Haplogroup E1b1b (Y-DNA)


Haplogroup E1b1b (Y-DNA)

Infobox haplogroup
name = E1b1b or E-M215
origin-place = East Africa or Middle East [https://www3.nationalgeographic.com/genographic/atlas.html National Geographic's Genographic Project - Haplogroup E3b (M35)] ]
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 [http://www.isogg.org/tree/ISOGG_HapgrpE08.html ISOGG: Y-DNA Haplogroup E and its Subclades - 2008] ] 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] Y Chromosome Consortium (2002) [http://genome.cshlp.org/cgi/content/abstract/12/2/339 Abstract|title= A Nomenclature System for the Tree of Human Y-Chromosomal Binary Haplogroups] ] .

In nearly all discussion, E1b1b is equivalent to it's sub-clade, E1b1b1 (E-M35), which contains nearly all of E1b1b. It was only in Cruciani et. al.'s 2004 articleCruciani et al. (2004), [http://www.familytreedna.com/pdf/hape3b.pdf 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.] 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, North Africa, parts of Eastern and Southern Africa, West Asia, and Europe (especially the Mediterranean and the Balkans).

A significant proportion of all types of Jewish male lines are made up of an interestingly wide variety of E1b1b1 (E-M35) sub-clades. Behar et al. (2003) found only haplogroup J lineages in higher numbers amongst Ashkenazim. [Behar et al. (2003), [http://www.familytreedna.com/pdf/Behar_contrasting.pdf 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.] The authors also found E1b1b to be, along with haplogroup J, one of the major founding lineages among Ashkenazi Jews.Behar et al. (2003), [http://www.familytreedna.com/pdf/Behar_contrasting.pdf 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."] E1b1b is observed in over 22.8% of Ashkenazis [Nebel et al. (2001), [http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B8JDD-4R29JBW-M&_user=10&_rdoc=1&_fmt=&_orig=search&_sort=d&view=c&_acct=C000050221&_version=1&_urlVersion=0&_userid=10&md5=311b0abad8d37d35a8776ade5baa84c4 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.] 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.Ellen Coffman-Levy (2005) [http://www.jogg.info/11/coffman.pdf A MOSAIC OF PEOPLE: THE JEWISH STORY AND A REASSESSMENT OF THE DNA EVIDENCE] Journal of Genetic Genealogy 1:12-33.]

Other Names, and history of the classification

The current phylogenetic terminology "E1b1b" and "E1b1b1" was proposed in 2008 by Karafet et. al.. This 2008 paper was intended to be an update of the 2002 "Y Chromosome Consortium"(YCC). The YCC first formalized the original phylogenetic nomenclature - "E3b" and "E3b1" - 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 [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.] , or as Eu4 according to Semino et al. (2000)'s classificationSemino et al. (2000) [http://hpgl.stanford.edu/publications/Science_2000_v290_p1155.pdf The Genetic Legacy of Paleolithic Homo sapiens sapiens in Extant Europeans: A Y Chromosome Perspective.] ] .

They were also within Underhill et al. (2001)'s "Group III" Underhill et al. (2001) [http://hpgl.stanford.edu/publications/AHG_2001_v65_p43.pdf The phylogeography of Y chromosome binary haplotypes and the origins of modern human populations. Ann Hum Genet. 65:43–62.] ] .

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)Bosch E, Calafell F, Comas D, Oefner PJ, Underhill PA, Bertranpetit J (2001) [http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pubmed&pubmedid=11254456 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 ] , Semino et al. (2004)Semino et al. (2004) [http://www.pubmedcentral.nih.gov/articlerender.fcgi?artid=1181965 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.] [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*."] , Cruciani et al. (2004 [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).] , 2006Cruciani et al. (2006), [http://www3.interscience.wiley.com/homepages/38515/pdf/916.pdf 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"] , and 2007Cruciani 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.] ), point to evidence that not only E1b1b (E-M215), but also both it's 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 Cruciani et al. (2007) are around 24,000 years ago for E-M215 [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.] or E-M35. [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.]

However, according to Coffman (2005), E1b1b is "often incorrectly described as “African,” leaving a misimpression regarding the origin and complex history of this haplogroup". A lot of misinformation about this haplogroup also continues to pervade the public and media.

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. [cite journal|url=https://www3.nationalgeographic.com/genographic/atlas.html|section=E3b|quote=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.|title=Atlas of the Human Journey>Genetic Markers>M35|author=The Genographic Project|date=2008|accessdate=2008-09-05]

As E1b1b1 dispersed, most major sub-branches of E1b1b1 are thought to have originated in North Africa, the Horn of Africa, and Western Asia.

ubclades 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, but still less well-known, sub-clade of E1b1b1 is E1b1b1c (defined by M123)cite web|url=http://www.isogg.org/tree/ISOGG_HapgrpE08.html|title= ISOGG: Y-DNA Haplogroup E and its Subclades - 2008|publisher=isogg.org|accessdate=2008-05-17|date=2008-05-05] .

A 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 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

Estimations of age are difficult and vary greatly, but E-M78's age has been estimated at about 18,600 years ago. This clade is thought to have originated in Northeast Africa, around Egypt and Libya (Cruciani et al. 2007). [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] The most recent estimates are as follows:

The geographic and quantitative analyses of haplogroup and microsatellite diversity is strongly suggestive of a northeastern African origin of E-M78, with a corridor for bidirectional migrations between northeastern and eastern Africa (at least 2 episodes between 23.9–17.3 ky and 18.0–5.9 ky ago), trans-Mediterranean migrations directly from northern Africa to Europe (mainly in the last 13.0 ky), and flow from northeastern Africa to western Asia between 20.0 and 6.8 ky ago. (Cruciani et. al. 2007)

It should be noted that the migrations to Europe mentioned above are the ones which are basically localized to Iberia and Southern Italy. Concerning the far more important part of E-M78 in Europe, see below concerning sub-clade E-V13.

E1b1b1a has been further divided into subclades by Fulvio Cruciani et al. (2004, 2006, 2007), on the basis of the following SNP mutations, and this is the basis of the updated phylogenies found in Karafet 2008, and [http://www.isogg.org/tree/ISOGG_HapgrpE08.html ISOGG] , as follows...

E1b1b1a1 (E-V12)

This is the oldest sub-clade of E-M78, found mainly in Southern Egyptians (arose ca. 15.2 kya). According to Cruciani et al. (2007), this lineage likely originated in North Africa. Lineages with this SNP mutation, but without any of the known sub-clade mutations such as V32 (so "V12*"), were formerly included 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.Hassan et al. (2008), [http://dirkschweitzer.net/E3b-papers/Hassan-Sudan-2008-AJPA.pdf Y-Chromosome Variation Among Sudanese: Restricted Gene Flow, Concordance With Language, Geography, and History] AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY,] The E-V12* paragroup is also observed in Europe (e.g. amongst French Basques) and Eastern Anatolia (e.g. Erzurum Turks).

:Sub Clades of E1b1b1a1 (E-V12):::*E1b1b1a1a (E-M224). This clade is apparently rare. It is discussed in Underhill et al. (2000, 2001); Cruciani et al. (2006). Cruciani et al. found no exemplars in their 2007 study.::*E1b1b1a1b (E-V32). This is a sub-clade of V12. Cruciani et al. (2007) suggest it originated in North Africa. It is prevalent in the Horn of Africa among Somalis, Ethiopians, and Eritreans. Before the discovery of V32, Cruciani et al. (2004) referred to the same lineages as the "gamma cluster", which is 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". 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 are also positive for V36. So V13 is currently considered "phylogenetically equivalent" to V13.)

E1b1b is common throughout Europe. [Firasat et al. (2007), [http://www.nature.com/ejhg/journal/v15/n1/full/5201726a.html 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"] Its E1b1b1a2 (E-V13) sub-clade is the most prevalent clade of E-M78 among Europeans, especially in the Balkans where high concentrations are reported amongst Albanians, Macedonians, Greeks, Bulgarians, Romanians, and Serbs.Rosser et al. (2000), [http://www.ajhg.org/AJHG/abstract/S0002-9297(07)63221-2 Y-Chromosomal Diversity in Europe Is Clinal and Influenced Primarily by Geography, Rather than by Language] , "American Journal of Human Genetics", 67: 1526-1543.] Peričic et. al. (2005), [http://mbe.oxfordjournals.org/cgi/content/full/22/10/1964 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] [King et al. (2008) [http://dirkschweitzer.net/E3b-papers/KingAHG-08-72-205.pdf Differential Y-chromosome Anatolian Influences on the Greek and Cretan Neolithic] Annals of Human Genetics 72,205–214] Particularly high frequencies have been observed in Kosovar Albanians (45.6%) and Peloponnesian Greeks (47%).

The 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.

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 - 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.

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 Cruciani et al. (2007) 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. He suggests that this might therefore 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. It then spread to Europe vie trade routes along the rivers of SouthEastern Europe.

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. See especially Cruciani et al. (2007).

: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)

This sub-clade of E-M78 is prevalent 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

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.Arredi et al. (2004), [http://www.pubmedcentral.nih.gov/articlerender.fcgi?artid=1216069 A Predominantly Neolithic Origin for Y-Chromosomal DNA Variation in North Africa] , "American Journal of Human Genetics", 75: 338–345.] 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, probably due to ancient migrations during the Islamic, Roman, and Carthaginian empires, as well as the influence of Sephardic Jews. [Gonçalves et al. (2005), [http://wysinger.homestead.com/portugal.pdf "Y-chromosome Lineages from Portugal, Madeira and Açores Record Elements of Sephardim and Berber Ancestry"] , Annals of Human Genetics (2005) 69,443–454] This sub-clade of E1b1b has also been observed in 40% of Europeans in the Pasiegos from Cantabria.

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-M165).

E1b1b1c (E-M123); formerly E3b1c, E3b3

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.

E1b1b1c (E-M123) is quite common among both Ashkenazi and Sephardic Jews, accounting for over 10% of all male lines. [Semino et al. 2004 [http://www.pubmedcentral.nih.gov/articlerender.fcgi?artid=1181965&rendertype=table&id=TB1 Table 1] ] 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 backmigrated 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 andAshkenazim possess this sub-clade in similar high frequency supports an Israelite/Middle Eastern origin.

:Sub Clades of E1b1b1c (E-M123):::*E1b1b1c1a. Defined by SNP mutations M84 and M136::*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.

E-M293

This sub-clade of E-M35 is associated by Henn et al. (2008) with the spread of pastoralism from Eastern Africa into Southern Africa.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.] 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 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 is the new name for 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 up-to-date phylogenetic clade name is in doubt for the time being.

ee also

*Human Y-chromosome DNA haplogroup
*Haplogroup E (Y-DNA)
*Haplogroup D (Y-DNA)
*Haplogroup DE (Y-DNA)

Notes

References

*Arredi et al. (2004) [http://www.pubmedcentral.nih.gov/articlerender.fcgi?artid=1216069 A Predominantly Neolithic Origin for Y-Chromosomal DNA Variation in North Africa] , "American Journal of Human Genetics", 75: 338–345.
*Beleza et al. (2006) [http://www3.interscience.wiley.com/cgi-bin/fulltext/118548798/PDFSTART Micro-Phylogeographic and Demographic History of Portuguese Male Lineages] Annals of Human Genetics (2006) 70,181–194
*Bosch E, Calafell F, Comas D, Oefner PJ, Underhill PA, Bertranpetit J (2001) [http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pubmed&pubmedid=11254456 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
*Bosch et al. (2006) [http://www3.interscience.wiley.com/journal/118548826/abstract?CRETRY=1&SRETRY=0 Paternal and maternal lineages in the Balkans show a homogeneous landscape over linguistic barriers, except for the isolated Aromuns] Annals of Human Genetics (2006) 70,459–487
*Ellen Coffman-Levy (2005) [http://www.jogg.info/11/coffman.pdf A MOSAIC OF PEOPLE: THE JEWISH STORY AND A REASSESSMENT OF THE DNA EVIDENCE] Journal of Genetic Genealogy 1:12-33.
*Cruciani et al. (2002) [http://www.sciencedirect.com/science?_ob=MImg&_imagekey=B8JDD-4RH3CKT-C-J&_cdi=43612&_user=10&_coverDate=05%2F31%2F2002&_sk=%23TOC%2343612%232002%23999299994%23677124%23FLA%23display%23Volume_70,_Issue_5,_Pages_i-ii,_1077-1388_(May_2002)%23tagged%23Volume%23first%3D70%23Issue%23first%3D5%23date%23(May_2002)%23&view=c&_gw=y&wchp=dGLbVzz-zSkzS&md5=49fa407673a5d86db8983413c144248a&ie=/sdarticle.pdf A Back Migration from Asia to Sub-Saharan Africa Is Supported by High-Resolution Analysis of Human Y-Chromosome Haplotypes] , "American Journal of Human Genetics" 70:1197–1214, 2002
*Cruciani et al. (2004) [http://www.familytreedna.com/pdf/hape3b.pdf 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.
*Cruciani et al. (2006) [http://www3.interscience.wiley.com/homepages/38515/pdf/916.pdf 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".
*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.
*Hassan et al. (2008) [http://dirkschweitzer.net/E3b-papers/Hassan-Sudan-2008-AJPA.pdf Y-Chromosome Variation Among Sudanese: Restricted Gene Flow, Concordance With Language, Geography, and History] AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY
*Henn et al. (2008) Y-chromosomal evidence of a pastoralist migration through Tanzania to southern Africa. See [http://dienekes.blogspot.com/2008/08/new-y-chromosome-haplogroup-e-m293.html comment on Dienekes blog] , [http://spittoon.23andme.com/2008/08/04/the-origins-of-pastoralism-in-africa-what-do-the-genes-say/ comment on the Spitoon blog] and [http://www.eurekalert.org/pub_releases/2008-08/sumc-ssu073108.php public release] .
*Jobling, M.A. and Tyler-Smith, C. 2000. New uses for new haplotypes the human Y chromosome, disease and selection. Trends Genet. 16: 356-362
*Karafet et al, (2008) New Binary Polymorphisms Reshape and Increase Resolution of the Human Y-Chromosomal Haplogroup Tree. [http://www.genome.org/cgi/content/abstract/gr.7172008v1 Abstract] . Genome Research, published online April 2, 2008. [http://www.genome.org/cgi/data/gr.7172008/DC1/1 Supplementary Material.]
*King et al. (2008) [http://dirkschweitzer.net/E3b-papers/KingAHG-08-72-205.pdf Differential Y-chromosome Anatolian Influences on the Greek and Cretan Neolithic] Annals of Human Genetics 72,205–214
*J. R. Luis "et al.": [http://hpgl.stanford.edu/publications/AJHG_2004_v74_p000-0130.pdf The Levant versus the Horn of Africa: Evidence for Bidirectional Corridors of Human Migrations] ( [http://hpgl.stanford.edu/publications/AJHG_2004_v74_errata.pdf Errata] ), "American Journal of Human Genetics", 74: 532-544.
*Peričic et. al. (2005) [http://mbe.oxfordjournals.org/cgi/content/full/22/10/1964 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
*Rosser et al. (2000) [http://www.ajhg.org/AJHG/abstract/S0002-9297(07)63221-2 Y-Chromosomal Diversity in Europe Is Clinal and Influenced Primarily by Geography, Rather than by Language] , "American Journal of Human Genetics", 67: 1526-1543.
*Semino et al. (2000) [http://hpgl.stanford.edu/publications/Science_2000_v290_p1155.pdf The Genetic Legacy of Paleolithic Homo sapiens sapiens in Extant Europeans: A Y Chromosome Perspective]
*Semino O, et al. (2002) Ethiopians and Khoisan share the deepest clades of the human Y-chromosome phylogeny. Am J Hum Genet 70:265–268
*Semino et al. (2004) [http://www.pubmedcentral.nih.gov/articlerender.fcgi?artid=1181965 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.
*Underhill PA, et al. (2000) [http://www.nature.com/ng/journal/v26/n3/full/ng1100_358.html Y chromosome sequence variation and the history of human populations] . Nat Genet 26:358–361.
*Underhill PA, et al. (2001) [http://hpgl.stanford.edu/publications/AHG_2001_v65_p43.pdf The phylogeography of Y chromosome binary haplotypes and the origins of modern human populations. Ann Hum Genet. 65:43–62.]
*Weale et al. (2003), [http://www.genetics.org/cgi/reprint/165/1/229 Rare Deep-Rooting Y Chromosome Lineages in Humans: Lessons for Phylogeography] , Genetics 165: 229–234
*Y Chromosome Consortium "YCC" (2002) [http://www.genome.org/cgi/content/abstract/12/2/339 A Nomenclature System for the Tree of Human Y-Chromosomal Binary Haplogroups] Genome Research, Vol. 12, Issue 2, 339-348, February 2002

External links

* [http://www.familytreedna.com/public/e3b E3b Y-DNA Project at FTDNA]
* [http://www.haplozone.net/e3b/project E3b Y-DNA Project (all labs site)]
* [http://community.haplozone.net/index.php E3b Public Forum]
* [http://www.familytreedna.com/public/JewishE3bProject Jewish E3b Project at FTDNA]
* [http://mbe.oxfordjournals.org/content/vol22/issue10/images/large/molbiolevolmsi185f04_ht.jpeg Map of E3b1 distribution in Europe]


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