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Evidence of early gene flow between Ashkenazi Jews and non-Jewish Europeans in mitochondrial DNA haplogroup H7



To investigate European introgression into Ashkenazi Jewry, the European-dominant haplogroup H mi-tochondrial DNA was examined. The results provided genetic evidence that gene flow between Jewish and non-Jewish populations occurred early in Jewish settlement in Europe with isolation of the groups thereafter. We targeted branch H7 and found three Ashkenazi Jewish clades, two that were not previously recognized as Jewish (H7e, H7c2) and one newly identified group (tentatively H7j) characterized by 1700C and 152C transitions. A total of 100 new complete mitochondrial DNA sequences (mitogenomes) are reported, including the largest collection of H7e to date. H7e is a deeply nested clade with several subclades; more than 85% of the carriers had Ashkenazi maternal ancestry from such diverse areas as Germany and Austria in Western Europe, Poland, and the Baltic states in Central Europe, and Moldova, Ukraine and Belarus in Eastern Europe. Between 10% and 15% of the carriers had European non-Jewish ancestry which, strikingly, showed the greatest number of mutational differences from ancestral H7e. Moreover, there was no overlap with the Jewish-affiliated sequences other than at the ancestral node. Earlier research proposing early mixing followed by isolation has relied on less direct inferences. The smaller groups of H7c2 and H7j were exclusively Ashkenazi Jewish, with interesting sequence patterns. H7c2 consisted of a number of non-nested sister branches, reflecting recent expansion in a large population , while H7j showed a possible in-progress vanishing of the ancestral group, well on its way to mothering an orphan node. The severe bottleneck and subsequent population explosion in the Ashkenazim provide a unique opportunity to view haplogroups in all states of evolution and provide a window into the Mediterranean-Hellenistic world of antiquity.
Journal of Genec Genealogy 8(1):21-34, 2016
To invesgate European introgression into Ashkenazi Jewry, the European-dominant haplogroup H mi-
tochondrial DNA was examined. The results provided genec evidence that gene ow between Jewish
and non-Jewish populaons occurred early in Jewish selement in Europe with isolaon of the groups
thereaer. We targeted branch H7 and found three Ashkenazi Jewish clades, two that were not previ-
ously recognized as Jewish (H7e, H7c2) and one newly idened group (tentavely H7j) characterized by
1700C and 152C transions. A total of 100 new complete mitochondrial DNA sequences (mitogenomes)
are reported, including the largest collecon of H7e to date. H7e is a deeply nested clade with several
subclades; more than 85% of the carriers had Ashkenazi maternal ancestry from such diverse areas as
Germany and Austria in Western Europe, Poland, and the Balc states in Central Europe, and Moldova,
Ukraine and Belarus in Eastern Europe. Between 10% and 15% of the carriers had European non-Jewish
ancestry which, strikingly, showed the greatest number of mutaonal dierences from ancestral H7e.
Moreover, there was no overlap with the Jewish-aliated sequences other than at the ancestral node.
Earlier research proposing early mixing followed by isolaon has relied on less direct inferences. The
smaller groups of H7c2 and H7j were exclusively Ashkenazi Jewish, with interesng sequence paerns.
H7c2 consisted of a number of non-nested sister branches, reecng recent expansion in a large popula-
on, while H7j showed a possible in-progress vanishing of the ancestral group, well on its way to moth-
ering an orphan node. The severe boleneck and subsequent populaon explosion in the Ashkenazim
provide a unique opportunity to view haplogroups in all states of evoluon and provide a window into
the Mediterranean–Hellenisc world of anquity.
Evidence of early gene ow between Ashkenazi Jews and non-Jewish Europeans
in mitochondrial DNA haplogroup H7
Doron Yacobi & Felice L Bedford, Ph.D.
Address for correspondence:
University of Arizona, P.O. Box 210068, Tucson, AZ 85721, USA
Over the last decade, evidence has accumulated
that the genec make-up of Ashkenazi Jewry is a
combinaon of Levanne and European sources.
Analyses of autosomal genes, reecng a combi-
naon of paternal and maternal inheritance, have
indicated a signicant degree of European admix-
ture among Ashkenazi Jews as well as a close re-
laonship between most contemporary Jews and
non-Jewish populaons from the Levant (Atzmon
et al., 2010; Behar et al., 2010). The source of the
European contribuon may come from the mater-
nal line. Costa and colleagues (2013) argued that
the majority of the Ashkenazi mitochondrial hap-
logroups, which are inherited only from the moth-
er, were present in Europe long before the arrival
of Jews. However, Behar and colleagues (2006)
suggested that these same maternal haplogroups
most likely originated in the Levant alongside pa-
ternally inherited Y chromosomes of Levanne or-
igin (Atzmon et al., 2010; Ostrer & Skorecki, 2013).
When haplogroups have a notable presence in
both the Near East and Europe, determining their
geographic origins can be challenging and lead to
diering interpretaons. An example involves T2e,
a haplogroup that harbors a couple of unique Jew-
ish clades. Bedford (2012) reported prevalence of
T2e in Italy, Egypt, and parts of Saudi Arabia and
favored a Near Eastern rather than European ori-
gin of the mutaons that dene T2e but le open
the possibility that either locale could be the origin
or recipient of migraon. On the other hand, Pala
et al. (2012), using similar geographic incidences,
concluded that T2e’s origin was European.
In principle, esmates of when mutaons emerged
can help resolve where they emerged. In pracce,
however, standard deviaons of me esmates
can extend across greater than a thousand years,
and me esmates themselves can dier by an
order of magnitude depending on the esmated
mutaon rate. In research on Jewish groups, we
(Bedford et al., 2013; Bedford & Yacobi, 2014) re-
ported on a Bulgarian Sephardic founding lineage
(T2e1b), originally idened by Behar, which we
found among both Ashkenazi and Sephardic Jews
from diverse regions. Full genomic sequencing
found much coding-region variability, with several
haplotypes. Coalescence me for the sequences
using a common mutaon-rate esmate suggest-
ed that the shared mutaon (9181G) predated
the split between the Jewish groups and therefore
likely arose in the Levant. However, a dierent ,
also jusable mutaon rate suggested the origin
was much more recent, implicang geneow in
Europe aer the split as the source as of the mu-
taon common to both Sephardic and Ashkenazi
Diculty in disnguishing between Levanne
and European sources for Ashkenazi mitochon-
drial haplogroups is further muddled by an oen
overlooked historical fact: that the boundaries of
Europe and the Levant are a relavely recent his-
torical construct dang back to the Arab conquest
in the 7th century CE.
To further invesgate the role of European ma-
ternal admixture into the Ashkenazi gene pool,
we took a dierent approach than previous in-
vesgaons. Rather than surveying a large num-
ber of haplogroups with ambiguous geographic
origins, we conducted a detailed invesgaon
into a haplogroup that is overwhelmingly Euro-
pean (e.g., Brotherton et al., 2013) yet sll found
among modern Ashkenazi Jews. Haplogroup H
is the dominant European mtDNA haplogroup.
Its numerical success nears half the populaon
in some countries, making it the most common
haplogroup in Europe. Among Ashkenazi Jews,
23% have haplogroup H (Costa et al., 2013), yet
despite being a “majorAshkenazi haplogroup, it
is oen overlooked. When examining Ashkenazi
H mitogenomes, Costa and colleagues found that
most of them nest within west/central European
subclades, with closely matching sequences in
Eastern Europe. As such, haplogroup H’s gener-
al European dominance may illuminate issues of
introgression of European DNA into the Ashke-
nazi gene pool. Does haplogroup H reect recent
unions of non-Jewish women and Ashkenazi men,
or does it point to events of more distant interest?
We focused on H7. While other choices were pos-
sible, we selected H7 as an understudied clade
within haplogroup H that our pilot study suggest-
ed had an unexpected notable presence among
the Ashkenazim. Finally, we also delved into Med-
iterranean and Jewish history to place the genec
results within their correct historical framework.
A consideraon of relevant Mediterranean and
Jewish history is given in Appendix A. The combi-
naon of genec results and accepted history may
lead to a greater understanding of Jewish mater-
nal lineages.
Materials and Methods
To idenfy Ashkenazi clusters within haplogroup
H7, we inially selected two individuals with
self-described Ashkenazi Jewish maternal lineages
belonging to two dierent subclades of H7 from
the customer base at Family Tree DNA (FTDNA;
Houston, Texas, USA). FTDNA oers genec test-
ing services direct to individuals and has one of
the largest databases in the world of individuals
who have had their full mitochondrial genomes
sequenced, including many with European and
Ashkenazi Jewish roots. The data from FTDNA cus-
tomers is increasingly being used as a scienc re-
source (Bedford, 2012; Bedford et al., 2013; Behar
et al., 2012; Pike, 2006; Pike et al., 2010).
These two sequences were used as “kernels”, or
seeds, to search the FTDNA database for other full
Journal of Genec Genealogy 8(1):21-34, 2016
Journal of Genec Genealogy 8(1):21-34, 2016
mitochondrial sequences that diered by 0–3 mu-
taons, as in our previous study (Bedford et al.,
2013). These people were contacted by email and
invited to be part of the research study. They were
asked about 1) the addional mutaons they car-
ried in their mtDNA, 2) who their matches were
within 0–3 genec dierences, and 3) their deep
maternal ancestry. In this manner, a large number
of dierent haplotypes belonging to both H7 sub-
clades was idened, and a robust picture of all
members of these Ashkenazi Jewish clusters was
Thereaer, the database of the H7 mtDNA ge-
nome project (“H7 MtGenome”), co-administered
by one of us (Yacobi), was mined for addional se-
quences not uncovered by the above procedure.
Within the H7 MtGenome project, 229 parci-
pants had tested their full mitochondrial genome
at the me of this study. The H7 MtGenome proj-
ect is open to anyone who has tested their mtDNA
full genomic sequence with FTDNA and belongs
to H7 or one of its subclades (hps://www.fam- All members
who were not contacted inially and whose data
showed they belonged to one of the groups of in-
terest (the two idened Jewish clades and any
cluster which suggested Jewish presence) were
also issued invitaons to parcipate in the study
and quesoned as above.
In addion, for each Ashkenazi cluster found, a
sister cluster was sought for comparison among
project members without regard to ethnicity. Sis-
ter clusters were dened as two disnct branches
deriving from the same mother node in the tree.
Sequences will also be deposited in GenBank (see
Supplementary Table 1).
We decided to use relave me origins, where ap-
propriate, rather than ambiguous absolute me
Three branches with a notable Jewish constu-
ency were idened within haplogroup H7, for
a total of 89 sequences. Two of these branches,
H7c2 and H7e, have been previously idened
but not previously connected to Ashkenazi Jewish
roots. The third branch is newly reported here; it
is dened by a nucleode transion from T to C
at posion 1700 in the coding region and by two
addional mutaons (152C, 573.1C), and thus was
not idenable from inspecon of the rst con-
trol region alone. We tentavely label this clade
H7j, following standard mtDNA nomenclature
(Phylotree Build 17; Van Oven, 2015). The three
branches likely represent three dierent mater-
nal founders. In addion, two sister clades were
idened for H7c2 among the project’s parci-
pants, namely H7c1 and H7c3, both documented
branches of H7c. We did not nd any sister clades
to H7e or H7j in our data set. An overview of the
ve branches in relaon to the H7 ancestral node
is shown in Figure 1.
A total of 14 individuals belonging to newly iden-
ed H7j were found. Of these, nine agreed to par-
cipate. All nine parcipants reported Ashkenazi
Jewish ancestry on their direct maternal line, with
one nong addional possible ancient Sephardic
Jewish roots. A notable paern was observed in
this small clade in which the most frequent se-
quence was not ancestral H7j, but rather a descen-
dant branch (see Figure 1, boom branch). There
is no known posive selecon pressure because
its single change in the coding region (T11137C)
is a synonymous mutaon. The success of this
branch within H7j may instead be due to random
dri during the populaon explosion following the
severe Ashkenazi boleneck. We may be witness-
ing the in-progress disappearance of the mother
node of H7j, which is becoming less prevalent
than its daughter node, presumably an interme-
diate step before being lost enrely to history and
producing breaks in the phylogenec tree.
H7c2 and sister clades H7c1 and H7c3
A total of 25 people were found in H7c2, 17 of
which responded to the invitaon. All 17 reported
Ashkenazi Jewish ancestry on the direct maternal
line. We do not think this reects sampling bias
because public informaon available on individ-
uals who did not respond pointed to Ashkenazi
Journal of Genec Genealogy 8(1):21-34, 2016
Jewish ancestry as well. H7c2 consisted of individ-
uals from regions of Austria, Hungary, Poland, Ro-
mania, and the Pale of Selement.
Of the 25 individuals conrmed as belonging to
H7c2, a large majority (20) belonged to the an-
cestral cluster (A13959T). The remaining ve each
had a unique haplotype. This is consistent with re-
cent expansion in a large populaon, large enough
for several branches to emerge contemporane-
ously. The deepest nesng was separated by two
mutaons from the ancestral H7c2, belonging to
an individual of Hungarian Jewish ancestry (see
Figure 1).
In contrast, the sister clade H7c1 (previously es-
mated to be over 3,000 years old; Behar et al.,
2012) had a wider geographic distribuon than
Ashkenazi dominated locales, with our parci-
pants reporng ancestry from Egypt, Asia Minor,
Italy, Germany, the Brish Isles, and the Ukraine.
H7c1 is also found among the Druze of Israel
(Shlush et al., 2008). One of our parcipants re-
ported Sephardic Jewish ancestry, and the re-
maining parcipants denied any Ashkenazi Jewish
ancestry. The current distribuon of H7c1 may re-
ect populaon movements around the Mediter-
ranean during and subsequent to the Roman era.
The second sister clade H7c3 (esmated by previ-
ous researchers to be 2440 years old) was distrib-
uted mainly in Northern and Eastern Europe with
ancestry reported from Finland, Sweden, Russia,
and Poland. As with H7c1, no individuals with
Ashkenazi Jewish ancestry were reported despite
the haplogroup being found in some of the areas
heavily populated by Ashkenazi Jews, such as Gali-
cia in Poland.
The Ashkenazi Jewish H7c2 appears to be a young-
er clade than sister H7c1 with one fewer mutaon
separang it from the mother haplogroup H7c and
less rich nesng structure. H7c2 has been dated
previously to 1,735 YBP (Behar et al. 2012), young-
er than the 3000+ YBP esmate for H7c1 and
2400+ YBP for H7c3. The relavely young cluster
of H7c2, found here only in Ashkenazim (although
among mulple diverse communies), favors a lo-
cal European emergence in early Ashkenazi sele-
ment predang their geographic dispersal. In view
of the wide geographic dispersal of the mother
clade H7c in both Western Asia and Europe (es-
mated TMRCA of over 7,000 YBP; Behar et al.,
2012), and the documented presence in the Le-
vant of the daughter branch H7c1, which includes
the Druze samples and at least one individual of
Sephardic origin, a Levanne source for the pre-
cursor of H7c2 is a possibility. However, consider-
ing that the sister clade H7c3, as well as some of
the H7c1 samples, trace their ancestry to Northern
Europe, it is dicult to reach a conclusion based
on this evidence. If the absolute me esmate for
H7c2 is correct, this ming would also support a
non-European origin for the maternal ancestress
of the local Ashkenazi H7c2 mutaon, because it
dates to the early period of the Jewish diaspora
(200–300 CE; i.e., it pre-dates 650 CE) when the
vast majority of Jews were found outside of Eu-
rope (see Appendix). However, as noted, absolute
me esmates from genec mutaons rates are
problemac and cannot presently be relied upon
to disambiguate origin. Brotherton and colleagues
(2013), for example, using dated haplogroup H ge-
nomes to calculate mutaon rates, found a mu-
taon rate 45% higher than current esmates for
human mitochondria.
In contrast with H7c2 and H7j, which were found to
be exclusively Jewish, H7e included a few individ-
uals of European ancestry with no known Jewish
ancestry. H7e was also the largest of the predom-
inately Jewish clusters within H7, with 54 of the
63 individuals of self-described certain Ashkenazi
Jewish. Behar and colleagues (2012) dated H7e to
the 5th–6th Century CE, but, as with other exam-
ples noted, use of a dierent mutaon rate or a
high standard deviaon means the cluster could
either predate or postdate the crical 650 CE me
boundary. We did not idenfy any individuals car-
rying only one of the dening mutaons of H7e
(8026T and 9527T), consistent with earlier work
by Atzmon et al. (2010). H7 itself has been es-
mated to be 8890 years old (Behar et al., 2012),
many thousands of years older than H7e. Overall,
no conclusion can be drawn about the origin of
H7e from looking at the haplotypes upstream.
Journal of Genec Genealogy 8(1):21-34, 2016
Journal of Genec Genealogy 8(1):21-34, 2016
Of the 63 individuals with H7e, 31 belonged to
the ancestral cluster and carried only the den-
ing mutaons of the clade, 8026T and 9527T. In
addion, 28 of these 31 individuals were either
self-described certain Ashkenazi Jewish or were
highly likely to have Ashkenazi roots based on the
informaon provided about their direct maternal
lines. For two individuals, there wasn’t sucient
informaon to determine whether they had Ash-
kenazi roots, and one individual had no known
Ashkenazi roots. None of those belonging to the
Ashkenazi cluster were aware of Sephardic or oth-
er Jewish roots.
Ashkenazi Jewish H7e
In addion to the ancestral cluster in H7e, a num-
ber of disnct Ashkenazi clades within H7e were
found. The cluster with the greatest internal di-
versity, which we tentavely labeled H7e1, was
idened by the addional mutaon 8994A in the
coding region. All known members of H7e1 report-
ed Ashkenazi ancestry on their maternal lines. The
sequence most distant from the ancestral cluster
had three addional mutaons (Figure 2). The
deep nesng provided evidence of the longevity
of H7e among Ashkenazi Jews. An addional large
Ashkenazi cluster, tentavely labeled H7e2, was
idened by the mutaon 12651A.
In total, 84% of the samples belonging to H7e had
or highly likely had Ashkenazi Jewish roots on their
direct maternal lines. The geographic distribuon
of these individuals in the ancestral cluster en-
compassed praccally all of the countries in which
Ashkenazi Jews lived at the beginning of the 20th
Century, from Germany and Austria in Western
Europe, through Poland and the Balc states in
Central Europe, to Moldova, Ukraine and Belarus
in Eastern Europe. Furthermore, within the Ashke-
nazi subclades of H7e, disnct regional paerns of
distribuon were discernable, with disproporon-
ate numbers reporng Lithuanian ancestry (60%)
in H7e1 (8994A) and Polish ancestry (50%) in H7e2
The wide distribuon of the ancestral cluster
along with the more regional distribuon of the
subclades indicate that H7e entered the Ashkenazi
gene pool at a relavely early stage in the history
of the haplogroup. The emergence most proba-
bly occurred no later than during the 9th and 10th
centuries during the formave stages of Ashkenazi
Jewry and prior to the movement eastwards to
Central and nally to Eastern Europe.
Non-Jewish H7e
Of the 63 H7e individuals, six had no known Ash-
kenazi ancestry (~10%), including two who can
trace their ancestry back to Germany and one to
the island of Susak in Croaa. The remainder could
not trace their ancestry beyond colonial America.
Another three individuals are unlikely to have Ash-
kenazi ancestry (~5%).
A striking aspect about the non-Jewish H7e results
is that they were found to be a considerable ge-
nec distance from the ancestral cluster and sep-
arated by several mutaons (see Figure 2). One
sequence had four possible independent muta-
ons (16218T, 292.1A, 294.1T, 11890R), and two
sequences had three mutaons (2222C, 11890G,
16305G). Furthermore, these clusters did not nest
within the exisng Jewish subclades of H7e, nor
did those nearer to the ancestral cluster with no
known Jewish roots. There seems to be a clear
disncon between those belonging to the sub-
clade with Ashkenazi Jewish roots and those with-
out Ashkenazi Jewish roots, bar one member of
the ancestral cluster with no known Jewish roots
(< 4% of the ancestral cluster) The non-Jewish
samples also show greater genec diversity than
the Jewish samples.
The current work idened three clades and sev-
eral subclades of H7 as predominantly Jewish. One
of these (H7j) was previously undiscovered, and
the others (H7e, H7c2) had not previously been
idened as mainly Jewish. We focused on the Eu-
ropean haplogroup H, rarely discussed within Ash-
kenazi genecs, to gain insight into early European
Jewish maternal origins.
Journal of Genec Genealogy 8(1):21-34, 2016
Journal of Genec Genealogy 8(1):21-34, 2016
The largest group was H7e, with 63 individuals.
This reects the largest collecon of complete
H7e sequences reported to date; adding to the
previous ve sequences available on GenBank. At
least two regionally disnct subgroups were new-
ly found within H7e. The relavely large sample
enabled several paerns to be revealed: 1) The
bulk of H7e individuals have Ashkenazi maternal
origins. 2) The geographic origins of Ashkenazi
H7e encompassed all regions in which Ashkenaz-
im were found including Germany and Austria in
Western Europe, Poland and the Balc states in
Central Europe, and Moldova, Ukraine and Be-
larus in Eastern Europe, with regional subclades
apparent. 3) Some H7e sequences were found in
individuals who knew of no Jewish ancestry. 4)
The Non-Jewish sequences showed rich nesng
and several mutaonal dierences from ancestral
H7e. And, 5) the non-Jewish clusters showed no
overlap with Jewish subclades. Taken together,
these ndings strongly implicate the introgression
of a mitochondrial lineage either from or into the
Jewish gene pool that occurred early in the sele-
ment of European Jews. This was followed by no
further genec contact between the two groups.
Genec isolaon led to separate expansions, es-
pecially among the Ashkenazi as they made their
way deep into Eastern Europe.
One challenge facing research into Jewish mater-
nal lineages has been their disncveness, which
makes their origins dicult to determine. That is,
many maternal lineages found among Jewish pop-
ulaons, despite having signicant coding region
variability, are restricted solely to the Jewish sub-
group to which they are found in. In H7e, on the
other hand, we found disnct evidence of both
Ashkenazi Jewish and European non-Jewish ma-
ternal lineages with clear relaonships based on
coding region variability. Thus we can see genec
evidence of an o-speculated but rarely seen early
exchange, followed by independent development,
in the gene pool between Jewish and non-Jewish
But in which direcon was the early genec con-
tribuon? The dominance of haplogroup H as an
early European rather than Near Eastern hap-
logroup may favor the hypothesis that one woman
belonging to Haplogroup H7e converted to Juda-
ism and married into the Jewish community. The
predominance of Jewish individuals within the
ancestral cluster would, in this view, be explained
by the Ashkenazi boleneck and subsequent pop-
ulaon boom (Carmi et al., 2014) which resulted
in an inated number of Ashkenazi Jewish women
carrying the ancestral version of H7e than in the
general European populaon.
One is also tempted to speculate that the non-Jew-
ish European origin of H7e was German. This
possibility is consistent with the fact that, of the
few individuals without Jewish roots, two could
trace their distant ancestry back to Germany. In
addion, Ashkenazi Jewish history considers set-
tlement in Germany to have occurred before ex-
pansion to Eastern European regions. If this is the
case, then H7 is younger than previously thought,
because there is praccally no evidence of a Jew-
ish presence during the 7th and 8th centuries in the
Rhineland area (see Appendix).
A second possibility consistent with an older age
for H7e is a European origin in Italy or Southern
France. The Jewish presence in the Rhineland
area, and later in central Europe, is considered the
outcome of the migraon of Jews from Southern
Europe that began in the 9th and 10th centuries
(Bocini & Eckstein, 2012). The gene ow, howev-
er, could have occurred in either direcon: for ex-
ample, non-Jewish French women marrying newly
arriving Near Eastern Jewish men or Jewish wom-
en arriving to Italy from the Near East and leav-
ing the Jewish community. Origin of H7e in Italy
or Southern France would require an explanaon
for why all traces of the haplogroup have vanished
from those areas. Such an explanaon may not be
hard to nd. In general, many — perhaps most —
haplogroups have likely vanished from existence;
the unusual situaon of the Ashkenazi extreme
boleneck and subsequent populaon explosion
allowed otherwise exnguished haplogroups to
survive in select demographics.
Finally, despite the predominance of haplogroup
H in Europe and the other factors suggesng a
European origin, we cannot denively rule out
the other extreme: that the ancestress of H7e was
herself part of the Jewish community in anqui-
ty. Regardless of where geographically the wom-
en were when the mutaons of H7e arose, they
sll could have arisen in women whose ancestors
were Jewish before leaving the Near East. H7 and
other H clades could nonetheless have been in the
Near East at the right mes even if they predomi-
nately expanded in Europe. In this view, the small
number of non-Jewish individuals belonging to
H7e represents the descendants of women who
le the Jewish community relavely early on in
the history of the subclade. This would include the
German, Croaan, and Colonial American parci-
pants in our study.
The present work also uncovered a small new
clade tentavely labelled H7j and idened the
previously known H7c2 group as Ashkenazi Jew-
ish. Neither had any non-Jewish aliaon. The
small sizes of the clusters may have precluded any
minor non-Jewish presence from being detected,
the small clusters may have vanished in all but
the large Ashkenazi populaon, or the mutaons
characterizing these branches may simply have
arisen among the isolated Ashkenazi communies
while in Europe. We favor the laer hypothesis.
Regardless, it is important to note that an ancient
Near Eastern source for the precursors of H7c2 or
H7j is possible under any of the hypotheses. We
also found interesng paerns in the smaller H7j
and H7c2 clusters. One cluster contained several,
non-overlapping, shallow branches that emerged
contemporaneously, reecng a relavely new
clade in a large populaon. The other paern re-
vealed a possible in-progress vanishing of the an-
cestral group, which may soon be lost to history
and lead to missing links in the phylogenec tree.
As analysis of H7 clades illustrates, determining
the direcon of gene ow with any degree of cer-
tainty is dicult, even when sequences belonging
to non-Jewish populaons are found (as for H7e).
The problem is even greater when a mitochon-
drial lineage is restricted exclusively to Ashkenazi
Jews, as oen occurs. Consequently, it is notable
that Costa and colleagues (2013) nonetheless con-
cluded that 80% of Ashkenazi maternal ancestry
is due to the assimilaon of mtDNAs indigenous
to Europe, most likely through conversion. We feel
this conclusion is premature and goes beyond the
available evidence for several reasons: the intri-
cacies of Jewish history are oen overlooked, the
methodology of looking at the immediate ances-
tral nodes is not always conclusive, me esmates
that can be grossly inaccurate are oen relied on
too heavily, and confusion exists between where
an individual lived when a de novo mutaon arose
and that person’s origins. We provide an example
and brief elaboraon from the Costa et al., 2013
paper to illustrate. We belabor the point because
of the importance of concluding such a denive
maternal origin for the vast number of Ashkenazi
The haplogroups surveyed by Costa and col-
leagues (2013) may have arisen in Europe be-
tween the last glacial period and the Neolithic
as maintained. However, when, considering the
complex history of migraon within the Mediter-
ranean basin over the last 3,000 years, as well as
Jewish history (see Appendix), it is apparent that
where a haplogroup rst arose many thousands of
years earlier need not have any bearing on where
and when a specic disncve mitochondrial hap-
logroup rst emerged among Jewish populaons.
Furthermore, a sizeable poron of the Mediterra-
nean–Hellenisc Jewry of anquity was comprised
of converts to Judaism rather than descendants
of the Iron-Age Israelites. While the majority of
these converted in the land of Israel prior to 65 CE,
they undoubtedly included some descendants of
merchants, colonists, and troops with roots trac-
ing back to Mediterranean Europe, which could
explain some of the European admixture found
amongst the Jewish populaons descending from
the Mediterranean-Hellenisc Jewry of anquity
based in the Eastern Mediterranean.
For a specic example, consider the oen dis-
cussed haplogroups K1a1b1a and K1a1b1a1
among Ashkenazi Jews. Costa and colleagues
(2013) used maximum likelihood to esmate that
K1a1b1a dates to approximately 4,400 YBP and
K1a1b1a1 to 2,300 YBP. To place these results in
their historical perspecve, 2,300 YBP predates
the dispersal of the Jewish populaon from the
Levant to Europe, and 4,400 YBP predates by more
than 1,000 years the earliest documented menon
of the name “Israel” in historical record (the Mer-
Journal of Genec Genealogy 8(1):21-34, 2016
Journal of Genec Genealogy 8(1):21-34, 2016
neptah Stele, dated to 1209 BC). As they esmate
the parent clade K1a1b1 to be over 10K years old,
in the interim ~6,000 years between the appear-
ance of K1a1b1 and the appearance of K1a1b1a,
the maternal lineage could have migrated to and
from the Levant on numerous occasions (in a man-
ner similar to the movement paern of H7c1). As
noted earlier, prior to the Arab conquest in the 7th
century CE the Western and Eastern sides of the
Mediterranean basin were as well, if not beer,
connected to each other than the Western Med-
iterranean was to parts of Northwestern Europe.
When considering the age of the haplogroup, its
presence (however limited) among Sephardic
Jews and its apparent absence in non-Jewish pop-
ulaons (Costa et al., 2013; Behar et al., 2006) all
seem to indicate that a Levanne origin is far more
likely for K1a1b1a than a European one, regard-
less of where K1a1b1 rst originated.
Turning aenon to mtDNA mutaon rates, our
nding of early exchange between the European
and Jewish gene pools in haplogroup H mtDNA
(H7e) suggests that the rates of mutaons are
much faster than commonly assumed. They
are closer to those esmated using pedigrees.
Madrigal and colleagues (2012) calculated a
mutaon rate of 1.24 × 10−6 per site per year in
an analysis of individual family pedigrees from a
well-documented populaon in Costa Rica, a rate
three mes faster than those commonly derived
from phylogenies. The disncveness of Ashke-
nazi Jewish maternal lineages and their isolaon
from non-Jewish maternal lineages, coupled with
a rapid populaon explosion and the relavely
well-documented history of Ashkenazi Jewry,
may provide a further basis for grounding the
widely varying mutaon rates oered by dierent
Finally, we can reconsider the high degree of
European admixture (30%–60%) observed among
Ashkenazi, Sephardic, Italian, and Syrian Jews
(Atzmon et al., 2010) in autosomal DNA studies,
as well as the higher proporon of European
admixture among North African Jews compared
with non-Jewish North African populaons
(Campbell et al., 2012). Part of this clearly reects
limited more recent European admixture, hence
the elevated levels of European admixture when
comparing Ashkenazi to Sephardic Jews or Mo-
roccan to Djerban Jews. However, part undoubt-
edly reects the legacy of the Mediterranean and
the movement of peoples around the Mediter-
ranean basin long before Chrisan Southern
Europe become isolated from the Islamic Levant
and North Africa, and results from conversions to
Judaism prior to 65 CE in the Hellenisc and then
Roman Levant and North Africa.
Lile is known about the earliest days of sele-
ment of the Ashkenazi Jews in Europe. Research
into Jewish populaon genecs holds the prom-
ise of illuminang migraons and expansions
that are poorly understood due to the scarcity of
reliable historical sources. We believe we have
provided one of the clearest views of this early
period through a branch of maternally inherited
mitochondrial DNA haplogroup H that strongly
implicates gene ow between the Ashkenazi and
non-Jewish European populaons pre-dang
the Ashkenazi expansion throughout Central and
Eastern Europe. We focused on the most preva-
lent haplogroup in Europe, which also contains
subclades found almost exclusively among Ash-
kenazi Jews, to provide further insight into the
origins of the European Jewish communies. We
found gene ow within haplogroup H7, evidence
that will be benecial in assessing the origin of
other mitochondrial subclades found among
Jewish groups.
The authors gratefully acknowledge the construc-
ve criques provided by Leah Larkin, Ian Logan,
and two anonymous reviewers as well as the
assistance of Jacques Beaugrand, administrator
of the H7 MtGenome Project.
Appendix A. A Brief Consideraon of Mediterra-
nean and Jewish History
Historical consideraons in mtDNA genec stud-
ies tend to focus on prehistoric Europe because
of the ages of many haplogroups and, in parc-
ular, the last glacial maximum and its impact on
human migraons (Roostalu et al., 2007). Oen
overlooked, , however, is that following these
events many thousands of years ago, human
migraon connued unabated and, with it, the
corresponding gene ow between dierent
parts of Europe, Western Asia, and North Africa
(e.g., Brotherton et al., 2013 re Haplogroup H in
One of the most important facilitators of migra-
on between these geographical areas was the
Mediterranean. As Abulaa (2003) pointed out,
thanks to the ease of movement across the open
sea, lands far removed from each other enjoyed
vibrant trading, cultural, and polical es. Fur-
thermore, from the Mediterranean, access could
be gained to the European network of big riv-
ers, such as the Danube and the Rhine, further
facilitang the movement of goods and people
from the Mediterranean basin inland into Central
Europe. There is no doubt that this movement
around the Mediterranean basin has very an-
cient roots. Archaeological sites in Israel reveal a
Stone Age culture quite similar to that known in
the Western Mediterranean from the limestone
caves of Spain, France, and Northern Italy (Suano,
The Mycenaeans in the 14th century BCE were the
rst to start intensively traversing the Mediterra-
nean carrying trade between the Aegean and the
Levanne coastal cies, thus linking these regions
to the central Mediterranean and, on occasion,
Iberia. Permanent selements of Mycenaeans
have been idened on the coast of southern
Italy, in Sicily, and in Sardinia (Torelli, 2003). The
commercial trac of the Mediterranean through-
out the pre-Roman age was marked by colonial
selement as much as by mercanle contact.
Following the collapse of the Mycenaean empire
and the rise of classical Greece and Phoenicia,
the trade rivalry between the Greeks and Phoeni-
cians and the ensuing bale over the Mediterra-
nean trading routes between 1,000 BCE and 300
BCE led to the development of a wide ranging
network of trading selements and colonies.
Colonies in Carthage and the ring of emporia in
Libya, Motya, and Soluntum in Sicily; the harbors
in Sardinia; and the bases and trading staons
at Ibiza in the Baleric Islands, Cadiz beyond the
straits of Gibraltar, and along the Moroccan Coast
allowed the Phoenicians to dominate many of the
trade routes straddling North Africa, Iberia, and
the Levant. The Greeks as well as the Etruscans
developed rival trading routes covering much of
Southern Europe, the Adriac, the Black Sea, and
Asia Minor (Torelli, 2003).
The key period of Mediterranean unicaon
occurred, however, under the rule of imperial
Rome. For a period of roughly 800 years (300
BCE–500 CE) the whole Mediterranean was
polically unied. As Rickman (2003) stated, “it
is hardly surprising that a sea which the Romans,
and the polyglot populaons under their control
had so thoroughly made their own …. should
witness not just the circulaon of goods, but
also of people”. Military conquests during the
Republic (300–100 BCE) and the expansion of the
Roman Empire brought to the Italian peninsula
signicant economic migraon of free immigrants
as well as slaves from Gaul, Hispania, Germania,
Magna Graecia, Asia Minor, Phoenicia, Egypt, and
North Africa (Noy, 2000; Scheidel, 2004). Scheidel
(2004) esmates that around 2 million people im-
migrated to Rome just during the last two centu-
ries BCE while, according to Noy (2000), over 10%
of foreigners buried at Rome came there from
North Africa, and most were civilians rather than
associated with the military (see Killgrove, 2010,
2013). The movements of people were not just
to Rome. The names of the units staoned on
Hadrian’s Wall reveal how widely Rome recruited
its auxiliary regiments, from Spain, Gaul, Germa-
ny, the lands along the Danube, Asia Minor, Syria,
and North Africa (Vindolanda, 2016).
Jewish history is intertwined with Mediterranean
history. The formave stages of the Jewish dias-
pora occur during the period of the Mare Nos-
trum (or ‘our [Roman] sea’). There is a tendency
to confuse the Iron-Age Israelites of the 8th and
Journal of Genec Genealogy 8(1):21-34, 2016
Journal of Genec Genealogy 8(1):21-34, 2016
9th centuries BCE with the Jewish populaon liv-
ing in the Roman province of Judea nearly 1,000
years later just prior to the great revolt of 65–70
CE, however, while undoubtedly some of those
living in Judea as Jews during the 1st century CE
were the genec descendants of the inhabitants
of the ancient kingdoms of Israel and Judah,
many others were not. The four centuries fol-
lowing the Babylonian conquest of Judah in 586
BCE had seen major polical and demographic
changes taking place in the land of Israel. Faust
(2012) has persuasively shown that, based on
the archaeological evidence, Judah experienced
drasc demographic decline due to the war, sub-
sequent famine, and epidemics that followed the
conquest. Connuity in the following centuries
with the Iron Age society of Judah was limited.
There were survivors, and some of the popula-
on exiled to Babylon must have returned, but
populaon recovery in the region must have also
been triggered by new selers from neighboring
regions (Faust, 2012). Following its conquest by
Alexander the Great in 332 BCE, Judea was no
longer merely a buer state between Egypt and
Mesopotamia; it now formed the eastern edge of
what was quickly becoming a pan-Mediterranean
empire — the Roman ‘Mare Nostrum’. By 63 BCE,
Judea was a client state of Rome and by 6 CE a
Roman province.
In Goodman’s (1994) thorough research into
proselytes and proselyzing to Judaism during
the period of the Roman Empire, he concluded
that there is evidence that prior to 65 CE, con-
verts made up a signicant proporon of the
Jewish populaon and that Jews accepted as
proselytes those genles who applied to join
their number, although they did not feel com-
pelled to encourage such conversions. As exam-
ples, Goodman (1994) referred to the spread of
Jewish selement in the diaspora, the increase in
the populaon of Judea apparent from archae-
ological survey, and Josephus’ recording of the
conversion en masse of neighboring populaons
such as the Idumeans and the Ituraeans by the
Hasmonaean dynasty.
In the post-70 CE period, ambivalence by Rabbini-
cal authories towards the proselyzaon of gen-
les meant that conversion to Judaism was far
less common, although there is some evidence
of proselytes to Judaism all the way through
into the medieval period (Goodman, 1994). This
was especially true aer the failed Bar Kokhba
rebellion during Hadrian’s rule and the passage of
legislaon by Hadrian and his successors against
the circumcision of non-Jews, the special Jewish
tax (the scus Judaicus), and a series of Roman
laws in the 4th and 5th centuries prohibing
conversion to Judaism, parcularly by Chrisans.
Furthermore, as Goodman (1994) pointed out,
some conversions to Judaism probably took place
to facilitate marriage. Considering the patriar-
chal nature of both Jewish and Roman sociees,
as well as the prohibion on circumcision that
prevented men (but not women) from convert-
ing, many of the converts to Judaism to facilitate
marriage were likely women.
How many of these conversions would have
taken place in Europe? As can be seen in Table 1
based on the esmates of Bocini and Eckstein
(2012), prior to 65 CE the majority of the Jewish
populaon throughout the Middle East and the
Mediterranean basin were located in the lands
of Israel, Mesopotamia, Persia, and North Afri-
ca (mainly Egypt), while the number of Jews in
Western Europe was relavely small and by 650
CE was negligible (~1,000). Thus the vast majority
of conversion to Judaism during this period must
have occurred outside of Europe in the Levant,
Egypt, and Mesopotamia.
Furthermore, in a detailed study by Toch (2005)
of Jews in Europe between 500–1050 CE, he
concluded that between the mid-7th and mid-8th
centuries, no source menons Jews in Frankish
lands (now France and Germany). Only in the 8th
and 9th centuries was there evidence of growing
numbers of Jews in the South of France, while in
the 9th and early 10th centuries, brief hints aest
to inerant merchants in Germany. Toch (2005),
therefore, concluded that no connuity could be
assumed between the Jews of the Roman Empire
and the Ashkenazi Jewish communies of the
Middle Ages.
From a genec perspecve, based on this histori-
cal overview, maternal lineages restricted to Jew-
ish populaons that pre-date 650 CE are highly
unlikely to have originated in either Western or
Eastern Europe, given the miniscule numbers of
Jews in these regions during this period.
Journal of Genec Genealogy 8(1):21-34, 2016
Table 1. Jewish populaon esmates in 65 CE and 650 CE (as per Bocini and Eckstein, 2012).
Region c. 65 CE c. 650 CE
Land of Israel 2,500,000 100,000
Mesopotamia and Persia (including the Arabian Peninsula) 1,000,000 700,000–900,000
North Africa (mainly Egypt) 1,000,000 4,000
Syria and Lebanon 200,000–400,000 5,000
Asia Minor and the Balkans 200,000–400,000 40,000
Western Europe (including Italy, France Germany, and Iberia) 100,000–200,000 1,000
Eastern Europe
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... was searched for potential participants. The large predominately United States customer database at Family Tree DNA (FTDNA) has proved to be a valuable research resource [4,5,[14][15][16][17]. Sequences from the control region of mtDNA harboring both the variant that identifies T2e, i.e. 16153A in the first hypervariable segment (HVS1), and the variant that identifies most of the T2e1 branch, 41T in the second hypervariable segment (HVS2), were selected for further consideration. ...
... Instead, the ancestral 2308G sequence is likely the only surviving example currently to be found, originally perhaps even from Spain, with the mother node now largely extinguished in favor of daughter node expansion. (Ashkenazim H7 provides another example of this phenomenon [16]). It instead likely reflects traces of dispersal of T2e1a as far as Eastern Europe, like the previously elaborated Jewish 9181G branch [5]. ...
... Estimates of variant rates based instead on pedigrees have found them at least three times faster [27], which would put the timeline back in the range of historical Jewish diaspora [17]. Time estimates with sufficient precision are currently not possible [4,5,16]. The best evidence for an older emergence of T2e1 that predates the Israelites comes from a sample of T2e1 found in skeletal remains in Germany estimated from the mid Neolithic about 5300 YBP [28]. ...
... We used those sequences to estimate relative frequencies of the haplogroups observed amongst AJ of different European origin (Fig. 1B) and contrasted them to the distribution of the same haplogroups amongst corresponding European populations (Fig. 1C). Both our data and data from literature show that all haplogroups established for the studied individuals correspond to lineages found in Ashkenazim, while being rare or missing in other European populations [5,6,[9][10][11][12]. According to earlier studies, the ancestors of modern AJ are considered to be mainly of Central and Eastern European descent [9], with some evidence also pointing towards the Near East. ...
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Six million Jews were killed by Nazi Germany and its collaborators during World War II. Archaeological excavations in the area of the death camp in Sobibór, Poland, revealed ten sets of human skeletal remains presumptively assigned to Polish victims of the totalitarian regimes. However, their genetic analyses indicate that the remains are of Ashkenazi Jews murdered as part of the mass extermination of European Jews by the Nazi regime and not of otherwise hypothesised non-Jewish partisan combatants. In accordance with traditional Jewish rite, the remains were reburied in the presence of a Rabbi at the place of their discovery.
... Increased interest in genetic genealogy and self-testing within the general population has allowed researchers access to a larger and more in-depth datasets. Genetic genealogy results are increasingly being used as a scientific resource [2,[7][8][9][10][11][12]. Furthermore, the development of Next-Generation Sequencing (NGS) technologies focused on the Y chromosome such as Family Tree DNA's "Big Y", FullGenomes Corporation's "Y-Elite", Whole-Genomic Sequencing (WGS) like FullGenomes Corporation's "Genome Guide" and such as undertaken in the 1000 Genomes Project [13,14] have allowed for collaborative research between professionals and citizens [15][16][17]. ...
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This article aims at researching the evolution of J-Z640 using an interdisciplinary approach in order to clarify the leading historical and anthropological events that shaped this particular branch of the human Y chromosome. We compiled a STR (short tandem repeat) and SNP (single nucleotide polymorphism) dataset of 145 known or predicted J-Z640 samples among the customers of Family Tree DNA and Full Genomes Corporation, as well as publicly available samples. Amongst these, we analyzed the results of 41 samples that had undergone Next-Generation Sequencing (NGS) and 32 samples that had undergone SNP testing using Sanger Sequencing. From this data, we constructed a J-Z640 phylogenetic tree that was dated using the method. Our data revealed that Haplogroup J-Z640 is a Y chromosome lineage found most notably, in several minority groups within the Near East such as the Samaritans, Druze, Armenians and Jews. J-Z640 originated during the Bronze Age, most likely in the Levant. During the Bronze Age the haplogroup rapidly expanded with multiple ancient branches surviving to the present, evidencing population growth. Further population expansion, and contraction, was also observed in later periods. Based on its geographic dispersal and age of the haplogroup and its subclades, the founder population most likely belonged to Canaanites found in the Levant. Following the collapse of the late Bronze age system, from the Iron Age onwards there followed a period of “differentiation by culture”, with many of the ancient branches surviving to the present separated along ethno-religious lines.
... Looking for possible mtDNA matches has involved the construction of a detailed ancestral tree (see Lineage DNB below), contacting matches in the commercial databases, joining the Richard III Society and giving interviews to the press. The methods are very similar to those described in a recent paper 28 which gives details of a search for people in haplogroup H7. ...
King Richard III has been a controversial figure for centuries and the finding of his skeletal remains under a car park in 2012 has only raised his profile. But what is generally unknown is that he belongs to an exceedingly uncommon mitochondrial haplogroup, J1c2c3. Two maternal descendants of his extended family have already been traced by orthodox genealogical methods, but no other possible descendants have been found in Europe. In our study we have identified a total of seven lineages containing people in haplogroup J1c2c3 through the use of direct-to-customer DNA testing. All the lineages are American, and unfortunately they do not link back to King Richard III, as yet. However, we do suggest that because of the small size of haplogroup J1c2c3 the people we have identified may well be descendants of the immediate family of King Richard III.
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We report on two of the oldest mitochondrial DNA clusters in existence with Jewish affiliation. Both are in haplogroup T2e1. Four unrelated individuals from the Mexico mtDNA project were found to have the control region mutations that characterize a Sephardic signature previously reported (motif 16114T-16192T within T2e). Full genomic sequencing found the identical coding region mutations as Sephardic individuals which provides genetic evidence for founders of Northern Mexico that were both female and Sephardic Jewish. This is in contrast to a more common finding of European male, but local female founders and additionally lends biological support to anecdotes and historical reports of Crypto-Jewish founding of the Coahuila, Nuevo Leon, and Tamaulipas regions of Mexico and influx to Southern Texas, USA. The haplotype is nested in an old tree with mutations at positions 2308 and 15499, presently of uncertain geographic origin. The second cluster, a Bulgarian Sephardic founding lineage (9181G within T2e) previously reported, was found here in a population of largely Americans of European descent, but only among Jewish individuals. The non-synonymous mutation in ATPase 6 was found among both Ashkenazi and Sephardic Jews from diverse regions of Czech Republic, Lithuania, the Netherlands, Poland, and Romania. Full genomic sequencing found great coding region variability with several haplotypes and suggested a Near East origin at least 3000 years old. This predates the split between Jewish groups, but more recent admixture between Sephardim and Ashkenazim cannot be ruled out. Together the two Jewish-affiliated clusters account for all the genetic distance found in branch T2e1 and much of T2e. The findings suggest reexamination of the origins of mitochondrial DNA haplogroup T2e as Levantine or early back migration to the Near East. New subclades of T2e are identified.
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The early Middle Ages saw the establishment of Jewish life in major parts of Europe. In this period the Mediterranean–Hellenistic Jewry of antiquity separated and developed into Byzantine–southern Italian, Roman, Catalan-southern French and Arabic–Sicilian branches. During the later part of the period Ashkenazic (north-western and northern European) and Sephardic (Iberian) Jewry came into being as distinctive entities very different from earlier patterns. There were (probably) also some Jews in eastern Europe, of which almost nothing is known but a vague association with Byzantium. Jews held to a common creed yet differed in language, religious custom and ritual, social organisation, occupations and legal standing, a fact that militates against easy historical inferences from an ostensibly fixed character. At different times and to differing degrees, they were under the influence of the centres of gravity – demographic, religious, intellectual – of Jewish life in the Middle East, especially Palestine and Babylonia. Compared to these, throughout most of the period under consideration, European Jews in numbers or intellectual creativity were yet barely remarkable. None the less, modern historians have usually accorded them a significance out of proportion. In this they faithfully followed medieval churchmen, to whom Jews presented a challenge far in excess of their actual presence and impact.
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Haplogroup H dominates present-day Western European mitochondrial DNA variability (440%), yet was less common (B19%) among Early Neolithic farmers (B5450 BC) and virtually absent in Mesolithic hunter-gatherers. Here we investigate this major component of the maternal population history of modern Europeans and sequence 39 complete haplogroup H mitochondrial genomes from ancient human remains. We then compare this 'real-time' genetic data with cultural changes taking place between the Early Neolithic (B5450 BC) and Bronze Age (B2200 BC) in Central Europe. Our results reveal that the current diversity and distribution of haplogroup H were largely established by the Mid Neolithic (B4000 BC), but with substantial genetic contributions from subsequent pan-European cultures such as the Bell Beakers expanding out of Iberia in the Late Neolithic (B2800 BC). Dated haplogroup H genomes allow us to reconstruct the recent evolutionary history of haplogroup H and reveal a mutation rate 45% higher than current estimates for human mitochondria.
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The Ashkenazi Jewish (AJ) population is a genetic isolate close to European and Middle Eastern groups, with genetic diversity patterns conducive to disease mapping. Here we report high-depth sequencing of 128 complete genomes of AJ controls. Compared with European samples, our AJ panel has 47% more novel variants per genome and is eightfold more effective at filtering benign variants out of AJ clinical genomes. Our panel improves imputation accuracy for AJ SNP arrays by 28%, and covers at least one haplotype in ≈67% of any AJ genome with long, identical-by-descent segments. Reconstruction of recent AJ history from such segments confirms a recent bottleneck of merely ≈350 individuals. Modelling of ancient histories for AJ and European populations using their joint allele frequency spectrum determines AJ to be an even admixture of European and likely Middle Eastern origins. We date the split between the two ancestral populations to ≈12–25 Kyr, suggesting a predominantly Near Eastern source for the repopulation of Europe after the Last Glacial Maximum.
In 70 CE, the Jews were an agrarian and illiterate people living mostly in the Land of Israel and Mesopotamia. By 1492 the Jewish people had become a small group of literate urbanites specializing in crafts, trade, moneylending, and medicine in hundreds of places across the Old World, from Seville to Mangalore. What caused this radical change?The Chosen Fewpresents a new answer to this question by applying the lens of economic analysis to the key facts of fifteen formative centuries of Jewish history. Maristella Botticini and Zvi Eckstein show that, contrary to previous explanations, this transformation was driven not by anti-Jewish persecution and legal restrictions, but rather by changes within Judaism itself after 70 CE--most importantly, the rise of a new norm that required every Jewish male to read and study the Torah and to send his sons to school. Over the next six centuries, those Jews who found the norms of Judaism too costly to obey converted to other religions, making world Jewry shrink. Later, when urbanization and commercial expansion in the newly established Muslim Caliphates increased the demand for occupations in which literacy was an advantage, the Jews found themselves literate in a world of almost universal illiteracy. From then forward, almost all Jews entered crafts and trade, and many of them began moving in search of business opportunities, creating a worldwide Diaspora in the process. The Chosen Fewoffers a powerful new explanation of one of the most significant transformations in Jewish history while also providing fresh insights to the growing debate about the social and economic impact of religion.