Haplogroup E1b1b (Y-DNA): Wikis


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Haplogroup E1b1b (E-M215)
Time of origin approx 22,400 years BP[1]
Place of origin Eastern Africa[2][3]
Ancestor E1b1
Descendants E1b1b1 (E-M35), which in turn has sub-clades:-
1. E1b1b1a (E-M78)
2. E1b1b1b (E-M81)
3. E1b1b1c (E-M123)
4. E1b1b1d (E-M281)
5. E1b1b1e (E-V6)
6. E1b1b1f (E-P72)
7. E1b1b1g (E-M293).
Defining mutations M215, most often found in conjunction with M35

In human genetics, Y Haplogroup E1b1b (E-M215) previously known as E3b (or "haplotype V" in the Lucotte classification [4]) is a Y-chromosome haplogroup, a sub-group of haplogroup E, which is defined by the single nucleotide polymorphism (SNP) mutation M215.[5][6][7] It is one of the major genetically distinguished paternal lines of the human race, linking from father-to-son back to a common male ancestor.


Current and previous names

E1b1b and E1b1b1 are the currently accepted names found in the proposals of the Y Chromosome Consortium (YCC).[6] The nomenclature E3b (E-M215) and E3b1 (E-M35) respectively were the YCC defined names used to designate the same haplogroups in older literature with E-M35 branching as a separate subclade of E-M215 in 2004.[2][7] Prior to 2002 these haplogroups were not designated in a consistent way, and nor was their relationship to other related clades within haplogroup E and haplogroup DE.[7]


E1b1b1 origins map

E1b1b (E-M215) and its dominant sub-clade E1b1b1 (E-M35) are believed to have first appeared in East Africa about 22,400 years ago.[1][Note 1][Note 2][Note 3][Note 4]

The ancient dispersals of the major E1b1b1 (E-M35) lineages. The map shows the earliest movements of E1b1b lineages as described in the most recent articles[2][1][8][9].

All major sub-branches of E1b1b1 are thought to have originated in the same general area as the parent clade: in North Africa, East Africa, or nearby areas of the Near East. Underhill (2002) believes that the structure and regional pattern of E-M35 sub-clades potentially give "reagents with which to infer specific episodes of population histories associated with the Neolithic agricultural expansion". Concerning European E-M35 within this scheme, Underhill and Kivisild (2007) have remarked that E1b1b seems to represent a late-Pleistocene migration from North Africa to Europe over the Sinai Peninsula in Egypt.[Note 5]

Concerning E1b1b, Coffman-Levy (2005) wrote that although E1b1b "arose in East Africa, approximately 25000 years ago, certain sub-clades appear to have been present in Europe and Asia for thousands of years" and so it is "often incorrectly described as 'African'" in a sense that creates a "misimpression regarding the origin and complex history of this haplogroup", which has pervaded the public and media.[Note 6]


E1b1b is distributed as far south as South Africa, and northwards into North Africa, from where it has in more recent millennia expanded to Europe and Asia.[2] E1b1b1 (E-M35) is the predominant subclade of E1b1b, representing almost exactly the same population. M215 was found to be older than M35 when individuals were found who have the M215 mutation, but do not have M35 mutation.[2] The E1b1b clade is presently found in various forms in the Horn of Africa, North Africa, parts of Eastern, Western, and Southern Africa, West Asia, and Europe (especially the Mediterranean and the Balkans)[2][3][10][11].

E1b1b and E1b1b1 are quite common amongst Afro-Asiatic speakers. The linguistic group and E1b1b1 may have dispersed together from the region of origin of this language family.[12][13][14] Amongst populations with an Afro-Asiatic speaking history, a significant proportion of Jewish male lineages are E1b1b1 (E-M35).[15] Haplogroup E1b1b1, which accounts for approximately 18%[3] to 20%[16][17] of Ashkenazi and 8.6%[18] to 30%[3] of Sephardi Y-chromosomes, appears to be one of the major founding lineages of the Jewish population.[19][Note 7]

Subclades of E1b1b1 (E-M35)

A large majority of E1b1b lineages are within E1b1b1 (defined by M35). Exceptions discovered so far are M215 positive/M35 negative ("E-M215*") cases found in two Amharic Ethiopians and 1 Yemeni.[2][20]

cladogram of E1b1b, E1b1b1, and its daughter clades

The E-M215 derivative, E1b1b1 (E-M35) is defined by the M35 SNP. E-M35 includes individuals with the "ancestral state" (no known sub-clade forming mutations). These are referred to as E1b1b1* or E-M35*. As of 2009, there are seven known branches that have resulted from different mutations on M35: M78, M81, M123, M281, V6, P72, and M293. In order to show what is known of their relationships to E1b1b1 and other related clades, these are also currently referred to as E1b1b1a to E1b1b1g, respectively (see image). The more frequently described sub-clades are E1b1b1a and E1b1b1b. Both are found in Mediterranean & West Asian peoples. These two sub-clades represent the largest proportion of E1b1b. E1b1b1a is found over most of the range where E1b1b is found excluding Southern Africa. E1b1b1b is found mainly in the Maghreb. E1b1b1c is less common but widely scattered, with significant populations in specific parts of the Horn of Africa, the Levant, Arabia, Iberia, and Anatolia. E1b1b1g is a fourth major sub-clade that has been found in parts of Eastern and Southern Africa, includes the majority of unique E1b1b1 lineages in sub-Saharan Africa (those that lack M78, M81, or M123 mutations).[8] Two smaller sub-clades are defined by mutations M281 and V6 appear to be unique to the Horn of Africa region.


E1b1b1a (E-M78)

Distribution density of E1b1b1a (E-M78) in select areas of Africa and Eurasia

E1b1b1a (E-M78), formerly E3b1a, is a commonly occurring subclade, widely distributed in North Africa, the Horn of Africa, West Asia, i.e. The Middle East and Near East "up to Southern Asia",[Note 8] and all of Europe.[Note 9] The European distribution has a frequency peak centered in parts of the Balkans (up to almost 50%[3][21]) and Italy and declining frequencies evident toward western, central, and northeastern Europe.

Based on genetic STR variance data, Cruciani et al. (2007) suggests that this subclade originated in "Northeastern Africa", which in the study refers specifically to Egypt and Libya.[Note 10][Note 11] about 18,600 years ago (17,300 - 20,000 years ago).[Note 12] Battaglia et al. (2008) describe Egypt as "a hub for the distribution of the various geographically localized M78-related sub-clades" and, based on archaeological data, they propose that the point of origin of E-M78 (as opposed to later dispersals from Egypt) may have been in a refugium which "existed on the border of present-day Sudan and Egypt, near Lake Nubia, until the onset of a humid phase around 8500 BC. The northward-moving rainfall belts during this period could have also spurred a rapid migration of Mesolithic foragers northwards in Africa, the Levant and ultimately onwards to Asia Minor and Europe, where they each eventually differentiated into their regionally distinctive branches". Towards the south, Hassan et al. (2008) also explain evidence that some subclades of E-M78, specifically E-V12 and E-22, "might have been brought to Sudan from North Africa after the progressive desertification of the Sahara around 6,000-8,000 years ago".

E1b1b1b (E-M81)

E1b1b1b (E-M81), formerly E3b1b or E3b2, is the most common Y chromosome haplogroup in the Maghreb, dominated by its sub-clade E-M183. It is thought to have originated in the area of North Africa 5,600 years ago.[2][22] It is colloquially referred to as the "Berber marker" for its prevalence among Mozabite, Moyen Atlas, Kabyle and other Amazigh groups, E-M81 is also quite common among North African Arab groups. It reaches frequencies of up to 80% in the Maghreb. This includes the Saharawish for whose men Bosch et al. (2001) reports that approximately 76% are M81+.

Distribution of E1b1b1b in select areas of Europe, Asia, and Africa

In this key area from Egypt to the Atlantic Ocean, Arredi et al. (2004) report a pattern of decreasing STR haplotype variation from East to West, accompanied by a substantial increasing frequency. At the eastern extreme of this core range, Kujanova et al. (2009) found M81 in 28.6% (10 out of 35 men) in El-Hayez in the Western desert in Egypt.

Arredi et al. (2004) believe the pattern of distribution and variance to be consistent with the hypothesis of a post Paleolithic "demic diffusion" from the East. The ancestral lineage of E-M81 in their hypothesis could have been linked with the spread of Neolithic food-producing technologies from the Fertile Crescent via the Nile, although pastoralism rather than agriculture. E-M81 and possibly proto-Afroasiatic language may have been carried either all the way from Asia, or they may represent a "local contribution to the North African Neolithic transition". According to Shomarka Keita, a Near Eastern origin of proto-Afroasiatic speakers carrying E-M81, or its ancestral lineage, is inconsistent with the linguistic evidence, which seems to indicate an African origin of Proto-Afro-Asiatic speakers. Keita argues that there is no autocthonous presence of E-M81 in the Near East, indicating that M81 most likely emerged from its parent clade M35 either in the Maghreb, or possibly as far south as the Horn of Africa.[23]

In Europe, E-M81 is found everywhere but mostly in the Iberian Peninsula, where unlike in the rest of Europe[Note 13] it is more common than E-M78, with an average frequency of 4-5.6%, and frequencies reaching 9% in Galicia, 10% in Western Andalusia and Northwest Castile and 13% in Cantabria.[24][25][26][27] The highest frequency of this clade found so far in Europe has been observed at 40% the Pasiegos from Cantabria.[2] Recent research suggests that E-M81 in Iberia and some other regions of Atlantic Europe resulted from pre-Neolithic Eurasian Berber migrations from North-west Africa.

E-M81 is also found in Sicily,[28] and in slightly lower frequencies in continental Italy (especially near Lucera)[27] and France,[2] possibly due to ancient migrations during the Islamic, Roman, and Carthaginian empires.

As a result of its old world distribution, this sub-clade is found throughout Latin America, for example 6.1% in Cuba,[29] 5.4% in Brazil (Rio de Janeiro), [Note 14] and among Hispanic men from California and Hawaii 2.4%.[30]

In smaller numbers, E-M81 men can be found in areas in contact with the Maghreb, both around the Sahara, in places like Sudan, and around the Mediterranean in places like Lebanon, Turkey, and amongst Sephardic Jews.

There are two recognized sub-clades, although one is much more important than the other.

Sub Clades of E1b1b1b (E-M81):
  • E1b1b1b1 (E-M107). Underhill et al. (2000) found one example in Mali.
  • E1b1b1b2 (E-M183). This clade is extremely dominant within E-M81. In fact, while Karafet et al. (2008) continues to describe this as a sub-clade of E-M81, and ISOGG defers to Karafet et al., all data seems to imply that it should actually be considered phylogenetically equivalent to M81. As of 24th November 2008, several SNPs are considered to define sub-clades of E-M183, although the phylogenetic structure is not yet known with confidence: M165, M243, M340, and L19.[5]

E1b1b1c (E-M123)

Distribution of E1b1b1c in select areas of Europe, Asia, and Africa

E1b1b1 (E-M35), formerly E3b1c or E3b3, is mostly known for its major sub-clade E1b1b1c1 (E-M34), which dominates this clade.[Note 15] However, earlier studies did not test for E-M34.

Concerning E-M123* (tested and definitely without E-M34) Cruciani et al. (2004) located one individual in Bulgaria after testing 3401 individuals from five continents, and Underhill et al. (2000) located one individual in Central Asia. In a 568 person study in Iberia, Flores et al. (2004) found 2 E-M123* individuals, both in Northern Portugal out of 109 people tested there. In a 553 person study of Portugal, Gonçalves et al. (2005) also found 2 E-M123* individuals in Northern Portugal, out of 101 people, as well as 2 in Madeira out of 129 people tested there. Flores et al. (2005) found one individual out of 146 Jordanians. Cadenas et al. (2007) found none amongst the significant presence of E-M34 they found in their study of the UAE, Yemen and Qatar. Arredi et al. (2004) found 1 Tunisian in their study of 275 men in Northern Africa. Zalloua et al. (2008) found 26 E-M123 cases in Cyprus, out of 164 men tested; and 27 Palestinians out of 291 tested[31]

Concerning E-M123 without checking for the M-34 SNP Bosch et al. (2006) found E-M123 examples in Greece, the Republic of Macedonia, and Roumania. Beleza et al. (2006) also found examples in Portugal, and Sanchez et al. (2005) found one sample in Somalia. Semino et al. (2004) reports relatively high levels of 13% in the Albanian community of Cosenza, in Calabria. A notably high regional frequency for E-M123 was reported in Oman, where it is apparently the dominant clade of E-M35. Luis et al. (2004) found 12 men out of 121 there were E-M123 positive, while in Egypt there were 7 out of 147. But in that study the Omani E-M123 diversity implied a younger age than the E-M123 found in Egypt. Shen et al. (2004) found 4 out of 20 tested Israeli Jews of Libyan ancestry to be M123+.

Concerning E1b1b1c1 (E-M34) Cruciani (2004) tested for E-M34 in Oman and found 7.7% to be E-M34+, with no E-M123*. According to Cruciani (2004), E-M34 is found at small frequencies in North Africa and Southern Europe (6.6% in Sicily for example), and has its highest concentration in Ethiopia and the Near East (with highest levels in Oman and Turkey). However, because the diversity is apparently low in Ethiopia, the authors suggest that E-M34 was likely introduced into Ethiopia from the Near East. In Turkey, Cinnioğlu et al. (2004) found slightly more E-M34 (29) than E-M78 (26) out of 523 individuals tested (a far different E1b1b population than found in the nearby Balkans). In Flores et al. (2004) E-M34 was found in several parts of Iberia, but most strikingly about 10% in Galicia. Gonçalves et al. (2005) found about the same levels of E-M34 in Portugal as E-M123*, but E-M34 mainly in Central Portugal (4 people out of 102 tested there) with one more person found in the Açores. Strikingly, Flores et al. (2005) found 14 out of 45 men tested in the Dead Sea area of Jordan to be M34 positive (31.1%), while in the capital Amman there were only 4 out of 101. Cadenas et al. (2007) found 8.1% of 62 men tested in Yemen were positive for M34, compared to much lower levels in Qatar (1.4%) and the UAE (3.1%). Arredi et al. (2004) found frequencies of 10.50% in Kabyles from Algeria, 9.5% in Egyptians and 1.50% in Tunisians.

E-M123 in Jews. Looking beyond simple regional concentrations, E1b1b1c (E-M123) is also quite common among both Ashkenazi and Sephardic Jews, accounting for over 10% of all male lines.[3] Coffman-Levy (2005) wrote that:

...the best candidate for possible E3b Israelite ancestry among Jews is E-M123. This sub-clade occurs in almost the same proportions (approximately 10-12%) among both Ashkenazim and Sephardim (Semino et al. (2004)). According to Cruciani (2004), E-M123 probably originated in the Middle East, since it is found in a large majority of the populations from that area, and then back-migrated to Ethiopia. He further notes that this sub-clade may have been spread to Europe during the Neolithic agricultural expansion out of the Middle East. However, because E-M123 is also found in low percentages (1-3%) in many southern European and Balkan populations, its origin among Jewish groups remains uncertain (Semino et al. (2004)). Yet the fact that both Sephardim and Ashkenazim possess this sub-clade in similar high frequency supports an Israelite/Middle Eastern origin.

However, Cruciani Cruciani (2004), do not rule out an East Africa origin for this haplogroup stating "the introduction of E-M34 from Africa directly to southern-central Europe cannot be excluded at the present."

Sub Clades of E1b1b1c1 (E-M34):
  • E1b1b1c1a. Defined by SNP mutation M84, with M136 defining a sub-clade, E1b1b1c1a1 as of October 2008.[5] The E-M35 Phylogeny Project estimates based on testing so far (in January 2009) that E-M84 is dominant in 6 out of the 8 clusters of E-M34 which that project identifies.
  • E1b1b1c1b. Defined by SNP mutation M290. Shen et al. (2004) found 1 Palestinian exemplar.

E1b1b1d (E-M281)

The discovery of the SNP mutation which defines this sub-clade of E-M35, M281, was announced Semino et al. (2002), who found it in two Ethiopian Oromo, but Cruciani et al. (2004) found no examples.

E1b1b1e (E-V6)

This sub-clade of E-M35 is defined by V6. Cruciani et al. (2004) (Table 1) identified a significant presence of these lineages in Ethiopia, and also some in the neighboring Somali population. Amongst the Ethiopian and Somali samples, the highest were 14.7% amongst the Ethiopian Amhara, and 16.7% amongst the Ethiopian Wolayta. One man in Kenya was also observed with the V6 mutation.

E1b1b1f (E-P72)

Appears in Karafet et al. (2008). Little has been published about this sub-clade of E-M35. Note also the potential for name confusion with E-M293 below.

E1b1b1g (E-M293)

This sub-clade of E-M35 was announced in Henn et al. (2008), which associated it with the spread of pastoralism from Eastern Africa into Southern Africa. So far high levels have been found in specific ethnic groups in Tanzania and Southern Africa. Highest were the Datog (43%), Khwe (Kxoe) (31%), Burunge (28%), and Sandawe (24%). Henn et al. (2008) in their study also found two Bantu-speaking Kenyan males with the M293 mutation.[8]

Other E1b1b sub-clades are rare in Southern Africa. The authors state...

Without information about M293 in the Maasai, Hema, and other populations in Kenya, Sudan, and Ethiopia, we cannot pinpoint the precise geographic source of M293 with greater confidence. However, the available evidence points to present-day Tanzania as an early and important geographic locus of M293 evolution.

They also say that "M293 is only found in sub-Saharan Africa, indicating a separate phylogenetic history for M35* (former) samples further north".

The authors Henn et al. referred to this sub-clade with the proposed name E3b1f. However, this name was already out of date by the time the article was published since E1b1b1 had become the new YCC and ISOGG name for former E3b1, the clade defined by SNP M35. The sub-clade under E1b1b1 with the suffix "f" had also already been proposed in Karafet et al. (2008) for SNP P72 (see above). So the phylogenetic clade name came to be E1b1b1g in late October 2008.[Note 16]


This phylogenetic tree of haplogroup subclades is based on the YCC 2008 tree [6] and subsequent published research as summarized by ISOGG.[5]

  • E1b1b (M215)
    • E1b1b1 (M35)
      • E1b1b1a (M78)
        • E1b1b1a1 (V12)
          • E1b1b1a1a (M224)
          • E1b1b1a1b (V32)
        • E1b1b1a2 (V13, V36)
          • E1b1b1a2a (V27)
          • E1b1b1a2b (P65)
          • E1b1b1a2c (L17)
        • E1b1b1a3 (V22)
          • E1b1b1a3a (M148)
          • E1b1b1a3b (V19)
        • E1b1b1a4 (V65)
        • E1b1b1a5 (M521)
      • E1b1b1b (M81)
        • E1b1b1b1 (M107)
        • E1b1b1b2 (M183, M310, L19)
          • E1b1b1b2a (M165)
      • E1b1b1c (M123)
        • E1b1b1c1 (M34)
          • E1b1b1c1a (M84)
            • E1b1b1c1a1 (M136)
          • E1b1b1c1b (M290)
      • E1b1b1d (M281)
      • E1b1b1e (V6)
      • E1b1b1f (P72)
      • E1b1b1g (M293)

See also

Human Y-chromosome DNA (Y-DNA) haplogroups (by ethnic groups · famous haplotypes)

most recent common Y-ancestor
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  1. ^ For E1b1b (M-215) Cruciani et al. (2007) reduced their estimate to 22,400 from 25,600 in Cruciani et al. (2004), re-calibrating the same data.
  2. ^ As explained above, the modern population of E-M215 and E-M35 lineages are almost identical, and therefore by definition age estimates based on these two populations are also.
  3. ^ Semino et al. (2004)"This inference is further supported by the presence of additional Hg E lineal diversification and by the highest frequency of E-P2* and E-M35* in the same region. The distribution of E-P2* appears limited to eastern African peoples. The E-M35* lineage shows its highest frequency (19.2%) in the Ethiopian Oromo but with a wider distribution range than E-P2*."
  4. ^ Cruciani et al. (2004): "Several observations point to eastern Africa as the homeland for haplogroup E3b—that is, it had (1) the highest number of different E3b clades (table 1), (2) a high frequency of this haplogroup and a high microsatellite diversity, and, finally, (3) the exclusive presence of the undifferentiated E3b* paragroup." As mentioned above, "E3b" is the old name for E1b1b (E-M215).
  5. ^ "Y chromosome data show a signal for a separate late-Pleistocene migration from Africa to Europe via Sinai as evidenced through the distribution of haplogroup E3b lineages, which is not manifested in mtDNA haplogroup distributions."Underhill and Kivisild (2007:547)
  6. ^ Coffman-Levy (2005): "Unfortunately, misinformation about these haplogroups continues to pervade the public and media. Haplogroup E3b is often incorrectly described as “African,” leaving a misimpression regarding the origin and complex history of this haplogroup. Haplogroup J2, as previously discussed, is often incorrectly equated with J1 and described as “Jewish” or “Semitic,” despite the fact that it is present in a variety of non-Jewish Mediterranean and Northern European populations. And haplogroup G is rarely discussed in depth; its origin and distribution remain poorly understood."
  7. ^ "Paragroup EM35* and haplogroup J-12f2a* fit the criteria for major AJ founding lineages because they are widespread both in AJ populations and in Near Eastern populations, and occur at much lower frequencies in European non-Jewish populations." Semino et al. (2008)
  8. ^ Cruciani et al. (2007):E-M78 shows "a wide geographic distribution" and is "relatively common not only in northeastern and eastern Africa but also found in Europe and western Asia, up to Southern Asia".
  9. ^ Cruciani et al. (2006): "The human Y chromosome haplogroup E-M78 (E3b1a) occurs commonly and is distributed in northern and eastern Africa, western Asia, and all of Europe."
  10. ^ Cruciani et al. (2007) use the term "Northeastern Africa" to refer to Egypt and Libya, as shown in Table 1 of the study.
  11. ^ Prior to Cruciani et al. (2007), Semino et al. (2004) had proposed the Horn of Africa as a possible place of origin of E-M78. This was because of the high frequency and diversity of E-M78 lineages in the region. For example, Sanchez et al. (2005) found that 77.6% of 201 male Somalis tested in Denmark were members of this clade. However, Cruciani et al. (2007) were able to study more data, including populations from North Africa who were not represented in the Semino et al. (2004) study, and found evidence that the E-M78 lineages in the Horn of Africa were relatively recent branches (see E1b1b1a1b (E-V32) below). They concluded that Northeast Africa was the likely place of origin of E-M78 based on "the peripheral geographic distribution of the most derived subhaplogroups with respect to northeastern Africa, as well as the results of quantitative analysis of UEP and microsatellite diversity". E-M215, the parent clade of E-M78, originated in East Africa during the paleolithic and subsequently, E-M215 spread to Northeast Africa. According to Cruciani et al. (2007), the presence of E-M78 in East Africa, is the result of a back migration of E-M215 chromosomes that had acquired the E-M78 mutation. Cruciani et al. (2007) also note this as evidence for "a corridor for bidirectional migrations" between Northeast Africa (Egypt and Libya in their data) on the one hand and East Africa on the other. The authors believe there were "at least 2 episodes between 23.9–17.3 ky and 18.0–5.9 ky ago".
  12. ^ Cruciani et al. (2007) use two calculation methods for estimating the age of E-M78 which give very different results. For the main 18,600 years ago, the ASD method is used, while for a second "ρ method", used as a check, gives 13.7kya with a standard deviation of 2.3kya, but the difference between the two methods is only large for the age estimation of E-M78, not its sub-clades. The authors state the the big difference is "attributable to the relevant departure from a star-like structure because of repeated founder effects"
  13. ^ Adams et al. (2008), shows an average frequency of 4% in the Iberian Peninsula with frequencies reaching 9% in Galicia, 10% in Western Andalusia and Northwest Castile, see table.
  14. ^ (6 out of 112), "The presence of chromosomes of North African origin (E3b1b-M81; Cruciani et al., 2004) can also be explained by a Portuguese-mediated influx, since this haplogroup reaches a frequency of 5.6% in Portugal (Beleza et al., (2006)), quite similar to the frequency found in Rio de Janeiro (5.4%) among European contributors." Silva et al. (2006)
  15. ^ As of 11 November 2008 for example, the E-M35 phylogeny project had records of four E-M123* tests, compared to 93 test results with E-M34.
  16. ^ For example the company Familytree DNA changed its webpages on or about 22 October and the relevant ISOGG reference page was changed on 23 October
  1. ^ a b c Cruciani et al. (2007)
  2. ^ a b c d e f g h i j Cruciani et al. (2004)
  3. ^ a b c d e f Semino et al. (2004)
  4. ^ Gérard et al. (2006)
  5. ^ a b c d ISOGG (2008)
  6. ^ a b c Karafet et al. (2008)
  7. ^ a b c Y Chromosome Consortium "YCC" (2002)
  8. ^ a b c Henn et al. (2008)
  9. ^ Hassan et al. (2008)
  10. ^ Rosser et al. (2000)
  11. ^ Firasat et al. (2006)
  12. ^ Ehret et al. (2004)
  13. ^ Keita and Boyce (2005)
  14. ^ Keita Shomarka (2008)
  15. ^ Behar et al. (2003)
  16. ^ Doron M. Behar, Daniel Garrigan, Matthew E. Kaplan et al., "Contrasting patterns of Y chromosome variation in Ashkenazi Jewish and host non-Jewish European populations," Human Genetics (2004) 114 : 354–365
  17. ^ Shen et al. (2004), "Reconstruction of Patrilineages and Matrilineages of Samaritans and Other Israeli Populations From Y-Chromosome and Mitochondrial DNA Sequence Variation," Human Mutation 24: 248, doi:10.1002/humu.20077
  18. ^ Susan M. Adams, Elena Bosch, Patricia L. Balaresque et al., "The Genetic Legacy of Religious Diversity and Intolerance: Paternal Lineages of Christians, Jews, and Muslims in the Iberian Peninsula," American Journal of Human Genetics 83, 725–736, December 12, 2008
  19. ^ Nebel et al. (2001)
  20. ^ Cadenas et al. (2007)
  21. ^ Peričic et al. (2005)
  22. ^ Arredi et al. (2004)
  23. ^ Keita (2008). "Geography, selected Afro-Asiatic families, and Y chromosome lineage variation". In Hot Pursuit of Language. http://books.google.com/books?hl=en&lr=&id=xxcdjUGfx40C&oi=fnd&pg=PA3. 
  24. ^ Flores et al. (2005)
  25. ^ Beleza et al. (2006)
  26. ^ Adams et al. (2008)
  27. ^ a b Capelli et al. (2009)
  28. ^ Gaetano et al. (2008)
  29. ^ (8 out of 132), Mendizabal et al. (2008)
  30. ^ (7 out of 295), Paracchini et al. (2003)
  31. ^ See also El-Sibai et al. (2009) for the same data in a different format.


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