.
YS578 = 8
GAGGCGGAACTTTCAGTGAGCCGAGATCAC
GCCACTCC
AAAACTCC
AATAAATAAATAAATAAA GTAAGTAAGACA
GACAACAACTGTGTGTAAGTGACAAAATGT
CCAGGGTTGTTGAAGC
[Eight repeats (AAAT) in 36 haplogroup base haploty
41 world-wide, except 9 in C3, F3 and O, NO and O
d 7 in J2 (Klyosov & Rozhan
Chimpanzee (Y-Chromosome)
GAGGCGGAACTTTCAGTGAGCCGAGATCGC
GCCACTCCGCTCTAGCCTGGG
AAGACTCCATCTTAAATAAAT
AATAAATAAATAAATAAAT AAATAAAGACA
CCAGGGTTGTTGAAGC
[Nine repeats (AAAT)]
YS590 = 8
GAACATAGTCGGGCT
AGTTGGGCAAGTTTT
GT
TTTTGTTT
GAGACGGA
[Eight repeats (TTTTG) in 39 haplogroup base ha
t of 41 world-wide, except 7 in A and L (Klyosov
nskii, 2012)].
himpanzee (Y-Chromosome)
NOTE: The chimpanzee Y-Chromosome has no DYS590
homologous fragments.
DYS641 = 10
CTTGAGCCCAGGAAGCATAGGTTGCAGTGA
GCTGAGATCGCCTGCTGCTCTCCAGCCTGG
TGATAGAGA
ATAAATAA
AAATGCTCCACAGCTAGGTGATATTGTTAA
TTGTTAGGTAACAGTTATTACAGACAAGAA
AGCCTAATTTACAAATAAAGGACAAAATTC
ATCGTGTGG
[Ten repeats (TAAA) in 39 haplogroup base haplo
41 world-wide, except 7 in NO and O (Klyosov &
ii, 2012)].
himpanzee (Y-Chromosome)
CTTGAGCCCAGGAAGCAGAGGTTGCAGTGA
GCTGAGATT
TGATAGAGAGAGACTCTTCCG
ATAAATAAATAAATAAAT AA
AAAAAGTTGCTCCACAGCTAGGTGATATTG
TTAATTGTTAGGTAACAGTTATTACAGACA
AGAAAGCCTAATTTACAAATCCAGGACAAA
ATTGCATCGTGTGG
[Ten repeats (TAAA)]
YS472 = 8
AGATTGTCCCACCT
ACACAGGAAGGTTCCATCTCAAATAGT
G
AATAATAA
CTGAACAC
[Eight repeats (AAT) in base haplotypes of all haplo
himpanzee (Y-Chromosome)
AGATTGTCCCACCTGCACTCCAGCCTGGCG
ACACAGGAAGGTTCCATCTCAAATAGTAAT
AATAATAATAATGATAATAA
AATAATAATAATAATGCCTCT
ACAGTGCCTC
[Five repeats (AAT) in this particular case if we fo
man homology].
NOTE: Macaques does not show homology on Y

A. A. KLYOSOV ET AL.
O
he 11th chromosome.
AA
CTATTGGAGGAGGAGAACCCT
AGAAGAAGAGAGAAACAGGCT
pes out
of
20
C
TATAGAAGGCAAGCAAGCTAA
GAGAACCCT
GAAACAGGCT
D
AATAAATCTAAAGCACATAAA
AATAAAATAAAATAAAATAAAA
of hap-
lo world-
weta-hap-
loh 13 (Klyosov &
R
GCACATAAA
TAAAAAAAAC
D
AGCAG
TGTTGTTGTTGT
TATCACCTCTAACGCAGCTCG
CTGTGTCCTGCTTCCATTGGC
f hap-
lo s world-
we beta-hap-
lolyosov & Rozhanskii, 2012)].
TGTCCTGCT
GC
D
ATTATTATTATT
AGTCTTGCTCTGTCACCCAGGC
GTGGCGTGATCTCAGCTCAGT
of hap-
lo s world-
wbeta-hap-
lo3, three with 15 and one with 16 (Klyosov
&
AGTCTTGCTCTGTCAC
ATCTCGG
D
TTTCTATTGCAGCA
ACATAGGTGTGTC
TGTGTGTGTGTGTAAGCATGTA
CGGTTGAACAAACTAGGATCAC
(Redd et
al er, among
41 are 33
w beta-haplogroup) and eight with 7
(K
ATAGGTGTGTG
TGTGTGTG
TGTTGAACA
rangutan shows (TAA)4 on the 18th chromosome and (TAA)2
(TGG)1 (TAA)4 on t
DYS425 = 12
GTGGGCTGAGAAATTTCTGGAGACACTTCT
CTTTCTGTCTATAGAAGGCAAGCTAGCT
CTCTCTTCA
GGATTGGAG
CTAGAATTTAGAAAAATGTTGTTGTTGTTG
TTGTTGTTGTTGTTGTTGTTGTTTAATTTC
CATTTTACCTCCAGAATTACT
[Twelve repeats (TGT) in 36 haplogroup base haploty
41, except G2a, G2c, H, H1, L (Klyosov & Rozhanskii,
12)].
himpanzee (Y-Chromosome)
GTGGGCTGAGAAATTTCTGGAAACATTTCT
CTTTCTGTC
CTCTCTTCACTATTGGAGGAG
GGATTGGAGAGAAGAAGAGA
CTAGAATTTAGAAAAATGTTGTTGTTGTTG
TTGTTGTTGTTGTTGTTGTTGTTTAATTTC
CATTTTACCTCCAGAATTACT
[Ten repeats (TGT)]
YS594 = 10
GATGTGCCTAATG
AAAATGGTAAATATTATATTATATTGTATA
CCACAGAATGTATACTT
TATAACAAT
AGAAATAA
TAAAATAAAATAAAATAAAATAA AAAA AAC
AGAAAATACTCGACTATTGGTAAGAACGTT
GCAGCCCCAATGGAAAAGAGCCAGAAAGTT
ACTCAAAGAAATAAAGAAATAGGAAGTTGG
ACTGAGGACACGATTAACACCAGGG
[Ten repeats (TAAAA) in the greatest number
groups: among 41 base haplotypes in haplogroups
ide there are 22 with the allele 10 (including the b
group), 14 with 11, four with 12 and one wit
ozhanskii, 2012)].
Chimpanzee (Y-Chromosome)
GATGTGCCTAATGCCACAGAATGTATACTT
AAAATGGTAAATATTATATTATATTGTATA
TATAACAATAATAAATCTAAA
AGAAATAAAATAAAATAA AA
AGAAAATACTCGACTATTGGTAAGAACGTT
GCAGCCCCAATGGAAAAGAGCCAG AAAGTT
ACTCAAAGGAATAAAGAAATAGGAAGTTGG
ACTGAGGACACGATTAGCACCAGGG
[4 repeats (TAAAA)]
YS436 = 12
CCAGGAGAGCACA
GGTTGTTGTTGTTGTTGT
CACAAAAAGGAA
TGTTGTTTT
TCCCTTTTA
TGAAGTTGGATTGC
[Twelve repeats (GTT) in the greatest number o
groups: among 41 base haplotypes in haplogroup
ide there are 36 with the allele 12 (including th
group) and five with 11 (K
Chimpanzee (Y-Chromosome)
CCACGAGAGCACACACAAAAAGGAAAGCAG
GGTTGTTGTTGTTATGTTTATCACCTCTAA
CGCAGCTCATCCCTTTTACTG
TCCATTGGCTGAAGTTGGATT
[4 repeats (GTT)]
YS490 = 12
AGTATGTCCTTG
ATTATTATTATTATTATT
ACAACATTCTTTATTATT
TGAGACGG
TGGAGTGCA
GCAAGCTCTG
[Twelve repeats (TTA) in the greatest number
groups: among 41 base haplotypes in haplogroup
ide there are 33 with the allele 12 (including the
group), four with 1
Rozhanskii, 2012)].
Chimpanzee (19th Chromosome)
AGTATGTCCTTACCTTATTATTATTATTAT
TATTATTGAGACAG
CCAGGCTGGAGTTCAGTGGCATG
CTCACTGCAGCCTCTG
[Seven repeats (TTA)]
YS450 = 8
AGAGACCAGCTTGG
TTTTGAGAGAACGTGAA
TA
CTGTTGTG
TACATAAC
TCACACAATGGAATATGATGCAGCTGTTTG
TAGATCTGGTAGAAATTATTCTGAATCTAT
CGGTTCCTGGGCCGTTTTTTATTTTATTTT
ATTTTATTTTATTTTATTTTATTTTATTTT
ATTTCATTTTATATTTTATTTTATTTTATT
TTTATTTTATTTTTGGCTGGTGGGTTATTT
ATTACTGATTCCATT
[Fourteen repeats (TTTTA) are shown according to
., 2002) as (TTTTA)9(TTTTA)3 (TTTTA)2; howev
base haplotypes in haplogroups world-wide there
ith the allele 8 (including
lyosov & Rozhanskii, 2012)].
Chimpanzee (Y-Chromosome)
AGAGGCCAGCTTGGCATTTCCACTGCAGCT
TTTTGAGAGAACATGAAAC
TGTTTGTATGTGTGTTTGTGTG
TGTAAGCATACACACAGAACC
AACTAGGATCACCCACACAATGGAATATGA
TGTAGCTTTTTGTAGATCTGGTAGAATTTT
CCAAATCTATCAGTTTCTGGGCTTTTTTTT
…
210

A. A. KLYOSOV ET AL.
NOTE: The chimpanzee Y-Chromosome has no DYS454 ho-
m
D17 = 12
TCTTTTTATTATTAT
TATTATTATTATTATTATACT
AGGGTACATGTGCACAATGTG
of hap-
lo s world-
wta-hap-
loith 11 and only one with 14
(K
TTCTTCTTCT
TCAAGTTTT
CAGGTTAGT
]
D
TGCAGTGAGCTGAGATTGGGTG
CAGCCTGGGCAACAAAAGCAA
lo ps world-
w beta-hap-
logroup), 10 with 12, three with 10 and one with 9 (Klyosov &
R
TGAGGCAGGAGAGTCACTTGAAACTGGAAG
GATTGTATC
GAGCAA
ology]
D
GATACCACCTTAAGTATATAC
TTCTGTACTTCTACCTGCTTC
lo ps world-
w beta-hap-
lo)].
C
AATATATAC
CATGCTTC
D
TCTTTCCATGTTGCTCAGGCTA
TCCTGAGCTGACATGATCCTT
, C, D, E,
G
C
ATTGATAGAG
CTAGTCTCA
CTTCCTCCT
D
ht roup_A/default.as
px
12
sogg.org/tree/.
ologous fragments.
YS6
AGCATGATGCCTTCAGCTTTGTTCTTTCTG
CTTAGTACTGTGTTT
TATTATTAT
TTAAGTTTT
CAGGTTAGTTAGTTACATACGTATATATGT
GCCATGCTGGTGTGCTGCACCCATTAACTT
GTCATTTAGCATAAGGTATGTCTCCTAATG
CTATCACTCCCCAATCC
[Twelve repeats (TTA) in the greatest number
groups: among 41 base haplotypes in haplogroup
ide there are 25 with the allele 12 (including the be
group), 12 with 13, three w
lyosov & Rozhanskii, 2012)].
Chimpanzee (Y-Chromosome)
AGCATGATGCCTTCAGGTTTGTTCTTTATG
CTTAGTATTGTGTTTTCTTC
TCTTCTTATTATTATTATACT
AGGGTACATGTGCACAATGTG
TAGTTACATATGTATATATGTGCCATGCTG
GTGTGCTGCACCCATTAACTTGTCATTTAG
CATAAGGTATGTCTCCTAATGCTATCACTC
CCCAATCC
[Repeats (TTA)4 if we follow the human homology
YS568 = 11
GTGGCAGACAAAACCCAGTTACTCGGCAGG
CTGAGGCAGGAGTGTCACTTGAAACCAGAA
GGTGGAGG
ACAACACTC
AACTCCTTCTCAAAAATAAATAAATAAATA
AATAAATAAATAAATAAATAAATA AATAAA
TAAGTCCTCTTTTCCATCTTCCTCTTCAAT
ACCATTGTTTTCCCACGCTTACTGCAAATT
CGCCTAGATGAGTCCCATCCCTTTTCAA
[Eleven repeats (TAAA) in the greatest number of hap-
groups: among 41 base haplotypes in haplogrou
ide there are 27 with the allele 11 (including the
ozhanskii, 2012)].
Chimpanzee (Y-Chromosome)
GTGGCAGGCATTACCAGCTACTCAGGAGGC
GCGGAGGTTGCAGTGAGCCGA
AGTACACTCCAGCCTGGGCAACAA
AACTCCTTCTCCAAAAATTAAATAAATAAA
TAAATAATAACATAAAGAAAGAAAG AAAGA
AAGAAAGAAAGAGTCCTCTTTTCCATCTTC
CTCTTCCATACCATTGTTCTCCTGCACTTA
CTATAAATTTGCCTAGATGAGTCCCACCCT
TTTCAA
[Four repeats (TAAA)4 if we follow the human hom
YS640 = 11
TGGGAAAAACCATGAGATCCTGTCTCAAAA
AATAAATAAATAAATAAATAAATAAATAAA
TAAATAAATAAATAAATCCATTATTGCCCA
ATAGTTTGA
TAAACTTCC
CAGATGTTCTTTTATATTCCTCTAGTCTTT
TTGTGTGTATGAGTGTTTTCATGCATCTGT
AACATATTATTTTCATTATAGATGGTTAAT
AATGTCTTTAAAATATGAACGGGCTTGACC
CTA
[Eleven repeats (TAAA) in the greatest number of hap-
groups: among 41 base haplotypes in haplogrou
ide there are 28 with the allele 11 (including the
group) and 13 with 12 (Klyosov & Rozhanskii, 2012
himpanzee (Y-Chromosome)
TGGGAAAAACCGTGAGATCCTGTCTCAAAA
AATAAATAAACAAATAAATAAATAAATAAA
TAAATAAACAAACCAACCCATTATTGCCCA
ATAGTTTGAGATACCACCTTA
TAAACTTCCTTCTGTACTTCTA
TAGATGTTCTTTTATATTCCTCTAGTCTTT
TTGTGGGTGTGTGTGTTTTCATGCATCTGT
AACATATTATTTTCATTATAGATGGTTAAT
AATGTCTTTAAAATATGAACGGGCTTGACC
CTA
[Repeats (TAAA)2(CAAA)1(TAAA)6 (CAAA)1]
YS492 = 11
AGATGAGCCAGGCTTCAGACCCATGCAGCT
CCATTTCAAGCCCATATCCTATCATTATTA
TTATTATTATTATTATTATTATTATTATTG
ATAGAGAG
GTCTCAAAC
CCTCCTCAGCCTCCAAAAAACCTGGGGTAA
CAAGTGCGAGCCATTGTGCCTGACCCCTAC
T
[Eleven repeats (TTA) in the oldest haplogroups: B
(Klyosov & Rozhanskii, 2012)].
himpanzee (Y-Chromosome)
AGATGAGCCAGGCTTCAGACCCATGCAGCT
CCATTTCAAGCCCATATCCTATCATTATTA
TTATTATTATTATTATTATT
AGTCTTTCCATGTTGCTCAGG
AACTCCTGAGCTGACATGATC
CAGCCTCCAAAAAACATGGGGTAACAAGTG
CAAGCCATTGTGCCTGACCCCTACT
[Nine repeats (TTA)]
atabases
The FTDNA Haplogroup A Project:
a.com/public/Haplogtp://www.familytreedn
?section=yresults.
ISOGG-20
http://www. i
European Nucleotide Archive (ENA)

A. A. KLYOSOV ET AL.
http://www.ebi.ac.uk/ena/
The National Center for Biotechnology Information (NCBI)
GenBank (http://www.ncbi.nlm.nih.gov/)
ts
ge expansion from Africa. Science, 332,
346-349. doi:10.1126/science.1199295
Acknowledgemen
The authors are indebted to Dr. Judith Remy Leder and Dr.
Alexander S. Zolotarev for their valuable help with the prepara-
tion of the manuscript.
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