Haplogroup E1b1b (Y-DNA) explained

E1b1b or E-M215
Origin-Place:East Africa or Middle East[1] ;
Origin-Date:approx 26,000 years BP
Ancestor:E1b1 or E-P2
Descendants:E1b1b1 or E-M35
Mutations:M215, and most often also M35

In human genetics, Y Haplogroup E1b1b (E-M215) is a Y-chromosome haplogroup, a sub-group of haplogroup E, which is defined by the single nucleotide polymorphism (SNP) mutation M215. 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.

In nearly all discussion, E1b1b is equivalent to its very dominant sub-clade, E1b1b1 (E-M35), which contains nearly all of E1b1b. announced the discovery that M215 was older than M35, because that survey found that some lineages which have the M215 mutation, do not have M35. On that basis this article covers both clades, but is named after the slightly larger one.

As discussed in more detail below, E1b1b is presently found in various forms in the Horn of Africa, North Africa, parts of Eastern and Southern Africa, West Asia, and Europe (especially the Mediterranean and the Balkans).

E1b1b and E1b1b1 are quite common amongst populations speaking an Afro-Asiatic language. According to at least some theories on the origins of this linguistic group, such as those advanced by Christopher Ehret, Afro-Asiatic and E1b1b1 may have dispersed amongst the same populations from the same point of origin, making E-M35 a useful tool for speculation into the origins of Afro-Asiatic . However, this hypothesis is disputed by those who suggest a Levantine origin for Afro-Asiatic.[2]

E1b1b is also common amongst populations with an Afro-Asiatic speaking history. A significant proportion of all Jewish male lines are made up of a wide variety of E1b1b1 (E-M35) sub-clades. found only haplogroup J lineages in higher numbers amongst Ashkenazim.[3] The authors also found E1b1b to be, along with haplogroup J, one of the major founding lineages among Ashkenazi Jews.[4] E1b1b is observed in over 22.8% of Ashkenazis and 30% of Sephardim. The variety of sub-clades is felt by many researchers to be a potential lead in seeking a better understanding of Jewish population movements over the centuries.

Other Names, and history of the classification

The current phylogenetic terminology "E1b1b" and "E1b1b1" was proposed by . This paper was intended to be an update of the "Y Chromosome Consortium"(. The YCC first formalized the original phylogenetic nomenclature - "E3b" (E-M215) and "E3b1" (E-M35) - which is still found widely especially in older literature.

It was also the 2002 consortium which proposed guidelines on the mutation nomenclature, "E-M215" and "E-M35". The mutation-based clade names have increasingly been used since then because they avoid the confusion which comes from the increasingly frequent discoveries of new SNP mutations - for example when older and newer literature is being compared.

Prior to, both E1b1b and E1b1b1, not yet distinguished at that time, had been referred to as Hg21 (Haplogroup 21) within 's nomenclature, or as Eu4 according to 's classification.

They were also within 's "Group III" (which included all E haplotypes).

Other older names are referred to in the YCC 2002 report in the referenced articles, but are less common in the literature.

Origins

Concerning the origins of the E1b1b lineage,, [5], [6], (2006), and (2007), point to evidence that not only E1b1b (E-M215), but also both its parent lineage E1b1 (E-P2), and its dominant sub-clade E1b1b1 (E-M35) probably all first appeared in East Africa between 20,000 and 47,500 years ago. There are different techniques available for such estimates, and a considerable range of possibilities, but the most recent estimates of are around 24,000 years ago for E-M215[7] or E-M35.[8]

Citing, wrote that E1b1b1 (E-M35) "arose in East Africa". However, she added that this haplogroup is "often incorrectly described as “African,” leaving a misimpression regarding the origin and complex history of this haplogroup", and that such misinformation about this haplogroup also continued to pervade the public and media at least until the time of writing in 2005.[9]

According to the International Society of Genetic Genealogy (ISOGG) and National Geographic's Genographic Project, E1b1b1 may have arisen instead in the Near East or the Middle East and then expanded into the Mediterranean with the spread of agriculture.,[10]

All major sub-branches of E1b1b1 are thought to have originated in the same general area as the parent clade: in North Africa, the Horn of Africa, or the Near East. 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".

Subclades of E1b1b1 (E-M35)

As mentioned above, nearly all E1b1b lineages are within E1b1b1 (defined by M35). found 2 out of 34 Ethiopian Amhara tested, to be M215 positive but M35 negative, and therefore in the paragroup "E-M215*". More recently, found one more E-M215* individual in Yemen, just across the Red Sea from the Amhara, out of 62 people tested there.

Turning to E-M35, the most current phylogeny of E1b1b1 includes the individuals with no known sub-clade mutations (who are therefore said to be in the "ancestral state" referred to as E1b1b1* or E-M35*) plus seven known "derived" branches, which are defined by the following SNPs: M78, M81, M123, M281, V6, P72, and M293, all of which are discussed below.

The two most written-about sub-clades of E1b1b1 are E1b1b1a (defined by M78) and E1b1b1b (defined by M81), both are associated with the Mediterranean. They are thought to represent the two sub-clades with the largest populations within E1b1b. E1b1b1a is by far the most common sub-clade of E1b1b in Europe, the Near East and generally outside of Africa. It is also common in North Africa and the Horn of Africa. E1b1b1b is found mainly in the Maghreb, but also in neighbouring areas of Europe such as Iberia.

A third very significant sub-clade of E1b1b1 is E1b1b1c (defined by M123)[11] . It is found both in and out of Africa, but probably had a Near Eastern origin. (See below.)

The fourth major sub-clade of E1b1b1 to be announced (Henn et al. 2008) is defined by M293, an SNP or polymorphism that has been found in parts of Eastern and Southern Africa, and is thought by the authors who announced it to include the majority of E-M35 lineages in sub-Saharan Africa which do not have the mutations M78, M81 or M123.

Smaller E1b1b1 sub-clades recognized are defined by the SNP mutations M281, V6, and P72. The first two at least appear to be unique to the Horn of Africa area.

E1b1b1a (E-M78); formerly E3b1a

E1b1b1a (E-M78) is a commonly occurring subclade, widely distributed in North Africa (especially Egypt and Libya), the Horn of Africa, West Asia "up to Southern Asia"[12], and all of Europe.[13] The European distribution has a frequency peak centered in south Europe and southeastern Europe (13%–16% in southern Italians and 17%–27% in the Balkans) and declining frequencies evident toward western (10% in northern and central Italians), central, and eastern Europe.Based on genetic data, this subclade is thought to have originated in or near the area of Egypt and Libya about 18,600 years ago (17,300 - 20,000 years ago).[14] 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". Prior to, 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, found that 77.6% of 201 male Somalis tested in Denmark were members of this clade. However, were able to study more data, including populations from North Africa who were not represented in the study, and found evidence that the E-M78 lineages in the Horn of Africa were relatively recent branches (see E1b1b1a1b (E-V32) below). They note this as evidence for "a corridor for bidirectional migrations" (conceivably the Nile River Valley) between Egypt and Libya on the one hand and the Horn of 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".

also note evidence for "trans-Mediterranean migrations directly from northern Africa to Europe (mainly in the last 13.0 ky)", and flow from North Africa (around Egypt and Libya) to western Asia between 20.0 and 6.8 ky ago. While there were apparently direct migrations from North Africa to Iberia and Southern Italy (E-V12, E-V22, and E-V65), the majority of E-M78 lineages found in Europe belong to the E-V13 sub-clade which appears to have entered Europe from the Near East, where it apparently originated, via the Balkans (see below).

The division of E1b1b1a into sub-clades such as E-V12, E-V13, etc has largely been the work of Fulvio Cruciani et al. (2004, 2006, 2007), on the basis of STR studies, and more recently the discovery of SNP mutations which define most of the branches with great clarity. This is the basis of the updated phylogenies found in, and ISOGG, which is in turn the basis of the phylogeny given below...

E1b1b1a1 (E-V12)

This sub-clade of E-M78 is the one which appears to have split from the others first (it arose ca. 13.7-15.2 kya[15]). According to, E-V12 likely originated in North Africa, in the area of Libya and Egypt. Undifferentiated E-V12* lineages (not E-V32 or E-M224, so therefore named "E-V12*") are found at especially high levels (44.3%) in Southern Egyptians, but also scattered widely in small amounts in both Northern Africa and Europe, but with very little sign in Western Asia, apart from Turkey[15] . These E-V12* lineages were formerly included (along with many E-V22* lineages[16]) in Cruciani et al.'s original (2004) "delta cluster", which he had defined using DYS profiles. With the discovery of the defining SNP, reported that V-12* was found in its highest concentrations in Egypt, especially Southern Egypt. report a significant presence of E-V12* in neighboring Sudan, including 5/33 Copts and 5/39 Nubians. E-V12* made up approximately 20% of the Sudanese E-M78. They propose that the E-V12 and E-V22 sub-clades of E1b1b1a (E-M78) might have been brought to Sudan from their place of origin in North Africa after the progressive desertification of the Sahara around 6,000–8,000 years ago. Sudden climate change might have forced several Neolithic cultures/people to migrate northward to the Mediterranean and southward to the Sahel and the Nile Valley. The E-V12* paragroup is also observed in Europe (e.g. amongst French Basques) and Eastern Anatolia (e.g. Erzurum Turks).[15]

Sub Clades of E1b1b1a1 (E-V12):

The STR data from concerning E-V12 can be summarized as follows...

HaplotypedescriptionYCAIIaYCAIIbDYS413aDYS413bDYS19DYS391DYS393DYS439DYS460DYS461A10
E-V12*modal192222221310131111913
min18212021111012118811
max1922222315121413121014
number4040404040404040404040
E-V32modal1921222311101312101013
min19192021119121191011
max2022222411111313121114
number3535353535353535353535
All E-V12modal1922222311101311111013
min1819202111912118811
max2022222415121413121114
number7575757575757575757575
E1b1b1a2 (E-V13)

The E-V13 clade is equivalent to the "alpha cluster" of E-M78 reported in, and was first defined by the SNP V13 in . Another SNP is known for this clade, V36, reported in . All known positive tests for V13 are also positive for V36. So E-V13 is currently considered "phylogenetically equivalent" to E-V36.

Within Europe, E-V13 is especially common in the Balkans and some parts of Italy. In different studies, particularly high frequencies have been observed in Kosovar Albanians (45.6%), Albanian speakers of the Former Yugoslavian Republic of Macedonia (34.4%), and Peloponnesian Greeks (47%) . More generally, high frequencies have also been found in other areas of Greece, and amongst Bulgarians, Romanians, and Serbs[18] .

Phylogenetic analysis strongly suggest that these lineages have spread through Europe, from the Balkans in a "rapid demographic expansion"[15] . Before then, the SNP mutation, V13 apparently first arose in West Asia around 10 thousand years ago, and although not widespread there, it is for example found in high levels (>10% of the male population) in Turkish Cypriot and Druze Arab lineages[15] . The Druze are considered a genetically isolated community, and are therefore of particular interest. Their STR DNA signature was actually originally classified in the delta cluster in . This means that Druze E-V13 clustered together with most E-V12 and E-V22, and not with European E-V13, which was mostly in the alpha cluster. This can be summarized in a table format...

haplotypedescriptionYCAIIaYCAIIbDYS413aDYS413bDYS19DYS391DYS393DYS439DYS460DYS461A10
All E-V13modal192123241310131291013
Druze V131192123231310131311912
Druze V132192123231310131311913
All E-V22modal1922222314101312111012
All E-V12*modal192222221310131111913
E-V13 is also found in scattered and small amounts in Libya (in the Jewish community) and Egypt, but this is considered most likely to be a result of migration from Europe or the Near East.[15]

E-V13 and Ancient MigrationsThe apparent movement of E-V13 lineages from the Near East to Europe, and their subsequent rapid expansion, make E-V13 particularly interesting subject for speculation about ancient human migrations.

Early Migration from the Middle East to EuropeThe haplogroup J2b (J-M12) is frequently also discussed in connection to V13, as a haplogroup with a seemingly very similar distribution and pre-history[19] .

says there were at least four major demographic events which have been envisioned for this geographic area:

The distribution and diversity of V13 were thought to be suggestive that it was brought to the Balkans along with early farming technologies, during the Neolithic expansion. However, more recently suggests that the timing for dispersal of European V13 from the Balkans to the rest of Europe may be much more recent, indeed no earlier than 5300 years ago. The authors therefore suggest that this might have been associated with an in situ population increase in the Balkans associated with the Balkan Bronze age, rather than an actual migratory movement of peoples from western Asia. In the next step, "the dispersion of the E-V13 and J-M12 haplogroups seems to have mainly followed the river waterways connecting the southern Balkans to north-central Europe"[15] .

On the other hand, and use age calculation which favor E-V13 dispersal scenarios even older than the Neolithic, in the Mesolithic. Battaglia et al. associate this migration also with the Y haplogroup I-M423.

Greek Soldiers in Pakistan:Both E-V13 and J-M12 have also been used in studies seeking to find evidence of a remaining Greek presence in Afghanistan and Pakistan, going back to the time of Alexander the Great.

Roman soldiers in Britain:Significant frequencies of E-V13 have also been observed in towns in Wales, England and Scotland. The old trading town of Abergele on the northern coast of Wales in particular showed 7 out of 18 local people tested were in this lineage (approximately 40%), as reported in . attributes the overall presence of E-V13 in Great Britain, especially in areas of high frequency, to settlement during the 1st through 4th centuries CE by Roman soldiers from the Balkan peninsula. Bird proposes a connection to the modern region encompassing Kosovo, southern Serbia, northern Macedonia and extreme northwestern Bulgaria (a region corresponding to the Roman province of Moesia Superior), which was identified by as harboring the highest frequency worldwide of this sub-clade[20] .

However, according to data published so far[21], E-V13 appears to be notably absent in Central England, a fact which Bird (2007) suggests reflects a genuine population replacement of Romano-British people with Anglo-Saxons:

Phoenician Traders in the Mediterranean: have also suggested E1b1b haplotypes (amongst others) to be a sign of Phoenician influence around the Mediterranean[22] .

Sub Clades of E1b1b1a2 (E-V13)Although most E-V13 individuals do not show any downstream SNP mutations, and are therefore categorized as E1b1b1a2* (E-V13*) there are two recognized sub-clades, both of which may be very small. These are one of two cases where remarked that at the time of that article, it was not certain that the two clades were truly separate ("the positions of these mutations have not been resolved because of a lack of a DNA sample containing the derived state at V27").

E1b1b1a3 (E-V22)

This clade comprises most of those classified in the "delta cluster" of . later noted that "E-V22 and E-V12* chromosomes are intermingled and not clearly differentiated by their microsatellite haplotypes".

This sub-clade of E-M78 is "relatively common"[15] in the Horn of Africa and Egypt, with higher microsatellite variance (0.35 vs. 0.46, respectively) in Egypt. In the article announcing this first information, described it as uncommon in Western Asia and they proposed Northeast Africa as this sub-clade's likely place of origin. also reported a significant presence in neighboring Sudan, making up about 30% of the diverse range of the country's E-M78 lineages in their study, including 8 out of 26 Fulani (about 31%), a widely-dispersed pastoral people[23] . E-V22 was also present in much smaller frequencies amongst the Shilluk (2 of 15 samples, 13%) and Dinka (3 of 26, 8%) Nilotes of Southern Sudan. Hassan et al. suggest that E-V22, like E-V12, might have entered Sudan from North Africa "after the progressive desertification of the Sahara around 6,000–8,000 years ago". They add that the gene flow to Sudan "is not only recent (Holocene onward) but also largely of focal nature", and that "most speakers of Nilo-Saharan languages, the major linguistic family spoken in the country, show very little evidence of gene flow and demonstrate low migration rate, with exception of the Nubians, who appear to have sustained considerable gene flow from Asia and Europe together with the Beja."

Other frequencies reported by include Asturians (4.44% out of 90 people), Sicilians (4.58% out of 153 people), Moroccan Arabs (7.27%, 55 people) and Moroccan Jews (8%, 50 people), Istanbul Turkish (5.71% out of 35 people), and Palestinians (6.9% out of 29 people). found a 6.7% presence in the UAE.

Sub Clades of E1b1b1a3 (E-V22): There are two recognized sub-clades, which are apparently separate, although remarked that at the time of that article, "the positions of these mutations have not been resolved because of a lack of a DNA sample containing the derived state at [...] V19".

E1b1b1a4 (E-V65)

This sub-clade, equivalent to the previously classified "beta cluster", is found in high levels in the Maghreb regions of far northern Africa. report levels of about 20% amongst Libyan Arab lineages, and about 30% amongst Morrocan Arabs. It appears to be less common amongst Berbers, but still present in levels of >10%. The authors suggest a North African origin for this lineage. In Europe, only a few individuals were found in Italy and Greece. The results from the article can be summarized as follows...

E-V65YCAIIaYCAIIbDYS413aDYS413bDYS19DYS391DYS393DYS439DYS460DYS461A10
modal1921212313101310101113
min19202022111013109912
max2121222314111411111213
number3838383838383838383838

studied the beta cluster in Europe. They found small amounts in Southern Italy, but also traces in Cantabria, Portugal and Galicia, with Cantabria having the highest level in Europe in their study, at 3.1% (out of 161 people).

E1b1b1a5 (E-VM521)

This sub-clade's discovery was announced in They found 2 out of 92 Greeks to have this mutation.

E1b1b1b (E-M81); formerly E3b1b, E3b2

E1b1b1b (E-M81) is the most common Y chromosome haplogroup in North Africa, dominated by its sub-clade E-M183. It is thought to have originated in North Africa 5,600 years ago. 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 reports that approximately 76% are M81+.

In Europe, E-M81 is found everywhere but mostly in the Iberian Peninsula, where it is more common than E-M78 unlike in the rest of Europe[24] at an average frequency of 4-5.6%, with frequencies reaching 10% in Western Andalusia, Northwest Castile and 13 % in Cantabria[25] . The highest frequency of this clade found so far in Europe has been observed at 40% the Pasiegos from Cantabria.

It is also common in Sicily, and in slightly lower frequencies in continental Italy (especially near Lucera) and France, possibly due to ancient migrations during the Islamic, Roman, and Carthaginian empires, as well as the influence of Sephardic Jews.

As a result of its old world distribution, this sub-clade is found throughout Latin America, for example 6.1% (8 out of 132) in Cuba, ; 5.4% (6 out of 112) in Brazil (Rio de Janeiro), [26] and among Hispanic men in the United States.[27]

In smaller numbers, E-M81 men can be found in Sudan, 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):

E1b1b1c (E-M123); formerly E3b1c, E3b3

This sub-clade of E1b1b1 (E-M35) is mostly known for its major sub-clade E1b1b1c1 (E-M34), which dominates this clade[28] . However, earlier studies did not test for E-M34.

Concerning E-M123* (tested and definitely without E-M34) 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, found 2 E-M123* individuals, both in Northern Portugal out of 109 people tested there. In a 553 person study of Portugal, 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. found one individual out of 146 Jordanians. found none amongst the significant presence of E-M34 they found in their study of the UAE, Yemen and Qatar.

Concerning E-M123 without checking for the M-34 SNP found E-M123 examples in Greece, the Republic of Macedonia, and Roumania. also found examples in Portugal, and found one sample in Somalia. 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. 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. (tested for E-M34 in Oman and found 7.7% to be E-M34+, with no E-M123*.) found 4 out of 20 tested Israeli Jews of Libyan ancestry to be M123+.

Concerning E1b1b1c1 (E-M34) According to, 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, 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 E-M34 was found in several parts of Iberia, but most strikingly about 10% in Galicia. 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, 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. 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%).

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.[29] wrote that:

Sub Clades of E1b1b1c1 (E-M34):

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. (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, 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.

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.[30]

See also

References

External links

Notes and References

  1. https://www3.nationalgeographic.com/genographic/atlas.html National Geographic's Genographic Project - Haplogroup E3b (M35)
  2. The only major linguist to argue for a Levantine origin of Afro-Asiatic is Alexander Militarev, but some archaeologists who argue for Neolithic "demic diffusion" such as Peter Bellwood argue that there is no significant archaeological evidence to support such an assumption, and that the subsequent spread of Semitic languages, in particular, has likely erased much of the original phylogenetic geography of the Afro-Asiatic language family, making it difficult to pinpoint the geographical source of this linguistic phylum .
  3. . See Table 2.
  4. . "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."
  5. "Both phylogeography and microsatellite variance suggest that E-P2 and its derivative, E-M35, probably originated in eastern Africa. 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*."
  6. "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).
  7. For E1b1b (M-215) Cruciani et al. reduced their 2004 estimates from 25,600 in 2004 to 22,400 in 2007, re-calibrating the same data.
  8. 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.
  9. "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."

  10. [https://www3.nationalgeographic.com/genographic/atlas.html Atlas of the Human Journey>Genetic Markers>M35]. E3b. The man who gave rise to marker M35 was born around 20,000 years ago in the Middle East. His descendants were among the first farmers and helped spread agriculture from the Middle East into the Mediterranean region.. The Genographic Project. 2008. 2008-09-05.
  11. Web site: ISOGG: Y-DNA Haplogroup E and its Subclades - 2008. isogg.org. 2008-05-17. 2008-05-05.
  12. 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".

  13. "The human Y chromosome haplogroup E-M78 (E3b1a) occurs commonly and is distributed in northern and eastern Africa, western Asia, and all of Europe."

  14. use the term "North-eastern Africa" for Libya and Egypt, as shown in their Table 1.
  15. See Figure 1.
  16. "E-V22 and E-V12* chromosomes are intermingled and not clearly differentiated by their microsatellite haplotypes". In the same authors show that a branch of E-V13 found amongst the Druze Arabs is also in the delta cluster. (Contrast the data tables of and .)

  17. Fig. 2/C

  18. ,,,
  19. See especially
  20. Doubts about this line of reasoning have been expressed because (a) new data appearing in indicates other high concentrations in Greece and (b) the data in show that the area with the highest frequency does not have the highest diversity, implying that V13 arrived there more recently than in Greece.
  21. Bird uses three sources:, and . Neither Capelli nor Weale have data from the area in the English Midlands where Bird suggests that there is a lack of E1b1b. In 2006 Bird mentioned that there were 193 Central English haplotypes in Sykes.
  22. write that "PCS3+ scores strongly as a Phoenician colonization candidate and is strongly associated with the SNP haplogroup E3b, but it does not show the wide geographic coverage that the other PCS+s demonstrate. It represents the strongest of the lower-coverage STR+s." However the authors admit that the number of STRs they use does not even distinguish between major haplogroups such as E and J in a clear way. They also admit that they could not design the testing so as to identify the influence of the Jewish diaspora.
  23. in a study of Guinea Bissau, showed that the Fulani there are about 10% E-M78. Note that this study did not test specifically for V12 or V22, so the E-M78 may have a different exact breakdown of diversity as well as a lower frequency.
  24. , shows an average frequency of 4% in the Iberian Peninsula with frequencies reaching 10% in Western Andalusia and Northwest Castile, see table.
  25. ,,,
  26. "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.",
  27. 2.4% (7 out of 295) among Hispanic men from California and Hawaii,
  28. 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.
  29. ) Table 1
  30. 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