Genetic studies on Jews

Recent studies have been conducted on a large number of genes, homologous chromosomes or autosomes (all chromosomes except chromosomes X and Y). A 2009 study was able to genetically identify individuals with full or partial Ashkenazi Jewish ancestry.[8] In August 2012, Dr. Harry Ostrer in his book Legacy: A Genetic History of the Jewish People, summarized his and other work in genetics of the last 20 years, and concluded that all major Jewish groups share a common Middle Eastern origin.[9] Ostrer also refuted the Khazar theory of Ashkenazi ancestry.[10] Citing autosomal DNA studies, Nicholas Wade estimates that "Ashkenazic and Sephardic Jews have roughly 30 percent European ancestry, with most of the rest from the Middle East." He further noticed that "The two communities seem very similar to each other genetically, which is unexpected because they have been separated for so long." Concerning this relationship he points to Atzmon's conclusions that "the shared genetic elements suggest that members of any Jewish community are related to one another as closely as are fourth or fifth cousins in a large population, which is about 10 times higher than the relationship between two people chosen at random off the streets of New York City".[11] Concerning North African Jews, autosomal genetic analysis in 2012 revealed that North African Jews are genetically close to European Jews. This finding "shows that North African Jews date to biblical-era Israel, and are not largely the descendants of natives who converted to Judaism,"[12] Y DNA studies examine various paternal lineages of modern Jewish populations. Such studies tend to imply a small number of founders in an old population whose members parted and followed different migration paths.[13] In most Jewish populations, these male line ancestors appear to have been mainly Middle Eastern. For example, Ashkenazi Jews share more common paternal lineages with other Jewish and Middle Eastern groups than with non-Jewish populations in areas where Jews lived in Eastern Europe, Germany and the French Rhine Valley. This is consistent with Jewish traditions in placing most Jewish paternal origins in the region of the Middle East.[14][3]

A study conducted in 2013 found no evidence of a Khazar origin for Ashkenazi Jews and suggested that "Ashkenazi Jews share the greatest genetic ancestry with other Jewish populations, and among non-Jewish populations, with groups from Europe and the Middle East. No particular similarity of Ashkenazi Jews with populations from the Caucasus is evident, particularly with the populations that most closely represent the Khazar region. In this view, analysis of Ashkenazi Jews, together with a large sample from the region of the Khazar Khaganate, corroborates earlier results that Ashkenazi Jews derive their ancestry primarily from populations of the Middle East and Europe, that they possess considerable shared ancestry with other Jewish populations, and that there is no indication of a significant genetic contribution either from within or from north of the Caucasus region."[15]

In 2016, together with R. Das, P. Wexler and M. Pirooznia, Elhaik advanced the view that the first Ashkenazi populations to speak the Yiddish language came from areas near four villages in Eastern Turkey along the Silk Road whose names derived from the word "Ashkenaz", arguing that Iranian, Greek, Turkish, and Slav populations converted on that travel route before moving to Khazaria, where a small-scale conversion took place.[16][17] The study was dismissed by Sergio DellaPergola as a "falsification", noting it failed to include Jewish groups such as the Italkim and Sephardic Jews, to whom Ashkenazi Jews are closely related genetically. Shaul Stampfer, a professor of Soviet and East European Jewry at the Hebrew University, called Elhaik's research "basically nonsense". Elhaik replied that the DNA of non-Ashkenazic Jews would not affect the origin of DNA hypothesized for the former.[18] Prof. Dovid Katz, founder of Vilnius University's Yiddish Institute criticized the study's linguistic analysis. “The authors have melded accurate but contextually meaningless genetic correlations with laughable linguistic theories that now proliferate, sadly, as a consequence of a much weakened Yiddish academic environment internationally ... there is not a single word or sound in Yiddish that comes from Iranian or Turkish".[19] In joint study published in 2016 by Genome Biology and Evolution, Pavel Flegontov from Department of Biology and Ecology, Faculty of Science, University of Ostrava, Czech Republic, A.A. Kharkevich Institute of Linguistics, Russian Academy of Sciences, Moscow, Mark G. Thomas from Research Department of Genetics, Evolution and Environment, University College London, UK, Valentina Fedchenko from Saint Petersburg State University, and George Starostin from Russian State University for the Humanities, dismissed both the genetic and linguistic components of Elhaik et al. study arguing that "GPS is a provenancing tool suited to inferring the geographic region where a modern and recently unadmixed genome is most likely to arise, but is hardly suitable for admixed populations and for tracing ancestry up to 1000 years before present, as its authors have previously claimed. Moreover, all methods of historical linguistics concur that Yiddish is a Germanic language, with no reliable evidence for Slavic, Iranian, or Turkic substrata." The authors concluded:

"In our view, Das and co-authors have attempted to fit together a marginal and unsupported interpretation of the linguistic data with a genetic provenancing approach, GPS, that is at best only suited to inferring the most likely geographic location of modern and relatively unadmixed genomes, and tells nothing of population history and origin."[20]

The authors, in a non peer-reviewed response, defended the methodological adequacy of their approach.[21] In 2016 Elhaik having reviewed the literature searching for a ‘Jüdische Typus’ argued that there is no genomic hallmark for Jewishness. While he allows that in the future it is possible that a ‘Jewish’ marker may turn up, so far, in his view, Jewishness turns out to be socially defined (a socionome), determined by non-genetic factors.[22] On 31 October 2016 a corrigendum to the initial GPS paper by Elhaik et al. 2014 was published in Nature Communications. The GPS tool, remained freely available on the lab website of Dr. Tatiana Tatarinova, but as of December 2016 the link is broken. In 2017, the same authors further supported a non-Levantine origin of Ashkenazi Jews claiming that "Overall, the combined results (of linguistics study and GPS tool) are in a strong agreement with the predictions of the Irano-Turko-Slavic hypothesis and rule out an ancient Levantine origin for AJs, which is predominant among modern-day Levantine populations (e.g., Bedouins and Palestinians)."[23] Elhaik's and Das' work was among others, strongly criticized by Marion Aptroot from University of Düsseldorf, who in the study published by Genome Biology and Evolution claimed that "Das et al. create a narrative based on genetic, philological and historical research and state that the findings of the three disciplines support each other...Incomplete and unreliable data from times when people were not counted regardless of sex, age, religion or financial or social status on the one hand, and the dearth of linguistic evidence predating the 15th century on the other, leave much room for conjecture and speculation. Linguistic evidence, however, does not support the theory that Yiddish is a Slavic language, and textual sources belie the thesis that the name Ashkenaz was brought to Eastern Europe directly from a region in the Near East. Although the focus and methods of research may be different in the humanities and the sciences, scholars should try to account for all evidence and observations, regardless of the field of research. Seen from the standpoint of the humanities, certain aspects of the article by Das et al. fall short of established standards".[24]

A 2020 study on remains from Bronze Age southern Levantine (Canaanite) populations suggests a significant degree of genetic continuity both in Jewish groups (including Mizrahi, Ashkenazi, and Sephardic Moroccan Jews) and non-Jewish Arabic-speaking Levantine populations (such as Lebanese, Druze, Palestinians, and Syrians) from the populations of the Bronze Age Levant, suggesting that the aforementioned groups derive at least about half or more of their ancestry from Canaanite/Bronze Age Levantine populations (over half in the Ashkenazi, with the remaining 41% of their ancestry being European, and a larger majority in Mizrahi Jewish, Moroccan Jewish and Levantine Arabic-speaking groups).[25][26]

In 1992 G. Lucotte and F. David were the first genetic researchers to have documented a common paternal genetic heritage between Sephardi and Ashkenazi Jews.[27][28] Another study published just a year later suggested the Middle Eastern origin of Jewish paternal lineages.[29]

In 2000, M. Hammer, et al. conducted a study on 1,371 men and definitively established that part of the paternal gene pool of Jewish communities in Europe, North Africa and Middle East came from a common Middle East ancestral population. They suggested that most Jewish communities in the Diaspora remained relatively isolated and endogamous compared to non-Jewish neighbor populations.[13][5][30]

Investigations made by Nebel et al.[14] on the genetic relationships among Ashkenazi Jews, Kurdish and Sephardi (North Africa, Turkey, Iberian Peninsula, Iraq and Syria) indicate that Jews are more genetically similar to groups in the northern Fertile Crescent (Kurds, Turks and Armenians) than their Arab neighbors, and suggest that some of this difference might be due to migration and admixture from the Arabian peninsula during the last two millennia (into certain current Arabic-speaking populations). Considering the timing of this origin, the study found that "the common genetic Middle Eastern background (of Jewish populations) predates the ethnogenesis in the region and concludes that the Y chromosome pool of Jews is an integral part of the genetic landscape of Middle East.[14]

Lucotte et al. 2003 study found that (Oriental, Sephardic, Ashkenazic Jews and Lebanese and Palestinians), "seem to be similar in their Y-haplotype patterns, both with regard to the haplotype distributions and the ancestral haplotype VIII frequencies." The authors stated in their findings that these results confirm similarities in the Y-haplotype frequencies of this Near-Eastern populations, sharing a common geographic origin."[31]

In a study of Israeli Jews from four different groups (Ashkenazi Jews, Kurdish Jews, North African Sephardi Jews, and Iraqi Jews) and Palestinian Muslim Arabs, more than 70% of the Jewish men and 82% of the Arab men whose DNA was studied, had inherited their Y chromosomes from the same paternal ancestors, who lived in the region within the last few thousand years. "Our recent study of high-resolution microsatellite haplotypes demonstrated that a substantial portion of Y chromosomes of Jews (70%) and of Palestinian Muslim Arabs (82%) belonged to the same chromosome pool."[32] All Jewish groups were found to be genetically closer to each other than to Palestinians and Muslim Kurds. Kurdish, North African Sephardi, and Iraqi Jews were found to be genetically indistinguishable while slightly but significantly differing from Ashkenazi Jews. In relation to the region of the Fertile Crescent, the same study noted; "In comparison with data available from other relevant populations in the region, Jews were found to be more closely related to groups in the north of the Fertile Crescent (Kurds, Turks, and Armenians) than to their Arab neighbors", which the authors suggested was due to migration and admixture from the Arabian Peninsula into certain current Arabic-speaking populations during the period of Islamic expansion.[14]

Approximately 35% to 43% of Jewish men are in the paternal line known as haplogroup J[lower-alpha 1] and its sub-haplogroups. This haplogroup is particularly present in the Middle East and Southern Europe.[33] 15% to 30% are in haplogroup E1b1b,[lower-alpha 2] (or E-M35) and its sub-haplogroups which is common in the Middle East, North Africa, and Southern Europe.

Studies of mitochondrial DNA of Jewish populations are more recent and are still debatable.[61][lower-alpha 8] The maternal lineages of Jewish populations, studied by looking at mitochondrial DNA, are generally more heterogeneous.[61] Scholars such as Harry Ostrer and Raphael Falk believe this may indicate that many Jewish males found new mates from European and other communities in the places where they migrated in the diaspora after fleeing ancient Israel.[62]

According to Thomas et al. in 2002, a number of Jewish communities reveal direct-line maternal ancestry originating from a few women. This was seen in independently founded communities in different geographic areas. What they shared was limited genetic additions later on the female side. Together, this is described as the founder effect. Those same communities had diversity in the male lines that was similar to the non-Jewish population.[63] Two studies in 2006 and 2008 suggested that about 40% of Ashkenazi Jews originate maternally from just four female founders who were likely of Near-Eastern origin, while the populations of Sephardi and Mizrahi Jewish communities "showed no evidence for a narrow founder effect".[64][61]

With the exception of Ethiopian Jews and Indian Jews, it has been argued that all of the various Jewish populations have components of mitochondrial genomes that were of Middle Eastern origin.[65][5] In 2013, however, Richards et al. published work suggesting that an overwhelming majority of Ashkenazi Jewish maternal ancestry, estimated at "80 percent of Ashkenazi maternal ancestry comes from women indigenous to Europe, and [only] 8 percent from the Near East, with the rest uncertain", suggesting that Jewish males migrated to Europe and took new wives from the local population, and converted them to Judaism, though some geneticists, such as Doron Behar, have expressed disagreement with the study's conclusions.[66] Another study by Eva Fernandez and her colleagues argues that the K lineages (claimed to be European in origin by Richards et al.) in Ashkenazi Jews might have an ancient Near Eastern source.[67]

Reflecting on previous mtDNA studies carried out by Behar, Atzmon et al. conclude that all major Jewish population groups are showing evidence for founder females of Middle Eastern origin with coalescence times >2000 years.[65] A 2013 study by Richards et al., based on a much larger sample base, drew differing conclusions, namely, that the Mt-DNA of Ashkenazi Jews originated among southern European women, where Diaspora communities had been established centuries before the fall of the Second Temple in 70 CE.[68] A 2014 study by Fernandez et al. found that Ashkenazi Jews display a frequency of haplogroup K which suggests an ancient Near Eastern origin, stating that this observation clearly contradicts the results of the study led by Richards which suggested a predominantly European origin for the Ashkenazi community's maternal lines. However, the authors of the 2014 study also state that definitively answering the question of whether this group was of Jewish origin rather than the result of a Neolithic migration to Europe would require the genotyping of the complete mtDNA in ancient Near Eastern populations.[67]

These studies focus upon autosomal chromosomes, the 22 homologous or autosomes (non sex chromosomes), rather than on the direct paternal or maternal lines. The technology has changed rapidly and so older studies are different in quality to newer ones.

An initial study conducted in 2001 by Noah Rosenberg and colleagues on six Jewish populations (Poland, Libya, Ethiopia, Iraq, Morocco, Yemen) and two non-Jewish populations (Palestinians and Druze) showed that while the eight groups had genetic links to each other, the Jews of Libya have a distinct genetic signature related to their genetic isolation and possible admixture with Berber populations.[77][lower-alpha 10] This same study suggested a close relationship between the Jews of Yemen and those of Ethiopia.[77]

A 2006 study by Seldin et al. used over five thousand autosomal SNPs to demonstrate European genetic substructure. The results showed "a consistent and reproducible distinction between 'northern' and 'southern' European population groups". Most northern, central, and eastern Europeans (Finns, Swedes, English, Irish, Germans, and Ukrainians) showed >90% in the 'northern' population group, while most individual participants with southern European ancestry (Italians, Greeks, Portuguese, Spaniards) showed >85% in the 'southern' group. Both Ashkenazi Jews as well as Sephardic Jews showed >85% membership in the "southern" group. Referring to the Jews clustering with southern Europeans, the authors state the results were "consistent with a later Mediterranean origin of these ethnic groups".[78]

A 2007 study by Bauchet et al. found that Ashkenazi Jews were most closely clustered with Arabic North African populations when compared to the global population of that study. In the European structure analysis, they share genetic similarities with Greeks and Sicilians, reflecting their east Mediterranean origins.[79]

A 2008 study by Price et al. sampled Southern Italians, Jews and other Europeans, and isolated the genetic markers that are most accurate for distinguishing between European groups, achieving results comparable to those from genome-wide analyses. It mines much larger datasets (more markers and more samples) to identify a panel of 300 highly ancestry-informative markers which accurately distinguish not just northwest and southeast European, but also Ashkenazi Jewish ancestry from Southern Europeans.[80]

A 2008 study by Tian et al. provides an additional example of the same clustering pattern, using samples and markers similar to those in their other study. European population genetic substructure was examined in a diverse set of >1,000 individuals of European descent, each genotyped with >300 K SNPs. Both STRUCTURE and principal component analyses (PCA) showed the largest division/principal component (PC) differentiated northern from southern European ancestry. A second PC further separated Italian, Spanish, and Greek individuals from those of Ashkenazi Jewish ancestry as well as distinguishing among northern European populations. In separate analyses of northern European participants other substructure relationships were discerned showing a west to east gradient.[81]

A 2009 study by Goldstein et al. shows that it is possible to predict full Ashkenazi Jewish ancestry with 100% sensitivity and 100% specificity, although the exact dividing line between a Jewish and non-Jewish cluster will vary across sample sets which in practice would reduce the accuracy of the prediction. While the full historical demographic explanations for this distinction remain to be resolved, it is clear that the genomes of individuals with full Ashkenazi Jewish ancestry carry an unambiguous signature of their Jewish ancestral DNA, and this seems more likely to be due to their specific Middle Eastern ancestry than to inbreeding. The authors note that there is almost perfect separation along PC 1, and, they note that most of the non-Jewish Europeans who are closest to the Jews on this PC are of Italian or Eastern Mediterranean origin.[8]

In a 2009 study by Kopelman et al., four Jewish groups, Ashkenazi, Turkish, Moroccan and Tunisian, were found to share a common origin from the Middle East, with more recent admixture that has resulted in "intermediate placement of the Jewish populations compared to European and Middle Eastern populations". The authors found that the "most similar to the Jewish populations is the Palestinian population". The Tunisian Jews were found to be distinct from three other Jewish populations, which suggests, according to the authors, a greater genetic isolation and/or a significant local Berber ancestry, as in the case of Libyan Jews. Concerning the theory of Khazar ancestry in Ashkenazi Jews, the authors found no direct evidence. Although they did find genetic similarities between Jews, especially Ashkenazi Jews, and the Adyghe people, a group from the Caucasus, whose region was formerly occupied by the Khazars, the Adyghe, living on the edge of geographical Europe, are more genetically related to Middle Easterners, including Palestinians, Bedouin, and non-Ashkenazi Jews, than to Europeans.[5][82]

Another study of L. Hao et al.[65] studied seven groups of Jewish populations with different geographic origin (Ashkenazi, Italian, Greek, Turkish, Iranian, Iraqi, and Syrian) and showed that the individuals all shared a common Middle Eastern background, although they were also genetically distinguishable from each other. In public comments, Harry Ostrer, the director of the Human Genetics Program at NYU Langone Medical Center, and one of the authors of this study, concluded, "We have shown that Jewishness can be identified through genetic analysis, so the notion of a Jewish people is plausible."[65]

A genome-wide genetic study carried out by Need et al. and published in 2009 showed that "individuals with full Jewish ancestry formed a clearly distinct cluster from those individuals with no Jewish ancestry." The study found that the Jewish cluster examined, fell between that of Middle Eastern and European populations. Reflecting on these findings, the authors concluded, "It is clear that the genomes of individuals with full Ashkenazi Jewish ancestry carry an unambiguous signature of their Jewish heritage, and this seems more likely to be due to their specific Middle Eastern ancestry than to inbreeding."[83]

The current study extends the analysis of European population genetic structure to include additional southern European groups and Arab populations. While the Ashkenazi are clearly of southern origin based on both PCA and STRUCTURE studies, in this analysis of diverse European populations, this group appears to have a unique genotypic pattern that may not reflect geographic origins.[84]

In June 2010, Behar et al. "shows that most Jewish samples form a remarkably tight subcluster with common genetic origin, that overlies Druze and Cypriot samples but not samples from other Levantine populations or paired Diaspora host populations. In contrast, Ethiopian Jews (Beta Israel) and Indian Jews (Bene Israel and Cochini) cluster with neighboring autochthonous populations in Ethiopia and western India, respectively, despite a clear paternal link between the Bene Israel and the Levant.".[4][11] "The most parsimonious explanation for these observations is a common genetic origin, which is consistent with an historical formulation of the Jewish people as descending from ancient Hebrew and Israelite residents of the Levant."[4] The authors say that the genetic results are concordant "with the dispersion of the people of ancient Israel throughout the Old World".[4] Regarding the samples he used, Behar says, "Our conclusion favoring common ancestry (of Jewish people) over recent admixture is further supported by the fact that our sample contains individuals that are known not to be admixed in the most recent one or two generations."[4]

A study led by Harry Ostrer published on 11 June 2010, found close links between Ashkenazi, Sephardi, and Mizrahi Jews, and found them to be genetically distinct from non-Jews. In the study, DNA from the blood of 237 Jews and about 2,800 non-Jews was analyzed, and it was determined how closely related they were through IBD. Individuals within the Ashkenazi, Sephardi, and Mizrahi groups shared high levels of IDB, roughly equivalent to that of fourth or fifth cousins. All three groups shared many genetic features, suggesting a common origin dating back more than 2,000 years. The study did find that all three Jewish groups did show various signs of admixture with non Jews, with the genetic profiles of Ashkenazi Jews indicating between 30% and 60% admixture with Europeans, although they clustered more closely with Sephardi and Mizrahi Jews.[85]

In July 2010, Bray et al., using SNP microarray techniques and linkage analysis,[86] "confirms that there is a closer relationship between the Ashkenazim and several European populations (Tuscans, Italians, and French) than between the Ashkenazim and Middle Eastern populations," and that European "admixture is considerably higher than previous estimates by studies that used the Y chromosome." They add their study data "support the model of a Middle Eastern origin of the Ashkenazim population followed by subsequent admixture with host Europeans or populations more similar to Europeans," and that their data imply that modern Ashkenazi Jews are perhaps more similar to Europeans than modern Middle Easterners. The level of admixture with European population was estimated between 35% and 55%. The study assumed Druze and Palestinian Arabs populations to represent the reference to world Jewry ancestor genome. With this reference point, the linkage disequilibrium in the Ashkenazi Jewish population was interpreted as "matches signs of interbreeding or 'admixture' between Middle Eastern and European populations". Also, in their press release, Bray stated: "We were surprised to find evidence that Ashkenazi Jews have higher heterozygosity than Europeans, contradicting the widely-held presumption that they have been a largely isolated group". The authors said that their calculations might have "overestimated the level of admixture" in case that the true Jewish ancestors were genetically closer to Southern Europeans than to Druze and Palestinian Arabs. They predict that using the non-Ashkenazi Jewish Diaspora populations as reference for a world Jewry ancestor genome would "underestimate the level of admixture" but that "however, using the Jewish Diaspora populations as the reference Jewish ancestor will naturally underestimate the true level of admixture, as the modern Jewish Diaspora has also undergone admixture since their dispersion.[87][86]

Zoossmann-Diskin (2010) argues, that based upon the analysis of X chromosome and seventeen autosomal markers, Eastern European Jewish populations and Jewish populations from Iran, Iraq and Yemen, do not have the same genetic origins. In particular, concerning Eastern European Jews, he believes the evidence points to a dominant amount of southern European, and specifically Italian, ancestry, which he argues is probably a result of conversions during the Roman empire. Concerning the similarity between Sephardi and Ashkenazi Jews, he argues that the reasons are uncertain, but that it is likely to be caused by Sephardic Jews having "Mediterranean" ancestry also, like the Ashkenazi Jews. Concerning mitochondrial DNA, and particularly Y DNA, he accepts that there are superficial signs of some Middle Eastern ancestry among Ashkenazi Jews, but he argues that this can be ignored as it may have come from a small number of ancestors.[6]

An autosomal DNA study carried out in 2010 by Atzmon et al. examined the origin of Iranian, Iraqi, Syrian, Turkish, Greek, Sephardic, and Ashkenazi Jewish communities. The study compared these Jewish groups with 1043 unrelated individuals from 52 worldwide populations. To further examine the relationship between Jewish communities and European populations, 2407 European subjects were assigned and divided into 10 groups based on geographic region of their origin. This study confirmed previous findings of shared Middle Eastern origin of the above Jewish groups and found that "the genetic connections between the Jewish populations became evident from the frequent IBD across these Jewish groups (63% of all shared segments). Jewish populations shared more and longer segments with one another than with non-Jewish populations, highlighting the commonality of Jewish origin. Among pairs of populations ordered by total sharing, 12 out of the top 20 were pairs of Jewish populations, and "none of the top 30 paired a Jewish population with a non-Jewish one". Atzmon concludes that "Each Jewish group demonstrated Middle Eastern ancestry and variable admixture from host population, while the split between Middle Eastern and European/Syrian Jews, calculated by simulation and comparison of length distributions of IBD segments, occurred 100–150 generations ago, which was described as "compatible with a historical divide that is reported to have occurred more than 2500 years ago" as the Jewish community in Iraq and Iran were formed by Jews in the Babylonian and Persian empires during and after Babylonian exile. The main difference between Mizrahi and Ashkenazi/Sephardic Jews was the absence of Southern European components in the former. According to these results, European/Syrian Jewish populations, including the Ashkenazi Jewish community, were formed latter, as a result of the expulsion and migration of Jews from Palestine, during Roman rule. Concerning Ashkenazi Jews, this study found that genetic dates "are incompatible with theories that Ashkenazi Jews are for the most part the direct lineal descendants of converted Khazars or Slavs". Citing Behar, Atzmon states that "Evidence for founder females of Middle Eastern origin has been observed in all Jewish populations based on non overlapping mitochondrial haplotypes with coalescence times >2000 years". The closest people related to Jewish groups were the Palestinians, Bedouins, Druze, Greeks, and Italians. Regarding this relationship, the authors conclude that "These observations are supported by the significant overlap of Y chromosomal haplogroups between Israeli and Palestinian Arabs with Ashkenazi and non-Ashkenazi Jewish populations".[2][65][5][88]

In 2011, Moorjani et al.[89] detected 3%–5% sub-Saharan African ancestry in all eight of the diverse Jewish populations (Ashkenazi Jews, Syrian Jews, Iranian Jews, Iraqi Jews, Greek Jews, Turkish Jews, Italian Jews) that they analyzed. The timing of this African admixture among all Jewish populations was identical The exact date was not determined, but it was estimated to have taken place between 1,600 and 3,400 years ago. Although African admixture was determined among South Europeans and Near Eastern population too, this admixture was found to be younger compared to the Jewish populations. This findings the authors explained as evidence regarding common origin of these 8 main Jewish groups. "It is intriguing that the Mizrahi Irani and Iraqi Jews—who are thought to descend at least in part from Jews who were exiled to Babylon about 2,600 years ago share the signal of African admixture. A parsimonious explanation for these observations is that they reflect a history in which many of the Jewish groups descend from a common ancestral population which was itself admixed with Africans, prior to the beginning of the Jewish diaspora that occurred in 8th to 6th century BC" the authors concludes.[5][90]

In 2012, two major genetic studies were carried out under the leadership of Harry Ostrer, from the Albert Einstein College of Medicine. The results were published in the Proceedings for the National Academy of Sciences. The genes of 509 Jewish donors from 15 different backgrounds and 114 non-Jewish donors of North African origin were analyzed. Ashkenazi, Sephardi, and Mizrahi Jews were found to be closer genetically to each other than to their long-term host populations, and all of them were found to have Middle Eastern ancestry, together with varying amounts of admixture in their local populations. Mizrahi and Ashkenazi Jews were found to have diverged from each other approximately 2,500 years in the past, approximately the time of the Babylonian exile. The studies also reconfirmed the results of previous studies which found that North African Jews were more closely related to each other and to European and Middle Eastern Jews than to their non-Jewish host populations.,[91] The genome-wide ancestry of North African Jewish groups was compared with respect to European (Basque), Maghrebi (Tunisian non-Jewish), and Middle Eastern (Palestinian) origins. The Middle Eastern component was found to be comparable across all North African Jewish and non-Jewish groups, while North African Jewish groups showed increased European and decreased level of North African (Maghrebi) ancestry [91] with Moroccan and Algerian Jews tending to be genetically closer to Europeans than Djerban Jews. The study found that Yemenite, Ethiopian, and Georgian Jews formed their own distinctive, genetically linked clusters. In particular, Yemenite Jews, who had previously been believed to have lived in isolation, were found to have genetic connections to their host population, suggesting some conversion of local Arabs to Judaism had taken place. Georgian Jews were found to share close connections to Iraqi and Iranian Jews, as well as other Middle Eastern Jewish groups. The study also found that Syrian Jews share more genetic commonality with Ashkenazi Jews than with other Middle Eastern Jewish populations.[92][93][94][95][96] According to the study:

distinctive North African Jewish population clusters with proximity to other Jewish populations and variable degrees of Middle Eastern, European, and North African admixture. Two major subgroups were identified by principal component, neighbor joining tree, and identity-by-descent analysis—Moroccan/Algerian and Djerban/Libyan—that varied in their degree of European admixture. These populations showed a high degree of endogamy and were part of a larger Ashkenazi and Sephardic Jewish group. By principal component analysis, these North African groups were orthogonal to contemporary populations from North and South Morocco, Western Sahara, Tunisia, Libya, and Egypt. Thus, this study is compatible with the history of North African Jews—founding during Classical Antiquity with proselytism of local populations, followed by genetic isolation with the rise of Christianity and then Islam, and admixture following the emigration of Sephardic Jews during the Inquisition.[91]

A 2012 study on Ethiopian Jews showed that while they are primarily related to the indigenous populations of Ethiopia, they do have very distant genetic links to the Middle East from some 2,000 years in the past, and are likely descended from a few Jewish founders. It was speculated that the community began when a few itinerant Jews settled in Ethiopia in ancient times, converted locals to Judaism, and married into the local populations.[97]

A 2012 study by Eran Elhaik analyzed data collected for previous studies and concluded that the DNA of Eastern and Central European Jewish populations indicates that their ancestry is "a mosaic of Caucasus, European, and Semitic ancestries".[98] For the study, Bedouins and Jordanian Hashemites, known to descend from Arabian tribes, were assumed to be a valid genetic surrogate of ancient Jews, whereas the Druze, known to come from Syria, were assumed to be non-Semitic immigrants into the Levant. Armenians and Georgians were also used as surrogate populations for the Khazars, who spoke a Turkic language unrelated to Georgian or Armenian. On this basis, a relatively strong connection to the Caucasus was proposed because of the stronger genetic similarity of these Jewish groups to modern Armenians, Georgians, Azerbaijani Jews, Druze and Cypriots, compared to a weaker genetic similarity with Hashemites and Bedouins. This proposed Caucasian component of ancestry was in turn taken to be consistent with the Khazarian Hypothesis as an explanation of part of the ancestry of Ashkenazi Jews.

A study by Haber et al. (2013) noted that while previous studies of the Levant, which had focused mainly on diaspora Jewish populations, showed that the "Jews form a distinctive cluster in the Middle East", these studies did not make clear "whether the factors driving this structure would also involve other groups in the Levant". The authors found strong evidence that modern Levant populations descend from two major apparent ancestral populations. One set of genetic characteristics which is shared with modern-day Europeans and Central Asians is most prominent in the Levant amongst "Lebanese, Armenians, Cypriots, Druze and Jews, as well as Turks, Iranians and Caucasian populations". The second set of inherited genetic characteristics is shared with populations in other parts of the Middle East as well as some African populations. Levant populations in this category today include "Palestinians, Jordanians, Syrians, as well as North Africans, Ethiopians, Saudis, and Bedouins". Concerning this second component of ancestry, the authors remark that while it correlates with "the pattern of the Islamic expansion", and that "a pre-Islamic expansion Levant was more genetically similar to Europeans than to Middle Easterners," they also say that "its presence in Lebanese Christians, Sephardi and Ashkenazi Jews, Cypriots and Armenians might suggest that its spread to the Levant could also represent an earlier event". The authors also found a strong correlation between religion and apparent ancestry in the Levant:

all Jews (Sephardi and Ashkenazi) cluster in one branch; Druze from Mount Lebanon and Druze from Mount Carmel are depicted on a private branch; and Lebanese Christians form a private branch with the Christian populations of Armenia and Cyprus placing the Lebanese Muslims as an outer group. The predominantly Muslim populations of Syrians, Palestinians and Jordanians cluster on branches with other Muslim populations as distant as Morocco and Yemen.[99]

A 2013 study by Doron M. Behar, Mait Metspalu, Yael Baran, Naama M. Kopelman, Bayazit Yunusbayev et al. using integration of genotypes on newly collected largest data set available to date (1,774 samples from 106 Jewish and non-Jewish populations) for assessment of Ashkenazi Jewish genetic origins from the regions of potential Ashkenazi ancestry: (Europe, the Middle East, and the region historically associated with the Khazar Khaganate) concluded that "This most comprehensive study... does not change and in fact reinforces the conclusions of multiple past studies, including ours and those of other groups (Atzmon and others, 2010; Bauchet and others, 2007; Behar and others, 2010; Campbell and others, 2012; Guha and others, 2012; Haber and others; 2013; Henn and others, 2012; Kopelman and others, 2009; Seldin and others, 2006; Tian and others, 2008). We confirm the notion that the Ashkenazi, North African, and Sephardi Jews share substantial genetic ancestry and that they derive it from Middle Eastern and European populations, with no indication of a detectable Khazar contribution to their genetic origins."

The authors also reanalyzed the 2012 study of Eran Elhaik, and found that "The provocative assumption that Armenians and Georgians could serve as appropriate proxies for Khazar descendants is problematic for a number of reasons as the evidence for ancestry among Caucasus populations do not reflect Khazar ancestry". Also, the authors found that "Even if it were allowed that Caucasus affinities could represent Khazar ancestry, the use of the Armenians and Georgians as Khazar proxies is particularly poor, as they represent the southern part of the Caucasus region, while the Khazar Khaganate was centered in the North Caucasus and further to the north. Furthermore, among populations of the Caucasus, Armenians and Georgians are geographically the closest to the Middle East, and are therefore expected a priori to show the greatest genetic similarity to Middle Eastern populations." Concerning the similarity of South Caucasus populations to Middle Eastern groups which was observed at the level of the whole genome in one recent study (Yunusbayev and others, 2012). The authors found that "Any genetic similarity between Ashkenazi Jews and Armenians and Georgians might merely reflect a common shared Middle Eastern ancestry component, actually providing further support to a Middle Eastern origin of Ashkenazi Jews, rather than a hint for a Khazar origin". The authors claimed "If one accepts the premise that similarity to Armenians and Georgians represents Khazar ancestry for Ashkenazi Jews, then by extension one must also claim that Middle Eastern Jews and many Mediterranean European and Middle Eastern populations are also Khazar descendants. This claim is clearly not valid, as the differences among the various Jewish and non-Jewish populations of Mediterranean Europe and the Middle East predate the period of the Khazars by thousands of years".[15][100]

Two 2014 studies by Paull and colleagues analyzed autosomal SNP data from FTDNA's Family Finder test for 100 study participants, divided into Jewish, non-Jewish, and interfaith study groups. It reported autosomal DNA test values, such as the size and number of shared DNA segments, the number of genetic matches, and the distribution of predicted relationships, varies between study groups. The study also investigates how shared autosomal DNA, and longest block values vary by strength-of-relationship for each study group. According to the results "The 40 participants in the Jewish study group were found to match an average of 24.8 or 62.0 % of the other Jewish study participants, while the 40 participants in the non-Jewish study group matched an average of 4.0 or 9.9 % of the other non-Jewish study participants. Hence, Jewish study participants had over 6 times more matches with each other than did non-Jewish study participants. With the exception of a single study participant, there were no matches between the Jewish and non-Jewish study groups."[101][102]

A 2014 study by Carmi et al. published by Nature Communications found that the Ashkenazi Jewish population originates from an even mixture between Middle Eastern and European peoples.[103] The study found that all Ashkenazi Jews descend from around 350 individuals who were genetically about half Middle Eastern and half European. This would make all Ashkenazi Jews related to the point of being at least 30th cousins.[104] According to the authors, this genetic bottleneck likely occurred some 600–800 years in the past , followed by rapid growth and genetic isolation (rate per generation 16–53%;). The principal component analysis of common variants in the sequenced AJ samples, confirmed previous observations, namely, the proximity of Ashkenazi Jewish cluster to other Jewish, European and Middle Eastern populations.[105][103]

A 2016 study by Elhaik et al. in the Oxford University Press published journal Genome Biology and Evolution found that the DNA of Ashkenazi Jews originated in northeastern Turkey.[17] The study found 90% of Ashkenazi Jews could be traced to four ancient villages in northeastern Turkey. The researchers speculated that the Ashkenazi Jews originated in the first millennium, when Iranian Jews converted Greco-Roman, Turkish, Iranian, southern Caucasian, and Slavic populations inhabiting Turkey, and speculated that the Yiddish language also originated there among Jewish merchants as a cryptic language in order to gain advantage in trade along the Silk Road.[106][107]

In joint study published in 2016 by Genome Biology and Evolution, a group of geneticists and linguists from the UK, Czech Republic, Russia and Lithuania, dismissed both the genetic and linguistic components of Elhaik's 2016 study. As for the genetic component, the authors argued that using a genetic "GPS tool" (as used by Elhaik et al.) would place Italians and Spaniards into Greece, all Tunisians and some Kuwaitis would be placed in the Mediterranean Sea, all Greeks were positioned in Bulgaria and in the Black Sea, and all Lebanese were scattered along a line connecting Egypt and the Caucasus; "These cases are sufficient to illustrate that mapping of test individuals has nothing to do with ancestral locations" the authors wrote. As for the linguistic component, the authors stated "Yiddish is a Germanic language, leaving no room for the Slavic relexification hypothesis and for the idea of early Yiddish-Persian contacts in Asia Minor. The study concluded that ‘Yiddish is a Slavic language created by Irano-Turko-Slavic Jewish merchants along the Silk Roads as a cryptic trade language, spoken only by its originators to gain an advantage in trade’ (Das et al. (2016) remains an assertion in the realm of unsupported speculation", the study concluded.[20]

A 2016 study of Indian Jews from the Bene Israel community by Waldman et al. found that the genetic composition of the community is "unique among Indian and Pakistani populations we analyzed in sharing considerable genetic ancestry with other Jewish populations. Putting together the results from all analyses point to Bene Israel being an admixed population with both Jewish and Indian ancestry, with the genetic contribution of each of these ancestral populations being substantial." The authors also examined the proportion and roots of the shared Jewish ancestry and the local genetic admixture: "In addition, we performed f4-based analysis to test whether Bene Israel are closer to Jews than to non-Jewish Middle-Eastern populations. We found that Middle-Eastern Jewish populations were closer to Bene Israel as compared to other Middle-Eastern populations examined (Druze, Bedouin and Palestinians). Non-Middle-Eastern Jewish populations were still closer to Bene Israel as compared to Bedouin and Palestinians, but not as compared to Druze. These results further support the hypothesis that the non-Indian ancestry of Bene Israel is Jewish specific, likely from a Middle-Eastern Jewish population."[108]

A 2017 study by Xue et al. found an approximately even mixture of Middle Eastern, particularly Levantine, and European ancestry in Ashkenazi Jews, with the majority of the European ancestry being of southern European origin. The study estimated that roughly 50% of the ancestral origin of Ashkenazi Jews was Middle Eastern, 34% was southern European, and 16% varied between western and eastern European.[109]

A 2020 genetic study on Bronze Age southern Levantine (Canaanite) remains found evidence of large scale migration of populations related to those of the Zagros or Caucasus into the southern Levant by the Bronze Age (resulting in a Bronze Age Canaanite population descended from both those migrants and earlier Neolithic Levantine peoples), and suggested a substantial genetic continuity from the Levantine Bronze Age both in modern-day non-Jewish Arabic-speaking Levantine populations (such as Syrians, Lebanese, Palestinians, and Druze) and Jewish groups (such as Moroccan Jews of Sephardic origin, Ashkenazi Jews, and Mizrahi Jews), who were all found to carry a significant degree of Bronze Age-era Levantine ancestry, with Ashkenazi Jews deriving over half of their ancestry from Bronze-Age Levantines and about 41% from European admixture, and Arabic-speaking Levantines, Moroccan Jews, and Mizrahi Jews deriving a larger majority of their ancestry from that population. Ethiopian Jews were found to derive the majority of their ancestry from an East African or Horn African component, but also carried some minor Canaanite-like ancestry.[25]

Wikimedia Commons has media related to Genetic studies on Jewish populations.

• List of publications from Microsoft Academic
• "DNA links prove Jews are a 'race,' says genetics expert". Haaretz. 7 May 2012. Retrieved 30 November 2013.
• Paul S. Appelbaum (12 February 2008). "Genetics and the Jewish identity". Jerusalem Post. Retrieved 27 August 2019.
• Wade, Nicholas (14 January 2006). "New Light on Origins of Ashkenazi in Europe". The New York Times. Retrieved 24 May 2006.
• Steve Olson (July–August 2007). "Who's Your Daddy?". The Atlantic. Retrieved 19 February 2009.
• "Priestly Gene Shared By Widely Dispersed Jews". American Society for Technion, Israel Institute of Technology. 14 July 1998.
• Jon Entine. "Jewish DNA - Genetic Research and The Origins of the Jewish People". Retrieved 11 October 2013.
• Shaul Stampfer (22 June 2017). "The Myth of the Khazar Conversion and the Origin of the Ashkenazi Jews".