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→‎E1b1b1a (E-M78); formerly E3b1a: removing awful English which came from an abstract. See discussion page, where this was discussed to death
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===E1b1b1a (E-M78); formerly E3b1a===
===E1b1b1a (E-M78); formerly E3b1a===


This clade is thought to have originated in [[North Africa]] (around [[Egypt]] and [[Libya]]) about 18,600 years ago (17,300 - 20,000 years ago).<ref>Cruciani et al. (2007) [http://dirkschweitzer.net/E3b-papers/MolecularBiologyandEvolution-07-24-6-1300.pdf Tracing Past Human Male Movements in Northern/Eastern Africa and Western Eurasia: New Clues from Y-Chromosomal Haplogroups E-M78 and J-M12], ''Molecular Biology and Evolution'', 24:1300-1311. (pp.11-13 & 29-30) See Table 1 for definition of areas sampled under "North-eastern Africa</ref> E1b1b1a (E-M78) occurs commonly and is distributed in [[North Africa]], the [[Horn of Africa]], [[West Asia]], and all of [[Europe]].<ref name=Cruciani2006/>
This clade is thought to have originated in [[North Africa]] (around [[Egypt]] and [[Libya]]) about 18,600 years ago (17,300 - 20,000 years ago).<ref>Cruciani et al. (2007) [http://dirkschweitzer.net/E3b-papers/MolecularBiologyandEvolution-07-24-6-1300.pdf Tracing Past Human Male Movements in Northern/Eastern Africa and Western Eurasia: New Clues from Y-Chromosomal Haplogroups E-M78 and J-M12], ''Molecular Biology and Evolution'', 24:1300-1311. (pp.11-13 & 29-30) See Table 1 for definition of areas sampled under "North-eastern Africa</ref> E1b1b1a (E-M78). E-M78 male lineages are common in [[North Africa]], the [[Horn of Africa]], [[West Asia]], and all of [[Europe]].<ref name=Cruciani2006/>
Prior to Cruciani et al.'s study from 2007, the Horn of Africa had also been proposed as a possible place of origin of E-M78 by Semino et al. (2004)<ref name=Semino2004/> given 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 [[Somali people|Somalis]] tested in Denmark carried the 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. study from 2004, and found evidence that the E-M78 lineages in the Horn of Africa were relatively recent branches (see E1b1b1a1b (E-V32) [[E3b#E1b1b1a1_.28E-V12.29|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".
Prior to Cruciani et al.'s study from 2007, the Horn of Africa had also been proposed as a possible place of origin of E-M78 by Semino et al. (2004)<ref name=Semino2004/> given 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 [[Somali people|Somalis]] tested in Denmark carried the 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. study from 2004, and found evidence that the E-M78 lineages in the Horn of Africa were relatively recent branches (see E1b1b1a1b (E-V32) [[E3b#E1b1b1a1_.28E-V12.29|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".

Revision as of 19:24, 30 October 2008

Haplogroup E1b1b or E-M215
Possible time of originapprox 26,000 years BP
Possible place of originEast Africa[1] or Middle East[2]
AncestorE1b1 or E-P2
DescendantsE1b1b1 or E-M35
Defining mutationsM215, 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.[3][4][5] 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. It was only in Cruciani et. al.'s 2004 article[1] that M215 was shown to be older than M35, because there are some lineages which have the M215 mutation, but not 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[6], North Africa[6], parts of Eastern and Southern Africa[1], West Asia[1], and Europe[7] (especially the Mediterranean and the Balkans[1][6][8]).

A significant proportion of all Jewish male lines are made up of a wide variety of E1b1b1 (E-M35) sub-clades. Behar et al. (2003) found only haplogroup J lineages in higher numbers amongst Ashkenazim.[9] The authors also found E1b1b to be, along with haplogroup J, one of the major founding lineages among Ashkenazi Jews.[10] E1b1b is observed in over 22.8% of Ashkenazis[11] and 30% of Sephardim.[6] 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.[12]

Other Names, and history of the classification

The current phylogenetic terminology "E1b1b" and "E1b1b1" was proposed in 2008 by Karafet et. al.[4]. This 2008 paper was intended to be an update of the 2002 "Y Chromosome Consortium"(YCC)[5]. 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 2002, both E1b1b and E1b1b1, not yet distinguished at that time, had been referred to as Hg21 (Haplogroup 21) within Zerjal et al. (1999)'s nomenclature[13], or as Eu4 according to Semino et al. (2000)'s classification[14].

They were also within Underhill et al. (2001)'s "Group III" [15].

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, Bosch et al. (2001)[16], Semino et al. (2004)[6][17], Cruciani et al. (2004[1][18], 2006[19], and 2007[20]), 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.[4] There are different techniques available for such estimates, and a considerable range of possibilities, but the most recent estimates of Cruciani et al. (2007) are around 24,000 years ago for E-M215[21] or E-M35.[22]

Referencing Cruciani et al.'s 2004 study, Coffman-Levy (2005) writes that E1b1b1 (E-M35) "arose in East Africa". However, she adds that this haplogroup is "often incorrectly described as “African,” leaving a misimpression regarding the origin and complex history of this haplogroup", and that a lot of misinformation about this haplogroup also continues to pervade the public and media.[23]

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.[3][24]

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.

Subclades of E1b1b1 (E-M35)

As mentioned above, nearly all E1b1b lineages are within E1b1b1 (defined by 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*) 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 and generally outside of Africa. It is also common in the Near East. And together, E1b1b1a (E-M78) and E1b1b1b (E-M81) form a very significant part of all male lineages in Northeast and North Africa.

A third very significant sub-clade of E1b1b1 is E1b1b1c (defined by M123)[3]. It is found both in and out of Africa, but probably had a Near Eastern origin. (See below.)

A fourth major sub-clade of E1b1b1 to be announced much later than the others (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 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.

E1b1b1a (E-M78); formerly E3b1a

This clade is thought to have originated in North Africa (around Egypt and Libya) about 18,600 years ago (17,300 - 20,000 years ago).[25] E1b1b1a (E-M78). E-M78 male lineages are common in North Africa, the Horn of Africa, West Asia, and all of Europe.[19]

Prior to Cruciani et al.'s study from 2007, the Horn of Africa had also been proposed as a possible place of origin of E-M78 by Semino et al. (2004)[6] given 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 carried the 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. study from 2004, 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".

Cruciani et al. (2007) 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 Karafet et al. (2008), 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[20]). According to Cruciani et al. (2007), E-V12 likely originated in North Africa. Undifferentiated E-V12* lineages (not E-V12 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[20]. These E-V12* lineages were formerly included (along with many E-V22* lineages[26]) in Cruciani et al.'s original (2004) "delta cluster", which he had defined using DYS profiles. With the discovery of the defining SNP, Cruciani et al. (2007) reported that V-12* was found in its highest concentrations in Egypt, especially Southern Egypt. Hassan et al. (2008) report a significant presence of E-V12* in neighboring Sudan. 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.[27] The E-V12* paragroup is also observed in Europe (e.g. amongst French Basques) and Eastern Anatolia (e.g. Erzurum Turks).[20]

Sub Clades of E1b1b1a1 (E-V12):
  • E1b1b1a1a (E-M224). This clade is apparently rare. It is discussed in Underhill et al. (2000, 2001) and Cruciani et al. (2006). Cruciani et al. found no exemplars in their 2007 study.
  • File:Cadow-pr2.gif
    Somali man in traditional white fez. A large majority of Somali male lines are in the E1b1b1a1b (E-V32) lineage, which originated in North Africa, around Egypt and Libya.
    E1b1b1a1b (E-V32). Cruciani et al. (2007) suggest that this sub-clade of E-V12 originated in North Africa (around Egypt and Libya), and then subsequently expanded further south into the Horn of Africa, where it is now prevalent.[20] Before the discovery of V32, Cruciani et al. (2004) referred to the same lineages as the "gamma cluster", which was estimated to have arisen about 8,500 years ago. They stated that "the highest frequencies in the three Cushitic-speaking groups: the Borana from Kenya (71.4%), the Oromo from Ethiopia (32.0%), and the Somali (52.2%). Outside of eastern Africa, it was found only in two subjects from Egypt (3.6%) and in one Arab from Morocco". Sanchez et al. (2005) found it extremely prominent in Somali men and stated that "the male Somali population is a branch of the East African population – closely related to the Oromos in Ethiopia and North Kenya (Boranas)" and that their gamma cluster lineages "probably were introduced into the Somali population 4000–5000 years ago". Hassan et al. (2008) in their study observed this to be the most common of the sub-clades of E-M78 found in Sudan, especially among the Beja, Masalit, and Fur.
E1b1b1a2 (E-V13)

(All known positive tests for V13 are also positive for V36. So E-V13 is currently considered "phylogenetically equivalent" to E-V36.)

"Haplogroup E-V13 is the only lineage that reaches the highest frequencies out of Africa"[20].

E1b1b1a2 (E-V13) is found in Europe "with a clinal pattern of frequency distribution from the southern Balkan peninsula (19.6%) to western Europe (2.5%). The same haplogroup is also present at lower frequencies in Anatolia (3.8%), the Near East (2.0%), and the Caucasus (1.8%). In Africa, haplogroup E-V13 is rare, being observed in northern Africa at a low frequency (0.9%)."[20]

This clade is the most common sub-clade of E-M78 in Europe,[28] especially in the Balkans where high concentrations are reported amongst Albanians, Macedonians, Greeks, Bulgarians, Romanians, and Serbs.[1][8][29][30] Particularly high frequencies have been observed in Kosovar Albanians (45.6%)[29] and Peloponnesian Greeks (47%).[6]

High 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 shows a frequency of nearly 39% of its male population in this lineage. Bird (2007) 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.[28] Peričić et al. (2005) likewise have identified the modern region encompassing Kosovo, southern Serbia, northern Macedonia and extreme northwestern Bulgaria (a region corresponding to the Roman province of Moesia Superior) as harboring the highest frequency worldwide of this sub-clade.[29] However, E-V13 is notably absent in Central England, a fact which Bird (2007) suggests reflects a genuine population replacement of Romano-British people with Anglo-Saxons in the form of either:[28]

  • a massive displacement of the native Romano-British population by invasion or,
  • the substantial genetic replacement of Romano-British Y-DNA through an elite dominance ("apartheid") model.

The E-V13 clade is equivalent to Cruciani et al's (2004) "alpha cluster" and phylogenetic analysis strongly suggest that these lineages have spread through Europe, from the Balkans in a "rapid demographic expansion"[20]. 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[20] - with the latter being considered a genetically isolated community, and therefore of particular interest. It 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.[20]

Druze Sheikh (ˤUqqāl) wearing religious dress. Many Druze men are in the E1b1b1a2 (E-V13) lineage most commonly found in Europe, especially the Balkans.

The 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. The haplogroup J2b (J-M12) is frequently also discussed in connection to V13, as a haplogroup with a seemingly very similar distribution and pre-history[31].

Cruciani et al. (2007) 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[6]. However, Cruciani et al. (2007) 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"[20][29].

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, with some success[7][6].

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:
  • E1b1b1a2a. Defined by V27.
  • E1b1b1a2b. Defined by P65.
E1b1b1a3 (E-V22)

According to 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".

This sub-clade of E-M78 is "relatively common"[20] in the Horn of Africa and Egypt, with higher microsatellite variance (0.35 vs. 0.46, respectively) in Egypt. Cruciani et al. (2007) propose Northeast Africa as this sub-clade's likely place of origin. Hassan et al. (2008) also reported a significant presence in neighboring Sudan. This clade comprises most of those classified in Cruciani et al's earlier (2004) "delta cluster".

Sub Clades of E1b1b1a3 (E-V22): There are two recognized sub-clades:
  • E1b1b1a3a. Defined by M148.
  • E1b1b1a3b. Defined by 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. Cruciani et al. (2007) 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.

E1b1b1b (E-M81); formerly E3b1b, E3b2

Libyan Arabs. E1b1b1b (E-M81): The most common Y haplogroup among North African Arabs and Berbers

E1b1b1b (E-M81) is the most common Y chromosome haplogroup in North Africa. It is thought to have originated in North Africa 5,600 years ago.[32][1] 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 haplogroup is also found in significant amounts in the Iberian Peninsula, Italy and France[1], probably due to ancient migrations during the Islamic, Roman, and Carthaginian empires, as well as the influence of Sephardic Jews.[33] This sub-clade of E1b1b has also been observed in 40% of Europeans in the Pasiegos from Cantabria.[1]

Individuals with the defining marker for this clade, M81, also test positive, in tests so far, for M183.

There are two recognized sub-clades, although at this time it seems that the majority of E-M81 is "E-M81*", meaning that they are not in any of the known sub-clades.

Sub Clades of E1b1b1b (E-M81):
  • E1b1b1b1 (E-M107).
  • E1b1b1b2 (E-M183). As of 23rd October 2008, the SNP M165 is currently considered to define a subclade, "E1b1b1b2a"[3].

E1b1b1c (E-M123); formerly E3b1c, E3b3

  • Jewish man seated. The E1b1b1c (E-M123) sub-clade of E1b1b is quite common among Jews.

This sub-clade of E1b1b1 (E-M35) is mostly known for its major sub-clade E1b1b1c1 (E-M34). According to Cruciani et al. (2004), E-M34 is found at very small frequencies in North Africa and southeastern Europe, and has its highest concentration in Ethiopia and the Near East. However, because the diversity is apparently low in Ethiopia, the authors suggest that E-M34 was likely introduced into Ethiopia from the Near East. This article located one E-M123* individual in Bulgaria after testing 3401 individuals from five continents, and Underhill et al. (2000) located one individual in Central Asia. Bosch (2006) found examples in Greece, Macedonia, and Roumania. Beleza (2006) also found examples in Portugal, and Sanchez et al. (2005) found one sample in Somalia.

E1b1b1c (E-M123) is quite common among both Ashkenazi and Sephardic Jews, accounting for over 10% of all male lines.[34] Coffman-Levy (2005)[12] 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.

Sub Clades of E1b1b1c1 (E-M34):
  • E1b1b1c1a. Defined by SNP mutation M84, with M136 defining a sub-clade, E1b1b1c1a1 as of October 2008[3].
  • E1b1b1c1b. Defined by SNP mutation M290

E1b1b1d (E-M281)

The SNP mutation which defines this sub-clade of E-M35, M281, was discussed in Underhill et al. (2000) and Semino et al. (2002), 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) identified a significant presence of these lineages in Ethiopia, but apparently not elsewhere.

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.[35] 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.[36]

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[37].

See also

Notes

  1. ^ a b c d e f g h i j Cruciani et al. (2004), Phylogeographic Analysis of Haplogroup E3b (E-M215) Y Chromosomes Reveals Multiple Migratory Events Within and Out Of Africa, American Journal of Human Genetics, 74: 1014-1022.
  2. ^ National Geographic's Genographic Project - Haplogroup E3b (M35)
  3. ^ a b c d e ISOGG: Y-DNA Haplogroup E and its Subclades - 2008 Cite error: The named reference "isogg" was defined multiple times with different content (see the help page).
  4. ^ a b c Karafet et al. (2008) [http://www.genome.org/cgi/content/abstract/gr.7172008v1 Abstract|title= New Binary Polymorphisms Reshape and Increase Resolution of the Human Y-Chromosomal Haplogroup Tree}}
  5. ^ a b Y Chromosome Consortium (2002)Abstract|title= A Nomenclature System for the Tree of Human Y-Chromosomal Binary Haplogroups
  6. ^ a b c d e f g h i Semino et al. (2004) Origin, Diffusion, and Differentiation of Y-Chromosome Haplogroups E and J: Inferences on the Neolithization of Europe and Later Migratory Events in the Mediterranean Area, American Journal of Human Genetics, 74: 1023–1034.
  7. ^ a b Firasat et al. (2007), Y-chromosomal evidence for a limited Greek contribution to the Pathan population of Pakistan, European Journal of Human Genetics (2007) 15, 121–126: "E3b is common in Europe"
  8. ^ a b Rosser et al. (2000), Y-Chromosomal Diversity in Europe Is Clinal and Influenced Primarily by Geography, Rather than by Language, American Journal of Human Genetics, 67: 1526-1543.
  9. ^ Behar et al. (2003), Contrasting patterns of Y chromosome variation in Ashkenazi Jewish and host non-Jewish European populations, Human genetics, 2004 Mar;114(4):354-65. See Table 2.
  10. ^ Behar et al. (2003), Contrasting patterns of Y chromosome variation in Ashkenazi Jewish and host non-Jewish European populations, Human genetics, 2004 Mar;114(4):354-65.: "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."
  11. ^ Nebel et al. (2001), The Y Chromosome Pool of Jews as Part of the Genetic Landscape of the Middle East, American Journal of Human Genetics 2001 November; 69(5): 1095–1112.
  12. ^ a b Ellen Coffman-Levy (2005) A Mosaic of People: The Jewish Story and a Reassessment of the DNA Evidence Journal of Genetic Genealogy 1:12-33.
  13. ^ Zerjal et al. (1999), The use of Y-chromosomal DNA variation to investigate population history; in Papiha SS, Deka R, Chakraborty R (eds): Genomic diversity: applications in human population genetics. New York: Kluwer Academic/Plenum Publishers, 1999, pp 91–101.
  14. ^ Semino et al. (2000)The Genetic Legacy of Paleolithic Homo sapiens sapiens in Extant Europeans: A Y Chromosome Perspective.
  15. ^ Underhill et al. (2001)The phylogeography of Y chromosome binary haplotypes and the origins of modern human populations. Ann Hum Genet. 65:43–62.
  16. ^ Bosch E, Calafell F, Comas D, Oefner PJ, Underhill PA, Bertranpetit J (2001) High-resolution analysis of human Y-chromosome variation shows a sharp discontinuity and limited gene flow between north-western Africa and the Iberian Peninsula. Am J Hum Genet 68:1019–1029
  17. ^ Semino et al. (2004): "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*."
  18. ^ 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).
  19. ^ a b Cruciani et al. (2006), Molecular Dissection of the Y Chromosome Haplogroup E-M78 (E3b1a): A Posteriori Evaluation of a Microsatellite-Network-Based Approach Through Six New Biallelic Markers, Human Mutation
  20. ^ a b c d e f g h i j k l Cruciani et al. (2007) Tracing Past Human Male Movements in Northern/Eastern Africa and Western Eurasia: New Clues from Y-Chromosomal Haplogroups E-M78 and J-M12, Molecular Biology and Evolution, 24:1300-1311. Cite error: The named reference "Cruciani2007" was defined multiple times with different content (see the help page).
  21. ^ 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.
  22. ^ 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.
  23. ^ Ellen Coffman-Levy (2005) A Mosaic of People: The Jewish Story and a Reassessment of the DNA Evidence Journal of Genetic Genealogy 1:12-33. From pp.22-23: "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."
  24. ^ The Genographic Project (2008). "Atlas of the Human Journey>Genetic Markers>M35". Retrieved 2008-09-05. 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. {{cite journal}}: |section= ignored (help); Cite journal requires |journal= (help)
  25. ^ Cruciani et al. (2007) Tracing Past Human Male Movements in Northern/Eastern Africa and Western Eurasia: New Clues from Y-Chromosomal Haplogroups E-M78 and J-M12, Molecular Biology and Evolution, 24:1300-1311. (pp.11-13 & 29-30) See Table 1 for definition of areas sampled under "North-eastern Africa
  26. ^ Cruciani et al. (2006) "E-V22 and E-V12* chromosomes are intermingled and not clearly differentiated by their microsatellite haplotypes"
  27. ^ Hassan et al. (2008), Y-Chromosome Variation Among Sudanese: Restricted Gene Flow, Concordance With Language, Geography, and History AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY,
  28. ^ a b c Bird (2007), Haplogroup E3b1a2 as a Possible Indicator of Settlement in Roman Britain by Soldiers of Balkan Origin, Journal of Genetic Genealogy. 3(2):26-46, 2007: "E3b1a-M78, of which E3b1a2 is, by far, the most common subclade in Europe"
  29. ^ a b c d Peričic et. al. (2005), High-Resolution Phylogenetic Analysis of Southeastern Europe Traces Major Episodes of Paternal Gene Flow Among Slavic Populations, Molecular Biology and Evolution 2005 22(10):1964-1975
  30. ^ King et al. (2008) Differential Y-chromosome Anatolian Influences on the Greek and Cretan Neolithic Annals of Human Genetics 72,205–214
  31. ^ See especially Cruciani et al. (2007)
  32. ^ Arredi et al. (2004), A Predominantly Neolithic Origin for Y-Chromosomal DNA Variation in North Africa, American Journal of Human Genetics, 75: 338–345.
  33. ^ Gonçalves et al. (2005), Y-chromosome Lineages from Portugal, Madeira and Açores Record Elements of Sephardim and Berber Ancestry, Annals of Human Genetics (2005) 69,443–454
  34. ^ Semino et al. 2004 Table 1
  35. ^ Henn et al. (2008), Y-chromosomal evidence of a pastoralist migration through Tanzania to southern Africa, Proc Natl Acad Sci U S A., 2008 Aug 5;105(31):10693-8.
  36. ^ Henn et al. (2008), Y-chromosomal evidence of a pastoralist migration through Tanzania to southern Africa, Proc Natl Acad Sci U S A., 2008 Aug 5;105(31); p.10695
  37. ^ 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

References

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