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{{Short description|Human Y-chromosome DNA haplogroup}}
{{redirect|E3b|the Pennsylvania Railroad locomotive|PRR E3b}}
{{Infobox haplogroup {{Infobox haplogroup
| name = E1b1b or E-M215 |name={{Hlist|E-M215|E1b1b}}
| origin-place = ]<ref name=Cruciani2004/> |origin-place=]{{sfnp|Cruciani|La_Fratta|Santolamazza|Sellitto|2004}}{{sfn|Trombetta|2015}}
|origin-date=47,500—22,400 BP{{sfn|Trombetta|2015}}<ref name="ReferenceC">{{cite journal | vauthors = Haber M, Jones AL, Connel BA, Asan, Arciero E, Huanming Y, Thomas MG, Xue Y, Tyler-Smith C | title = A Rare Deep-Rooting D0 African Y-chromosomal Haplogroup and its Implications for the Expansion of Modern Humans Out of Africa | journal = Genetics | pages = 1421–1428 | date = June 2019 | pmid = 31196864 | doi = 10.1534/genetics.119.302368 | doi-access = free | pmc = 6707464 | volume = 212 | issue = 4 }}</ref><ref name = Cruciani2007/>
| origin-date = approx 26,000 years BP
|TMRCA=34,800 BP<ref name="YFull">{{Cite web |url=https://www.yfull.com/tree/E-M215/ |title=E-M215 YTree}}</ref>
| ancestor = ]
|ancestor=]
| descendants = E1b1b1 or E-M35
|descendants={{Hlist|]|E-M281}}
| mutations = M215, and most often also M35
|mutations=M215
}}
|map=E1b1b.png|caption=Geographic distribution of the haplogroup E1b1b}}


'''E-M215''' or '''E1b1b''', formerly known as '''E3b''', is a major ]. E-M215 has two basal branches, ] and E-M281. E-M35 is primarily distributed in ] and the ], and occurs at moderate frequencies in the ], ], and ]. E-M281 occurs at a low frequency in ].
In ], ] '''E1b1b (E-M215)''' is a ] ], a sub-group of ], which is defined by the ] (SNP) ] M215.<ref name=isogg>{{Harvcoltxt|ISOGG|2008}}</ref><ref name=Karafet>{{Harvcoltxt|Karafet et al.|2008}}</ref><ref name=YCC>{{Harvcoltxt|Y Chromosome Consortium "YCC"|2002}}</ref> 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. E1b1b probably first came into existence somewhere in the area of the ], most probably in or near the ], and then expanded both into the ] and westwards, north and south of the ].


==Origins==
In nearly all discussion, E1b1b is equivalent to its very dominant sub-], E1b1b1 (E-M35), which contains nearly all of E1b1b. {{Harvcoltxt|Cruciani et al.|2004}} 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.
The origins of E-M215 were dated by Cruciani in 2007 to about 22,400 years ago in ].<ref name = Cruciani2007>{{Harvcoltxt | Cruciani|La Fratta | Trombetta | Santolamazza | 2007}}</ref><ref group = "Note">{{Harvcoltxt | Cruciani |La Fratta|Santolamazza| Sellitto | 2004 | ps =: "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* ]." As mentioned above, "E3b" is the old name for E-M215.}} {{Harvcoltxt |Semino |Magri |Benuzzi |Lin |2004 | ps =: "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*."}} For E-M215 {{Harvcoltxt|Cruciani|La Fratta | Trombetta | Santolamazza | 2007}} reduced their estimate to 22,400 from 25,600 in {{Harvcoltxt|Cruciani|La Fratta| Santolamazza | Sellitto | 2004}}, re-calibrating the same data.</ref>


]
As discussed in more detail below, E1b1b is presently found in various forms in the ]<ref name=Semino2004>{{Harvcoltxt|Semino et al.|2004}}</ref>, ]<ref name=Semino2004/>, parts of ] and ]<ref name=Cruciani2004>{{Harvcoltxt|Cruciani et al.|2004}}</ref>, ]<ref name=Cruciani2004/>, and ]<ref name=Firasat2006>{{Harvcoltxt|Firasat et al.|2006}}</ref> (especially the ] and the ]<ref name=Cruciani2004/><ref name=Semino2004/><ref name=Rosser2000>{{Harvcoltxt|Rosser et al.|2000}}</ref>).


==Ancient DNA==
E1b1b and E1b1b1 are quite common amongst populations speaking an ]. According to at least some theories on the origins of this linguistic group, such as those advanced by ], 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 ({{Harvcoltxt|Ehret et al.|2004}}<ref>Also see {{Harvcoltxt|Keita and Boyce|2005}}.</ref>). However, this hypothesis is disputed by those who suggest a Levantine origin for Afro-Asiatic.<ref>The only major linguist to argue for a Levantine origin of Afro-Asiatic is ], but some archaeologists who argue for ] "]" such as ] argue that there is no significant archaeological evidence to support assuming an African origin, 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 ({{Harvcoltxt|Ehret et al.|2004}}).</ref>
According to Lazaridis et al. (2016), ] skeletal remains from the ancient ] predominantly carried the Y-DNA haplogroup E1b1b. Of the five Natufian specimens analyzed for paternal lineages, three belonged to the E1b1b1b2(xE1b1b1b2a, E1b1b1b2b), E1b1(xE1b1a1, E1b1b1b1) and E1b1b1b2(xE1b1b1b2a, E1b1b1b2b) subclades (60%). Haplogroup E1b1b was also found at moderate frequencies among fossils from the ensuing ] culture, with the E1b1b1 and E1b1b1b2(xE1b1b1b2a, E1b1b1b2b) subclades observed in two of seven PPNB specimens (~29%). The scientists suggest that the Levantine early farmers may have spread southward into East Africa, bringing along Western Eurasian and ] ancestral components separate from that which would arrive later in North Africa.


Additionally, haplogroup E1b1b1 has been found in an ]ian mummy excavated at the ] archaeological site in Middle Egypt, which dates from a period between the late ] and the ].<ref>{{cite journal|last1=Schuenemann, Verena J.|display-authors=etal|title=Ancient Egyptian mummy genomes suggest an increase of Sub-Saharan African ancestry in post-Roman periods|journal=Nature Communications|date=2017|volume=8|page=15694|pmid=28556824|doi=10.1038/ncomms15694|pmc=5459999|bibcode=2017NatCo...815694S}}</ref> Fossils at the ] site of ] in ], which have been dated to around 5,000 BCE, also carried haplotypes related to the E1b1b1b1a (E-M81) subclade. These ancient individuals bore an autochthonous Maghrebi genomic component that peaks among modern ], indicating that they were ancestral to populations in the area.<ref>{{cite bioRxiv|last1=Fregel|display-authors=etal|year=2018|title=Ancient genomes from North Africa evidence prehistoric migrations to the Maghreb from both the Levant and Europe|biorxiv=10.1101/191569}}</ref> The E1b1b haplogroup has likewise been observed in ancient ] fossils excavated in ] and ] on the ], which have been radiocarbon-dated to between the 7th and 11th centuries CE. The clade-bearing individuals that were analysed for paternal DNA were inhumed at the Tenerife site, with all of these specimens found to belong to the E1b1b1b1a1 or E-M183 subclade (3/3; 100%).<ref>{{cite journal|last1=Rodrı́guez-Varela|display-authors=etal|title=Genomic Analyses of Pre-European Conquest Human Remains from the Canary Islands Reveal Close Affinity to Modern North Africans|journal=Current Biology|date=2017|volume=27|issue=1–7|pages=3396–3402.e5|doi=10.1016/j.cub.2017.09.059|pmid=29107554|doi-access=free|bibcode=2017CBio...27E3396R |hdl=2164/13526|hdl-access=free}}</ref>
Amongst populations with an Afro-Asiatic speaking history, a significant proportion of all ] male lines are made up of a wide variety of E1b1b1 (E-M35) sub-clades. {{Harvcoltxt|Behar et al.|2003}} found only ] lineages in higher numbers amongst ].<ref>{{Harvcoltxt|Behar et al.|2003}}. See Table 2.</ref> The authors also found E1b1b to be, along with haplogroup J, one of the major founding lineages among ] Jews.<ref>{{Harvcoltxt|Behar et al.|2003}}. "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."</ref> E1b1b is observed in over 22.8% of Ashkenazis<ref>{{Harvcoltxt|Nebel et al.|2001}}</ref> and 30% of ].<ref name=Semino2004/> 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.<ref name="Coffman2005">{{Harvcoltxt|Coffman-Levy|2005}}</ref>


Loosdrecht et al. (2018) analysed genome-wide data from seven ancient ] individuals from the Grotte des Pigeons near ] in eastern Morocco. The fossils were directly dated to between 15,100 and 13,900 calibrated years before present. The scientists found that five male specimens with sufficient nuclear DNA preservation belonged to the E1b1b1a1 (M78) subclade, with one skeleton bearing the E1b1b1a1b1 parent lineage to E-V13, another male specimen belonged to E1b1b (M215*).<ref name="Taforalt">{{cite journal | doi=10.1126/science.aar8380 | title=Pleistocene North African genomes link Near Eastern and sub-Saharan African human populations | journal=Science | date=4 May 2018 | volume=360 | issue=6388 | pages=548–552 | last1=Van De Loosdrecht | first1=Marieke | last2=Bouzouggar | first2=Abdeljalil | last3=Humphrey | first3=Louise | last4=Posth | first4=Cosimo | last5=Barton | first5=Nick | last6=Aximu-Petri | first6=Ayinuer | last7=Nickel | first7=Birgit | last8=Nagel | first8=Sarah | last9=Talbi | first9=El Hassan | last10=El Hajraoui | first10=Mohammed Abdeljalil | last11=Amzazi | first11=Saaïd | last12=Hublin | first12=Jean-Jacques | last13=Pääbo | first13=Svante | last14=Schiffels | first14=Stephan | last15=Meyer | first15=Matthias | last16=Haak | first16=Wolfgang | last17=Jeong | first17=Choongwon | last18=Krause | first18=Johannes | pmid=29545507 | bibcode=2018Sci...360..548V | s2cid=206666517 | doi-access=free }}</ref>
==Other Names, and history of the classification==


==Distribution==
The current ] terminology "E1b1b" and "E1b1b1" was proposed by {{Harvcoltxt|Karafet et al.|2008}}. This paper was intended to be an update of the "Y Chromosome Consortium"({{Harvcoltxt|YCC|2002}}). The YCC first formalized the original phylogenetic nomenclature - "E3b" (E-M215) and "E3b1" (E-M35) - which is still found widely especially in older literature.


In Africa, E-M215 is distributed in highest frequencies in the ] and ], specifically in the countries ] and ], whence it has in recent millennia expanded as far south as ], and northwards into ] and ] (especially the ] and the ]).<ref name=Cruciani2004>{{Harvcoltxt|Cruciani|La Fratta|Santolamazza|Sellitto|2004}}</ref><ref name=Semino2004>{{Harvcoltxt|Semino|Magri|Benuzzi|Lin|2004}}</ref><ref name=Rosser2000>{{Harvcoltxt|Rosser|Zerjal|Hurles|Adojaan|2000}}</ref><ref name=Firasat2006>{{Harvcoltxt|Firasat|Khaliq|Mohyuddin|Papaioannou|2006}}</ref> E-M281 has been found in ].<ref name="Semino2004" />
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.


Almost all E-M215 men are also in E-M35. In 2004, M215 was found to be older than M35 when individuals were found who have the M215 mutation, but do not have M35 mutation.<ref name=Cruciani2004/>
Prior to {{Harvcoltxt|Cruciani et al.|2004}}, both E1b1b and E1b1b1, not yet distinguished at that time, had been referred to as '''Hg21''' ('''Haplogroup 21''') within {{Harvcoltxt|Zerjal et al.|1999}}'s nomenclature, or as '''Eu4''' according to {{Harvcoltxt|Semino et al.|2000}}'s classification.
In 2013, Di Cristofaro et al. (2013) found one individual in Khorasan, North-East Iran to be positive for M215 but negative for M35.<ref name="Di Cristofaro">{{cite journal |last1=Di Cristofaro |first1=Julie |display-authors=etal |title=Afghan Hindu Kush: Where Eurasian Sub-Continent Gene Flows Converge |journal=PLOS ONE |date=October 18, 2013 |volume=8 |issue=10 |page=e76748 |doi=10.1371/journal.pone.0076748 |pmid=24204668 |issn=1932-6203 |pmc=3799995 |oclc=5534533323 |bibcode=2013PLoSO...876748D |s2cid=16455960 |doi-access=free }}</ref>


E-M215 and E-M35 are quite common among ]. The linguistic group and carriers of E-M35 lineage have a high probability to have arisen and dispersed together from the ].{{sfnmp|1a1=Ehret|1a2=Keita|1a3=Newman|1y=2004|2a1=Keita|2a2=Boyce|2y=2005|3a1=Keita|3y=2008}} Amongst populations with an Afro-Asiatic speaking history, a significant proportion of ] male lineages are E-M35.<ref name="pmid13680527">{{Harvcoltxt|Behar|Thomas|Skorecki|Hammer|2003}}</ref> Haplogroup E-M35, which accounts for approximately 18%<ref name="Semino2004" /> to 20%<ref name="Behar2004">{{Harvcoltxt|Behar|Garrigan|Kaplan|Mobasher|2004}}</ref><ref name="Shen2004">{{Harvcoltxt|Shen|Lavi|Kivisild|Chou|2004}}</ref> of ] and 8.6%<ref name="Adams2008">{{Harvcoltxt|Adams|Bosch|Balaresque|Ballereau|2008}}</ref> to 30%<ref name="Semino2004" /> of ] Y-chromosomes, appears to be one of the major founding lineages of the Jewish population.<ref name=Nebel2001>{{Harvcoltxt|Nebel|Filon|Brinkmann|Majumder|2001}}</ref><ref group="Note">"Paragroup E-M35 * 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." {{Harvcoltxt|Behar|Garrigan|Kaplan|Mobasher|2004}}</ref>
They were also within {{Harvcoltxt|Underhill et al.|2001}}'s "Group III", and in terms of "p49/TaqI" tests, E-M35 came within Haplotype V. (It should be generally be kept in mind that some older haplogroup testing methods could not distinguish between related clades in a way perfectly consistent with more recent findings.)


===E-M215 association with endurance===
Other older names are referred to in the YCC 2002 report in the referenced articles, but are less common in the literature.


Moran et al. (2004) observed that among Y-DNA (paternal) clades borne by elite endurance athletes in Ethiopia, the haplogroup E3b1 was negatively correlated with elite athletic endurance performance,<ref name="Moran-a"/> whereas the haplogroups ], ], ],<ref name="Moran-a">{{cite journal|last1=Moran, Colin N.|display-authors=etal|title=Y chromosome haplogroups of elite Ethiopian endurance runners|journal=Human Genetics|date=2004|volume=115|issue=6|pages=492–7|url=https://www.researchgate.net/publication/8214295|access-date=6 February 2017|doi=10.1007/s00439-004-1202-y|pmid=15503146|s2cid=13960753}}</ref> and ] were significantly more frequent among the elite endurance athletes.<ref name="Moran-a"/>
==Origins==
]
Concerning the origins of the E1b1b lineage, {{Harvcoltxt|Bosch et al.|2001}}, {{Harvcoltxt|Semino et al.|2004}}<ref>"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*."</ref>, {{Harvcoltxt|Cruciani et al.|2004}}<ref>"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* ]." As mentioned above, "E3b" is the old name for E1b1b (E-M215).</ref>, (2006)<ref name=Cruciani2006>{{Harvcoltxt|Cruciani et al.|2006}}</ref>, and (2007)<ref name=Cruciani2007>{{Harvcoltxt|Cruciani et al.|2007}}</ref>, 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.<ref name=Karafet/> There are different techniques available for such estimates, and a considerable range of possibilities, but the most recent estimates of {{Harvcoltxt|Cruciani et al.|2007}} are around 22,400 years ago for E-M215<ref>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.</ref> or E-M35.<ref>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.</ref>


==Subclades==
All major sub-branches of E1b1b1 are thought to have originated in the same general area as the parent clade: in ], the ], or the ]. {{Harvcoltxt|Underhill|2002}} believes that the structure and regional pattern of E-M35 sub-clades potentially give "reagents with which to infer specific episodes of population histories associated with the ] agricultural expansion". Concerning European E-M35 within this scheme, {{Harvcoltxt|Underhill and Kivisild|2007}} have remarked that E1b1b seems to represent a late-] migration from Africa to Europe over the ] in ].<ref>"Y chromosome data show a signal for a separate late-Pleistocene migration from Africa to Europe via Sinai as evidenced through the distribution of haplogroup E3b lineages, which is not manifested in mtDNA haplogroup distributions."{{Harvcoltxt|Underhill and Kivisild|2007|p=547}}</ref>


===E-M35===
Citing {{Harvcoltxt|Cruciani et al.|2004}}, {{Harvcoltxt|Coffman-Levy|2005}} 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.<ref>{{Harvcoltxt|Coffman-Levy|2005}}: "Unfortunately, misinformation about these haplogroups continues to pervade the public and media. Haplogroup E3b is often incorrectly described as “African,” leaving a misimpression regarding the origin and complex history of this haplogroup. Haplogroup J2, as previously discussed, is often incorrectly equated with J1 and described as “Jewish” or “Semitic,” despite the fact that it is present in a variety of non-Jewish Mediterranean and Northern European populations. And haplogroup G is rarely discussed in depth; its origin and distribution remain poorly understood."</ref>
{{Main|Haplogroup E-M35}}


Haplogroup E-M35 is a subclade of E-M215.
==Subclades of E1b1b1 (E-M35)==


===E-M281===
As mentioned above, nearly all E1b1b lineages are within E1b1b1 (defined by M35). {{Harvcoltxt|Cruciani et al.|2004}} found 2 out of 34 Ethiopian ] tested, to be M215 positive but M35 negative, and therefore in the paragroup "E-M215*". More recently, {{Harvcoltxt|Cadenas et al.|2007}} found one more E-M215* individual in ], just across the ] from the Amhara, out of 62 people tested there.
{{Main|Haplogroup E-M281}}


Haplogroup E-M281 is a subclade of E-M215.
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.


==Phylogenetics==
The two most written-about sub-clades of E1b1b1 are ] (defined by M78) and ] (defined by M81), both are associated with the ]. 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 ], the ] and generally outside of Africa. It is also common in ] and the ]. E1b1b1b is found mainly in the ], but it is also found mainly in very low frequencies in neighbouring areas of ], ] and the ].


===Phylogenetic history===
A third very significant sub-clade of E1b1b1 is E1b1b1c (defined by M123)<ref name=isogg>{{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}}</ref>. It is found both in and out of Africa, but probably had a Near Eastern origin. (See ].)
{{main|Conversion table for Y chromosome haplogroups}}


Prior to 2002, there were in academic literature at least seven naming systems for the Y-Chromosome phylogenetic tree. This led to considerable confusion. In 2002, the major research groups came together and formed the Y-Chromosome Consortium (YCC). They published a joint paper that created a single new tree that all agreed to use. Later, a group of citizen scientists with an interest in population genetics and genetic genealogy formed a working group to create an amateur tree aiming at being above all timely. The table below brings together all of these works at the point of the landmark 2002 YCC Tree. This allows a researcher reviewing older published literature to quickly move between nomenclatures.
The fourth major sub-clade of E1b1b1 to be announced (Henn et al. 2008) is defined by M293, an SNP or ] 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.


{| class="wikitable"
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.
! align="center" style="background:#c63;"|'''YCC 2002/2008 (Shorthand)'''

! align="center" style="background:#f96;"|'''(α)'''
===E1b1b1a (E-M78); formerly E3b1a===
! align="center" style="background:#f96;"|'''(β)'''
]
! align="center" style="background:#f96;"|'''(γ)'''
E1b1b1a (E-M78) is a commonly occurring subclade, widely distributed in ] , the ], ] "up to Southern Asia"<ref>{{Harvcoltxt|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".</ref>, and all of ].<ref>{{Harvcoltxt|Cruciani et al.|2006}}: "The human Y chromosome haplogroup E-M78 (E3b1a) occurs commonly and is distributed in northern and eastern Africa, western Asia, and all of Europe."</ref> The European distribution has a frequency peak centered in parts of the ] (up to almost 50%<ref name=Semino2004/><ref name="Pericic2005">{{Harvcoltxt|Peričic et al.|2005}}</ref>) and Italy (see below) and declining frequencies evident toward western, central, and northeastern Europe.
| align="center" style="background:#f96;"|'''(δ)'''

| align="center" style="background:#f96;"|'''(ε)'''
Based on genetic ] variance data, Cruciani et al. (2007) suggests that this subclade originated in "Northeastern Africa", which in the study refers specifically to ] and ].<ref>{{Harvcoltxt|Cruciani et al.|2007}} use the term "Northeastern Africa" to refer to Egypt and Libya, as shown in Table 1 of the study.</ref> The E-M78 mutation is thought to have occurred about 18,600 years ago (17,300 - 20,000 years ago), with some possibility that it may have been more recent.<ref>{{Harvcoltxt|Cruciani et al.|2007}} use two calculation methods for estimating the age of E-M78 which give very different results. For the main 18,600 years ago, the ] is used, while for a second "]", used as a check, gives 13.7kya with a ] of 2.3kya, but the difference between the two methods is only large for the age estimation of E-M78, not its sub-clades. The authors state the the big difference is "attributable to the relevant departure from a star-like structure because of repeated ]s"</ref> {{Harvcoltxt|Battaglia et al.|2008}} describe Egypt as "a hub for the distribution of the various geographically localized M78-related sub-clades" and, based on archaeological data, they propose that the point of origin of E-M78 (as opposed to later dispersals from Egypt) may have been in a ] which "existed on the border of present-day ] and Egypt, near ], 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 ] foragers northwards in Africa, the ] and ultimately onwards to ] and Europe, where they each eventually differentiated into their regionally distinctive branches". Towards the south, {{Harvcoltxt|Hassan et al.|2008}} also explain evidence that some subclades of E-M78, specifically E-V12 and E-22, "might have been brought to Sudan from North Africa after the progressive desertification of the Sahara around 6,000-8,000 years ago".
| align="center" style="background:#f96;"|'''(ζ)'''
], and ].]]
| align="center" style="background:#f96"|'''(η)'''
Prior to {{Harvcoltxt|Cruciani et al.|2007}}, {{Harvcoltxt|Semino et al.|2004}} had proposed the Horn of Africa as a possible place of origin of E-M78. This was because of the high frequency and diversity of E-M78 lineages in the region. For example, {{Harvcoltxt|Sanchez et al.|2005}} found that 77.6% of 201 male ] tested in ] were members of this clade. However, {{Harvcoltxt|Cruciani et al.|2007}} were able to study more data, including populations from North Africa who were not represented in the {{Harvcoltxt|Semino et al.|2004}} study, and found evidence that the E-M78 lineages in the Horn of Africa were relatively recent branches (see E1b1b1a1b (E-V32) ]). They concluded that Northeast Africa was the likely place of origin of E-M78 based on "the peripheral geographic distribution of the most derived subhaplogroups with respect to northeastern Africa, as well as the results of quantitative analysis of UEP and microsatellite diversity". E-M215, the parent clade of E-M78, originated in East Africa during the paleolithic and subsequently, E-M215 spread to Northeast Africa. According to {{Harvcoltxt|Cruciani et al.|2007}}, the presence of E-M78 in East Africa, is the result of a back migration of E-M215 chromosomes that had acquired the E-M78 mutation. {{Harvcoltxt|Cruciani et al.|2007}} also note this as evidence for "a corridor for bidirectional migrations" (conceivably the ]) between Northeast Africa (Egypt and Libya in their data) on the one hand and East Africa on the other. The authors believe there were "at least 2 episodes between 23.9–17.3 ky and 18.0–5.9 ky ago".
| align="center" style="background:#f96"|'''YCC 2002 (Longhand)'''

| align="center" style="background:#c96;"|'''YCC 2005 (Longhand)'''
{{Harvcoltxt|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 to western Asia between 20.0 and 6.8 ky ago. While there were apparently direct migrations from North Africa to ] and ] (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 ], where it apparently originated, via the ] (see ]).
| align="center" style="background:#c96;"|'''YCC 2008 (Longhand)'''

| align="center" style="background:#c96;"|'''YCC 2010r (Longhand)'''
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 ] 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 {{Harvcoltxt|Karafet et al.|2008}}, and , which is in turn the basis of the phylogeny given below...
| align="center" style="background:#ff9;"|'''ISOGG 2006'''

| align="center" style="background:#ff9;"|'''ISOGG 2007'''
=====E1b1b1a1 (E-V12)=====
| align="center" style="background:#ff9;"|'''ISOGG 2008'''
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<ref name=Cruciani2007>See Figure 1.</ref>). According to {{Harvcoltxt|Cruciani et al.|2007}}, E-V12 likely originated in ].
| align="center" style="background:#ff9;"|'''ISOGG 2009'''
| align="center" style="background:#ff9;"|'''ISOGG 2010'''
'''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 ], but also scattered widely in small amounts in both Northern Africa and Europe, but with very little sign in Western Asia, apart from Turkey<ref name=Cruciani2007/>. These E-V12* lineages were formerly included (along with many E-V22* lineages<ref>{{Harvcoltxt|Cruciani et al.|2004}}: "E-V22 and E-V12* chromosomes are intermingled and not clearly differentiated by their microsatellite haplotypes". In {{Harvcoltxt|Cruciani et al.|2007}} 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 {{Harvcoltxt|Cruciani et al.|2007}} and {{Harvcoltxt|Cruciani et al.|2004}}.)</ref>) in Cruciani et al.'s original (2004) "delta cluster", which he had defined using ] profiles. With the discovery of the defining SNP, {{Harvcoltxt|Cruciani et al.|2007}} reported that V-12* was found in its highest concentrations in Egypt, especially Southern Egypt. {{Harvcoltxt|Hassan et al.|2008}} report a significant presence of E-V12* in neighboring Sudan, including 5/33 ] and 5/39 ]. 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.<ref name="Hassan2008">{{Harvcoltxt|Hassan et al.|2008}}</ref> The E-V12* ] is also observed in Europe (e.g. amongst French ]) and Eastern ] (e.g. ] ]).<ref name=Cruciani2007/>
| align="center" style="background:#ff9;"|'''ISOGG 2011'''

| align="center" style="background:#ff9;"|'''ISOGG 2012'''
'''Sub Clades of E1b1b1a1 (E-V12):'''
======<big>E1b1b1a1a (E-M224)</big>======
E1b1b1a1a (E-M224) has been found in ] among ]i population (5%) and appears to be a minor subclade. Its discovery was announced in {{Harvcoltxt|Underhill et al.|2001}} and {{Harvcoltxt|Cruciani et al.|2004}} found 1 Yemeni exemplar. {{Harvcoltxt|Cruciani et al.|2006}} called M224 "rare and rather uninformative", and they found no exemplars in the data they collected and discussed in their 2004, 2006, and 2007 studies.

======<big>E1b1b1a1b (E-V32)</big>======
{{Harvcoltxt|Cruciani et al.|2007}} suggest that this sub-clade of E-V12 originated in ] , and then subsequently expanded further south into the ], where it is now prevalent.<ref name=Cruciani2007>{{Harvcoltxt|Cruciani et al.|2007}}: Fig. 2/C</ref> Before the discovery of V32, {{Harvcoltxt|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 ]-speaking groups: the ] from ] (71.4%), the ] from ] (32.0%), and the ] (52.2%). Outside of eastern Africa, it was found only in two subjects from Egypt (3.6%) and in one Arab from Morocco". {{Harvcoltxt|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". {{Harvcoltxt|Hassan et al.|2008}} in their study observed this to be the most common of the sub-clades of E-M78 found in ], especially among the ], ], and ]. The Beja, like Somalis and Oromos, speak an ] language and live along the "corridor" from Egypt to the Horn of Africa. {{Harvcoltxt|Hassan et al.|2008}} interpret this as reinforcing the "strong correlation between linguistic and genetic diversity" and signs of contact between the Beja and the peoples of the Horn of Africa such as the ] and Oromo. On the other hand, the Masalit and Fur live in ] and speak a ] language. The authors observed in their study that "the Masalit possesses by far the highest frequency of the E-M78 and of the E-V32 haplogroup", which they believe suggests "either a recent ] in the population or a proximity to the origin of the haplogroup."

The STR data from {{Harvcoltxt|Cruciani et al.|2007}} concerning E-V12 can be summarized as follows...
<center>
{| {{table}}
| align="center" style="background:#f0f0f0;"|'''Haplotype'''
| align="center" style="background:#f0f0f0;"|'''description'''
| align="center" style="background:#f0f0f0;"|'''YCAIIa'''
| align="center" style="background:#f0f0f0;"|'''YCAIIb'''
| align="center" style="background:#f0f0f0;"|'''DYS413a'''
| align="center" style="background:#f0f0f0;"|'''DYS413b'''
| align="center" style="background:#f0f0f0;"|'''DYS19'''
| align="center" style="background:#f0f0f0;"|'''DYS391'''
| align="center" style="background:#f0f0f0;"|'''DYS393'''
| align="center" style="background:#f0f0f0;"|'''DYS439'''
| align="center" style="background:#f0f0f0;"|'''DYS460'''
| align="center" style="background:#f0f0f0;"|'''DYS461'''
| align="center" style="background:#f0f0f0;"|'''A10'''
|- |-
| E-V12*||modal||19||22||22||22||13||10||13||11||11||9||13 | ]||21||III||3A||13||Eu3||H2||B||E*||E||E||E||E||E||E||E||E||E||E
|- |-
| ||min||18||21||20||21||11||10||12||11||8||8||11 | ]||21||III||3A||13||Eu3||H2||B||E1*||E1||E1a||E1a||E1||E1||E1a||E1a||E1a||E1a||E1a
|- |-
| ]||21||III||3A||13||Eu3||H2||B||E1a||E1a||E1a1||E1a1||E1a||E1a||E1a1||E1a1||E1a1||E1a1||E1a1
| ||max||19||22||22||23||15||12||14||13||12||10||14
|- |-
| ||number||40||40||40||40||40||40||40||40||40||40||40 | ]||21||III||3A||13||Eu3||H2||B||E2a||E2||E2||E2||E2||E2||E2||E2||E2||E2||E2
|- |-
| E-V32||modal||19||21||22||23||11||10||13||12||10||10||13 | ]||21||III||3A||13||Eu3||H2||B||E2b||E2b||E2b||E2b1||-||-||-||-||-||-||-
|- |-
| ]||25||III||4||14||Eu3||H2||B||E3*||E3||E1b||E1b1||E3||E3||E1b1||E1b1||E1b1||E1b1||E1b1
| ||min||19||19||20||21||11||9||12||11||9||10||11
|- |-
| ]||8||III||5||15||Eu2||H2||B||E3a*||E3a||E1b1||E1b1a||E3a||E3a||E1b1a||E1b1a||E1b1a||E1b1a1||E1b1a1
| ||max||20||22||22||24||11||11||13||13||12||11||14
|- |-
| ]||8||III||5||15||Eu2||H2||B||E3a1||E3a1||E1b1a1||E1b1a1||E3a1||E3a1||E1b1a1||E1b1a1||E1b1a1||E1b1a1a1a||E1b1a1a1a
| ||number||35||35||35||35||35||35||35||35||35||35||35
|- |-
| ]||8||III||5||15||Eu2||H2||B||E3a2||E3a2||E1b1a2||E1b1a2||E3a2||E3a2||E1b1a2||E1b1a2||E1ba12||removed||removed
| All E-V12||modal||19||22||22||23||11||10||13||11||11||10||13
|- |-
| ]||8||III||5||15||Eu2||H2||B||E3a3||E3a3||E1b1a3||E1b1a3||E3a3||E3a3||E1b1a3||E1b1a3||E1b1a3||E1b1a1a1c||E1b1a1a1c
| ||min||18||19||20||21||11||9||12||11||8||8||11
|- |-
| ]||8||III||5||15||Eu2||H2||B||E3a4||E3a4||E1b1a4||E1b1a4||E3a4||E3a4||E1b1a4||E1b1a4||E1b1a4||E1b1a1a1g1c||E1b1a1a1g1c
| ||max||20||22||22||24||15||12||14||13||12||11||14
|- |-
| ]||8||III||5||15||Eu2||H2||B||E3a5||E3a5||E1b1a5||E1b1a5||E3a5||E3a5||E1b1a5||E1b1a5||E1b1a5||E1b1a1a1d||E1b1a1a1d
| ||number||75||75||75||75||75||75||75||75||75||75||75
|- |-
| ]||8||III||5||15||Eu2||H2||B||E3a6||E3a6||E1b1a6||E1b1a6||E3a6||E3a6||E1b1a6||E1b1a6||E1b1a6||E1b1a1a1e||E1b1a1a1e
|
|}
</center>

=====E1b1b1a2 (E-V13)=====

The E-V13 clade is equivalent to the "alpha cluster" of E-M78 reported in {{Harvcoltxt|Cruciani et al.|2004}}, and was first defined by the SNP V13 in {{Harvcoltxt|Cruciani et al.|2006}}. Another SNP is known for this clade, V36, reported in {{Harvcoltxt|Cruciani et al.|2007}}. All known positive tests for V13 are also positive for V36. So E-V13 is currently considered "phylogenetically equivalent" to E-V36.

{{Quotation|Haplogroup E-V13 is the only lineage that reaches the highest frequencies out of Africa. In fact, it represents about 85% of the European E-M78 chromosomes 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 only in northern Africa at a low frequency (0.9%).|{{Harvcoltxt|Cruciani et al.|2007}}}}

Within Europe, E-V13 is especially common in the ] and some parts of Italy. In different studies, particularly high frequencies have been observed in ] Albanians (45.6%) ({{Harvcoltxt|Peričic et al.|2005}}), Albanian speakers of the Former Yugoslavian Republic of Macedonia (34.4%) ({{harvcoltxt|Battaglia et al.|2008}}), and ] ] (47%) <ref name=Semino2004/>. More generally, high frequencies have also been found in other areas of ], and amongst ], ], ], and ]<ref>{{Harvcoltxt|Cruciani et al.|2004}}, {{Harvcoltxt|Rosser et al.|2000}}, {{Harvcoltxt|Peričic et al.|2005}}, {{Harvcoltxt|King et al.|2008}}</ref>.

Within ], frequencies tend to be higher in ]<ref name=Cruciani2007/>, with particularly high results sometimes seen in particular areas, for example ] and ] in ]<ref>{{Harvcoltxt|Di Gaetano et al.|2008}}</ref>. High frequencies have also been observed in some northern areas, for example around ]<ref>{{Harvcoltxt|Scozzari et al.|2001}} See clade 25.1. The same data set was later used in {{Harvcoltxt|Cruciani et al.|2004}} and {{Harvcoltxt|Cruciani et al.|2007}}.</ref>, and ]<ref>{{Harvcoltxt|Pelotti et al.|2007}}</ref>, both in the northwest of Italy, as well as on the island of ]<ref>{{Harvcoltxt|Francalacci et al.|2003}}</ref>

Phylogenetic analysis strongly suggest that these lineages have spread through Europe, from the Balkans in a "rapid demographic expansion"<ref name=Cruciani2007/>. 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 ] and ] Arab lineages<ref name=Cruciani2007/>. The Druze are considered a genetically isolated community<ref>{{Harvcoltxt|Shlush et al.|2008}}</ref>, and are therefore of particular interest. Their STR DNA signature was actually originally classified in the delta cluster in {{Harvcoltxt|Cruciani et al.|2004}}. 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...
<center>
{| {{table}}
| align="center" style="background:#f0f0f0;"|'''haplotype'''
| align="center" style="background:#f0f0f0;"|'''description'''
| align="center" style="background:#f0f0f0;"|'''YCAIIa'''
| align="center" style="background:#f0f0f0;"|'''YCAIIb'''
| align="center" style="background:#f0f0f0;"|'''DYS413a'''
| align="center" style="background:#f0f0f0;"|'''DYS413b'''
| align="center" style="background:#f0f0f0;"|'''DYS19'''
| align="center" style="background:#f0f0f0;"|'''DYS391'''
| align="center" style="background:#f0f0f0;"|'''DYS393'''
| align="center" style="background:#f0f0f0;"|'''DYS439'''
| align="center" style="background:#f0f0f0;"|'''DYS460'''
| align="center" style="background:#f0f0f0;"|'''DYS461'''
| align="center" style="background:#f0f0f0;"|'''A10'''
|- |-
| All E-V13||modal||19||21||23||24||13||10||13||12||9||10||13 | ]||25||III||4||14||Eu4||H2||B||E3b*||E3b||E1b1b1||E1b1b1||E3b1||E3b1||E1b1b1||E1b1b1||E1b1b1||removed||removed
|- |-
| ]||25||III||4||14||Eu4||H2||B||E3b1*||E3b1||E1b1b1a||E1b1b1a1||E3b1a||E3b1a||E1b1b1a||E1b1b1a||E1b1b1a||E1b1b1a1||E1b1b1a1
| Druze V13||1||19||21||23||23||13||10||13||13||11||9||12
|- |-
| ]||25||III||4||14||Eu4||H2||B||E3b1a||E3b1a||E1b1b1a3a||E1b1b1a1c1||E3b1a3a||E3b1a3a||E1b1b1a3a||E1b1b1a3a||E1b1b1a3a||E1b1b1a1c1||E1b1b1a1c1
| Druze V13||2||19||21||23||23||13||10||13||13||11||9||13
|- |-
| ]||25||III||4||14||Eu4||H2||B||E3b2*||E3b2||E1b1b1b||E1b1b1b1||E3b1b||E3b1b||E1b1b1b||E1b1b1b||E1b1b1b||E1b1b1b1||E1b1b1b1a
| All E-V22||modal||19||22||22||23||14||10||13||12||11||10||12
|- |-
| ]||25||III||4||14||Eu4||H2||B||E3b2a||E3b2a||E1b1b1b1||E1b1b1b1a||E3b1b1||E3b1b1||E1b1b1b1||E1b1b1b1||E1b1b1b1||E1b1b1b1a||E1b1b1b1a1
| All E-V12*||modal||19||22||22||22||13||10||13||11||11||9||13
|- |-
| ]||25||III||4||14||Eu4||H2||B||E3b2b||E3b2b||E1b1b1b2||E1b1b1b1b1||E3b1b2||E3b1b2||E1b1b1b2a||E1b1b1b2a||E1b1b1b2a||E1b1b1b2a||E1b1b1b1a2a
|
|}
</center>
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.<ref name=Cruciani2007/>

======<big>E-V13 and Ancient Migrations</big>======
The apparent movement of E-M78 lineages from the Near East to Europe, and their subsequent rapid expansion, make its E-V13 sub-clade a particularly interesting subject for speculation about ancient human migrations.
]

======<big>Early Migration from the Middle East to Europe</big>======
The ] is frequently also discussed in connection to V13, as a haplogroup with a seemingly very similar distribution and pre-history<ref>See especially {{Harvcoltxt|Cruciani et al.|2007}}</ref>.

{{Harvcoltxt|Cruciani et al.|2007}} says there were at least four major demographic events which have been envisioned for this geographic area:
*The "post-] expansion (about 20 kya)"
*The "]-] reexpansion (about 12 kya)"
*The "population growth associated with the introduction of ] practices (about 8 kya)"
*The "development of ] technology (about 5kya)"

The distribution and diversity of V13 are generally thought to be suggestive that it was brought to the Balkans along with early farming technologies, during the ] expansion<ref name=Semino2004/>. However, {{Harvcoltxt|Battaglia et al.|2008}} propose that the E-M78* lineage ancestral to all modern E-V13 men moved rapidly out of a Southern Egyptian homeland, in the wetter conditions of the early ]; arrived in Europe with only ] technologies and then only subsequently integrated with Neolithic cultures which arrived later in the Balkans. They then suggest that the E-V13 sub-clade of E-M78 originated in situ in Europe, and propose that the first major dispersal of E-V13 from the Balkans may have been in the direction of the ] with the ] ] culture often referred to as ] or ].

Concerning dispersal from the Balkans {{Harvcoltxt|Cruciani et al.|2007}} suggest in contrast to Battaglia et al. that this may have been more recent than 5300 years ago. The authors suggest that this might have been associated with an ''in situ'' population increase in the Balkans associated with the Balkan ], 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". {{Harvcoltxt|Peričic et al.|2005}} specifically propose the Vardar-Morava-Danube rivers as a possible route of Neolithic dispersal into central Europe.

E-V13 is in any case generally described in ] as one of the components of the ] which shows the contribution made by the populations who dispersed Neolithic technology<ref>{{Harvcoltxt|Semino et al.|2000}}, {{Harvcoltxt|King and Underhill|2002}} {{Harvcoltxt|Underhill|2002}}</ref>. As such, it also represents a relatively recent genetic movement ] into ], and has been described "a signal for a separate late-] migration from Africa to Europe over ] ... which is not manifested in ] haplogroup distributions"<ref>{{Harvcoltxt|Underhill and Kivisild|2007}}</ref>.

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

{{Quotation|An extensive analysis of Y diversity within Greeks and three Pakistani populations – the ], ] and ] – who claim descent from Greek soldiers allowed us to compare Y lineages within these populations and re-evaluate their suggested Greek origins. This study as a whole seems to exclude a large Greek contribution to any Pakistani population, confirming previous observations. However, it provides strong evidence in support of the Greek origins for a small proportion of Pathans, as demonstrated by the clade E network and the low pairwise genetic distances between these two populations.|{{Harvcoltxt|Firasat et al.|2006}}}}

======<big>Roman soldiers in Britain</big>======
Significant frequencies of E-V13 have also been observed in towns in ], ] and ]. The old trading town of ] 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 {{Harvcoltxt|Weale et al.|2002}}. {{Harvcoltxt|Bird|2007}} attributes the overall presence of E-V13 in ], especially in areas of high frequency, to settlement during the 1st through 4th centuries CE by ] 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 '']''), which was identified by {{Harvcoltxt|Peričic et al.|2005}} as harboring the highest frequency worldwide of this sub-clade<ref>Doubts about this line of reasoning because (a) new data appearing in {{Harvcoltxt|King et al.|2008}} indicates other high concentrations in Greece and (b) the data in {{Harvcoltxt|Peričic et al.|2005}} show that the area with the highest frequency does not have the highest diversity, implying that V13 arrived there more recently than in Greece.</ref>.

However, according to data published so far<ref>Bird uses three sources: {{Harvcoltxt|Weale et al.|2002}}, {{Harvcoltxt|Capelli et al.|2003}} and {{Harvcoltxt|Sykes|2006}}. Neither Capelli nor Weale have data from the area in the English Midlands where Bird suggests that there is a lack of E1b1b. Bird mentioned that there were 193 Central English haplotypes in Sykes.</ref>, E-V13 appears to be notably absent in Central England, a fact which Bird (2007) suggests reflects a genuine population replacement of ] people with ]s:
{{Quotation|The "E3b hole" suggests that either (a) a massive displacement of the native Romano-British population by invasion or, (b) the substantial genetic replacement of Romano-British Y-DNA through an elite dominance ("apartheid") model (Thomas, 2006), has occurred in Central England. Regardless of the mechanism, the Central England region of Britain, with its lack of E3b haplotypes, is the area having the most "striking similarity in the distribution of Y-chromosomes" with Friesland {{Harvcoltxt|Thomas et al.|2006}}.|{{Harvcoltxt|Bird|2007}}}}

======<big>Phoenician Traders in the Mediterranean</big>======
{{Harvcoltxt|Zalloua et al.|2008}} have also suggested E1b1b haplotypes (amongst others) to be a sign of Phoenician influence around the Mediterranean<ref>{{Harvcoltxt|Zalloua et al.|2008}} 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.</ref>.

======<big>Sub Clades of E1b1b1a2 (E-V13)</big>======
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 {{Harvcoltxt|Karafet et al.|2008}} 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").
::*'''E1b1b1a2a.''' Defined by V27. {{Harvcoltxt|Cruciani et al.|2007}} found one case in ].
::*'''E1b1b1a2b.''' Defined by P65.

=====E1b1b1a3 (E-V22)=====
This clade comprises most of those classified in the "delta cluster" of {{Harvcoltxt|Cruciani et al.|2004}}. {{Harvcoltxt|Cruciani et al.|2006}} 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"<ref name=Cruciani2007/> in the ] and ], with higher microsatellite variance (0.35 vs. 0.46, respectively) in Egypt. In the article announcing this first information, {{Harvcoltxt|Cruciani et al.|2007}} described it as uncommon in Western Asia and they proposed Northeast Africa (Libya/Egypt) as this sub-clade's likely place of origin. {{Harvcoltxt|Hassan et al.|2008}} 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 ] (about 31%), a widely-dispersed pastoral people<ref>{{Harvcoltxt|Rosa et al.|2007}} in a study of ], 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.</ref>. E-V22 was also present in much smaller frequencies amongst the ] (2 of 15 samples, 13%) and ] (3 of 26, 8%) ] of Southern Sudan. Hassan et al. suggest that E-V22, like E-V12, might have entered Sudan from ] "after the progressive desertification of the Sahara around 6,000–8,000 years ago". They add that the ] to Sudan "is not only recent (] 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 ], who appear to have sustained considerable gene flow from ] and ] together with the ]."

Other frequencies reported by {{Harvcoltxt|Cruciani et al.|2007}} include ] (4.44% out of 90 people), ] (4.58% out of 153 people), ] (7.27%, 55 people) and ] (8%, 50 people), ] Turkish (5.71% out of 35 people), and ] (6.9% out of 29 people). {{Harvcoltxt|Cadenas et al.|2007}} found a 6.7% presence in the ].

:'''Sub Clades of E1b1b1a3 (E-V22):''' There are two recognized sub-clades, which are apparently separate, although {{Harvcoltxt|Karafet|2008}} 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".
::*'''E1b1b1a3a.''' Defined by M148. {{Harvcoltxt|Underhill et al.|2000}} found 1 example in the ]. {{Harvcoltxt|Cruciani et al.|2006}} calls M148 "rare and rather uninformative".
::*'''E1b1b1a3b.''' Defined by V19. {{Harvcoltxt|Cruciani et al.|2007}} found 2 exemplars in ].

=====E1b1b1a4 (E-V65)=====
This sub-clade, equivalent to the previously classified "beta cluster", is found in high levels in the ] regions of far northern Africa. {{Harvcoltxt|Cruciani et al.|2007}} report levels of about 20% amongst ] lineages, and about 30% amongst ]. It appears to be less common amongst ], 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...
<center>
{| {{table}}
| align="center" style="background:#f0f0f0;"|'''E-V65'''
| align="center" style="background:#f0f0f0;"|'''YCAIIa'''
| align="center" style="background:#f0f0f0;"|'''YCAIIb'''
| align="center" style="background:#f0f0f0;"|'''DYS413a'''
| align="center" style="background:#f0f0f0;"|'''DYS413b'''
| align="center" style="background:#f0f0f0;"|'''DYS19'''
| align="center" style="background:#f0f0f0;"|'''DYS391'''
| align="center" style="background:#f0f0f0;"|'''DYS393'''
| align="center" style="background:#f0f0f0;"|'''DYS439'''
| align="center" style="background:#f0f0f0;"|'''DYS460'''
| align="center" style="background:#f0f0f0;"|'''DYS461'''
| align="center" style="background:#f0f0f0;"|'''A10'''
|- |-
| ]||25||III||4||14||Eu4||H2||B||E3b3*||E3b3||E1b1b1c||E1b1b1c||E3b1c||E3b1c||E1b1b1c||E1b1b1c||E1b1b1c||E1b1b1c||E1b1b1b2a
| modal||19||21||21||23||13||10||13||10||10||11||13
|- |-
| ]||25||III||4||14||Eu4||H2||B||E3b3a*||E3b3a||E1b1b1c1||E1b1b1c1||E3b1c1||E3b1c1||E1b1b1c1||E1b1b1c1||E1b1b1c1||E1b1b1c1||E1b1b1b2a1
| min||19||20||20||22||11||10||13||10||9||9||12
|- |-
| ]||25||III||4||14||Eu4||H2||B||E3ba1||E3b3a1||E1b1b1c1a||E1b1b1c1a1||E3b1c1a||E3b1c1a||E1b1b1c1a1||E1b1b1c1a1||E1b1b1c1a1||E1b1b1c1a1||E1b1b1b2a1a1
| max||21||21||22||23||14||11||14||11||11||12||13
|- |-
| number||38||38||38||38||38||38||38||38||38||38||38
|-
|
|} |}
</center>


====Research publications====
{{Harvcoltxt|Capelli et al.|2009}} 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).


The following research teams per their publications were represented in the creation of the YCC Tree.
=====E1b1b1a5 (E-VM521)=====
{{columns-list|colwidth=22em|
This sub-clade's discovery was announced in {{Harvcoltxt|Battaglia et al.|2008}} They found 2 out of 92 Greeks to have this mutation.
* '''α''' {{harvp|Jobling|Tyler-Smith|2000}} and {{harvp|Kaladjieva|2001}}
* '''β''' {{harvp|Underhill|2000}}
* '''γ''' {{harvp|Hammer|2001}}
* '''δ''' {{harvp|Karafet|2001}}
* '''ε''' {{harvp|Semino|2000}}
* '''ζ''' {{harvp|Su|1999}}
* '''η''' {{harvp|Capelli|2001}}
}}


====Discussion====
===E1b1b1b (E-M81); formerly E3b1b, E3b2===
]
E1b1b1b (E-M81) is the most common Y chromosome haplogroup in the ], dominated by its sub-clade E-M183. It is thought to have originated in the area of North Africa 5,600 years ago.<ref name=Cruciani2004/><ref name="Arredi">{{Harvcoltxt|Arredi et al.|2004}}</ref> It is colloquially referred to as the "] marker" for its prevalence among ], ], ] and other ] groups, E-M81 is also quite common among North African ] groups. It reaches frequencies of up to 80% in the ]. This includes the ]sh for whose men {{Harvcoltxt|Bosch et al.|2001}} reports that approximately 76% are M81+.


'''E-M215''' and '''E1b1b1''' are the currently accepted names found in the proposals of the ] (YCC), for the clades defined by mutation M215 and M35 respectively, which can also be referred to as E-M215 and E-M35.<ref name=Karafet2008>{{Harvcoltxt|Karafet|Mendez|Meilerman|Underhill|2008}}</ref> The nomenclature ''E3b'' (E-M215) and ''E3b1'' (E-M35) respectively were the YCC defined names used to designate the same ]s in older literature with E-M35 branching as a separate ] of E-M215 in 2004.<ref name=Cruciani2004/> Prior to 2002 these haplogroups were not designated in a consistent way, and nor was their relationship to other related ] within ] and ]. But in non-standard or older terminologies, E-M215 is for example approximately the same as "haplotype V", still used in publications such as {{Harvcoltxt|Gérard|Berriche|Aouizérate|Diéterlen|2006}}.<ref name=YCC>{{Harvcoltxt|Y Chromosome Consortium "YCC"|2002}}</ref>
In this key area from Egypt to the ], {{Harvcoltxt|Arredi et al.|2004}} report a pattern of decreasing ] haplotype variation from East to West, accompanied by a substantial increasing frequency. On the other hand {{Harvcoltxt|Kujanova et al.|2009}} found M81 in 28.6% (10 out of 35 men) in ] in the ] in Egypt.


====Phylogenetic trees====
{{Harvcoltxt|Arredi et al.|2004}} believe the pattern of distribution and variance to be consistent with the hypothesis of a post ] "]" from the East. The ancestral lineage of E-M81 in their hypothesis could have been linked with the spread of ] food-producing technologies from the ] via the ], although ] rather than ]. E-M81 and possibly ] language may have been carried either all the way from ], or they may represent a "local contribution to the North African Neolithic transition".


Cladogram with the main subclades:
In Europe, E-M81 is found everywhere but mostly in the ], where it is more common than E-M78 unlike in the rest of Europe<ref>{{Harvcoltxt|Adams et al.|2008}}, shows an average frequency of 4% in the Iberian Peninsula with frequencies reaching 9% in ], 10% in Western] and Northwest ], .</ref> at an average frequency of 4-5.6%, with frequencies reaching 9% in ], 10% in Western ] and Northwest ] and 13 % in ]<ref>{{Harvcoltxt|Flores et al.|2005}}, {{Harvcoltxt|Beleza et al.|2006}}, {{Harvcoltxt|Adams et al.|2008}}, {{Harvcoltxt|Capelli et al.|2009}}</ref>. The highest frequency of this clade found so far in Europe has been observed at 40% the ] from ].<ref name=Cruciani2004/>


{{Clade
E-M81 is also found in ]<ref>{{Harvcoltxt|Gaetano et al.|2008}}</ref>, and in slightly lower frequencies in continental ] (especially near ])<ref>{{Harvcoltxt|Capelli et al.|2009}}</ref> and ]<ref name=Cruciani2004/>, possibly due to ancient migrations during the ], ], and ] empires, as well as the influence of ] ].<ref>{{Harvcoltxt|Gonçalves et al.|2005}}</ref>
| label1= '''E1b1b''' ('''M215''')&nbsp;
| 1={{Clade
| label1= ]
| 1={{Clade
| label1= ]
| 1={{Clade
| label1= ]
| 1={{Clade
| 1=]
| 2=E-V65
| 3=]
| 4=]}}
| 2=E-V2729}}
| label2= ]
| 2={{Clade
| 1=]
| 2=]}} }}
| 2=E1b1b2 (M281)}} }}


The following ] is based on the YCC 2008 tree and subsequent published research as summarized by ISOGG. It includes all known subclades as of June 2015 (Trombetta et al. 2015)<ref name=isogg08>{{Harvcoltxt|ISOGG|2011}}</ref><ref name=Karafet2008/><ref name="YCC"/>
As a result of its old world distribution, this sub-clade is found throughout ], for example 6.1% in ]<ref> (8 out of 132), {{Harvcoltxt|Mendizabal et al.|2008}}</ref>, 5.4% in ] (Rio de Janeiro), <ref>(6 out of 112), "The presence of chromosomes of North African origin (E3b1b-M81; Cruciani et al., 2004) can also be explained by a Portuguese-mediated influx, since this haplogroup reaches a frequency of 5.6% in ] (Beleza et al., 2006), quite similar to the frequency found in ] (5.4%) among European contributors.", {{Harvcoltxt|Silva et al.|2006}}</ref> and among ] men in the ].<ref>2.4% (7 out of 295) among Hispanic men from ] and ], {{Harvcoltxt|Paracchini et al.|2003}}</ref>


* '''E-M215''' (E1b1b)
In smaller numbers, E-M81 men can be found in areas in contact with the Maghreb, both around the Sahara, in places like ], and around the Mediterranean in places like ], ], and amongst ].
** '''E-M215*'''. Rare or non-existent.
** '''E-M35''' (E1b1b1)
*** ''']''' (E1b1b1a)
**** '''E-V2009'''. Found in individuals in Sardinia and Morocco.
**** '''E-M78''' (E1b1b1a1). North Africa, Horn of Africa, West Asia, Sicily. (Formerly "E1b1b1a".)
***** '''E-M78*'''
***** '''E-V1477'''. Found in Tunisian Jews.
***** '''E-V1083'''.
****** '''E-V1083*'''. Found only in Eritrea (1.1%) and Sardinia (0.3%).
****** '''E-V13'''
****** '''E-V22'''
***** '''E-V1129'''
****** '''E-V12'''
******* '''E-V12*'''
******* '''E-V32'''
****** '''E-V264'''
******* '''E-V259'''. Found in North Cameroon.
******* '''E-V65'''
******** E-CTS194
*** ''']''' (E1b1b1b)<ref name="ISOGG 2015">ISOGG 2015</ref>
**** '''E-V257/L19''' (L19, V257) – E1b1b1b1<ref name="ISOGG 2015"/>
***** '''E-PF2431'''
***** '''E-M81''' (M81)
****** E-PF2546
******* E-PF2546*
******* E-CTS12227
******** E-MZ11
********* E-MZ12
******* E-A929
******** E-Z5009
********* E-Z5009*
********* E-Z5010
********* E-Z5013
********** E-Z5013*
********** E-A1152
******** E-A2227
********* E-A428
********* E-MZ16
******** E-PF6794
********* E-PF6794*
********* E-PF6789
********** E-MZ21
********** E-MZ23
********** E-MZ80
******** E-A930
******** E-Z2198/E-MZ46
********* E-A601
********* E-L351
**** '''E-Z830''' (Z830) – E1b1b1b2<ref name="ISOGG 2015"/>
***** ''']''' (M123)
****** '''E-M34''' (M34)
******* '''E-M84''' (M84)
******** '''E-M136''' (M136)
******* '''E-M290''' (M290)
******* '''E-V23''' (V23)
******* '''E-L791''' (L791,L792)
***** '''E-V1515'''. E-V1515 and its subclades are mainly restricted to eastern Africa.
****** '''E-V1515*'''
****** '''E-V1486'''
******* '''E-V1486*'''
******* '''E-V2881'''
******** '''E-V2881*'''
******** '''E-V1792'''
******** '''E-V92'''
******* '''E-M293''' (M293)
******** '''E-M293*'''
******** '''E-P72''' (P72)
******** '''E-V3065*'''
****** '''E-V1700'''
******* '''E-V42''' (V42)
******* '''E-V1785'''
******** '''E-V1785*'''
******** '''E-V6''' (V6)
*** '''E-V16/E-M281''' (E1b1b2). Rare. Found in individuals in Ethiopia, Yemen and Saudi Arabia.


==See also==
There are two recognized sub-clades, although one is much more important than the other.
:'''Sub Clades of E1b1b1b (E-M81):'''
::*'''E1b1b1b1 (E-M107).''' {{Harvcoltxt|Underhill et al.|2000}} found one example in ].
::*'''E1b1b1b2 (E-M183).''' This clade is extremely dominant within E-M81. In fact, while {{Harvcoltxt|Karafet et al.|2008}} continues to describe this as a sub-clade of E-M81, and ISOGG defers to Karafet et al., all data seems to imply that it should actually be considered phylogenetically equivalent to M81. As of 24th November 2008, several SNPs are considered to define sub-clades of E-M183, although the phylogenetic structure is not yet known with confidence: M165, M243, M340, and L19<ref name=isogg/>.


{{wikiquote}}
===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<ref>As of 11 November 2008 for example, had records of four E-M123* tests, compared to 93 test results with E-M34.</ref>. However, earlier studies did not test for E-M34.


===Genetics===
'''Concerning E-M123* (tested and definitely without E-M34)''' {{harvcoltxt|Cruciani et al.|2004}} located one individual in ] after testing 3401 individuals from five continents, and Underhill et al. (2000) located one individual in ]. In a 568 person study in ], {{Harvcoltxt|Flores et al.|2004}} found 2 E-M123* individuals, both in ] out of 109 people tested there. In a 553 person study of Portugal, {{Harvcoltxt|Gonçalves et al.|2005}} also found 2 E-M123* individuals in Northern Portugal, out of 101 people, as well as 2 in ] out of 129 people tested there. {{Harvcoltxt|Flores et al.|2005}} found one individual out of 146 ]. {{Harvcoltxt|Cadenas et al.|2007}} found none amongst the significant presence of E-M34 they found in their study of the ], ] and ]. {{Harvcoltxt|Arredi et al.|2004}} found 1 Tunisian in their study of 275 men in Northern Africa.


{{columns-list|colwidth=22em|
'''Concerning E-M123 without checking for the M-34 SNP''' {{harvcoltxt|Bosch et al.|2006}} found E-M123 examples in ], the ], and ]. {{Harvcoltxt|Beleza et al.|2006}} also found examples in Portugal, and {{Harvcoltxt|Sanchez et al.|2005}} found one sample in Somalia. {{Harvcoltxt|Semino et al.|2004}} reports relatively high levels of 13% in the Albanian community of ], in ]. A notably high regional frequency for E-M123 was reported in ], where it is apparently the dominant clade of E-M35. {{Harvcoltxt|Luis et al.|2004}} found 12 men out of 121 there were E-M123 positive, while in Egypt there were 7 out of 147. But in that study the Omani E-M123 diversity implied a younger age than the E-M123 found in Egypt. {{Harvcoltxt|Shen et al.|2004}} found 4 out of 20 tested Israeli Jews of Libyan ancestry to be M123+.
*]
*]
*]
*]
*]
*]
*]
*]
*]
*]
*]
*]
*]
}}


===Y-DNA E subclades===
'''Concerning E1b1b1c1 (E-M34)''' {{Harvcoltxt|Cruciani|2004}} tested for E-M34 in Oman and found 7.7% to be E-M34+, with no E-M123*. According to {{Harvcoltxt|Cruciani|2004}}, E-M34 is found at small frequencies in North Africa and Southern Europe (6.6% in Sicily for example), and has its highest concentration in ] and the ] (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, {{harvcoltxt|Cinnioğlu et al.|2004}} found slightly more E-M34 (29) than E-M78 (26) out of 523 individuals tested (a far different E1b1b population than found in the nearby Balkans). In {{Harvcoltxt|Flores et al.|2004}} E-M34 was found in several parts of Iberia, but most strikingly about 10% in ]. {{Harvcoltxt|Gonçalves et al.|2005}} found about the same levels of E-M34 in Portugal as E-M123*, but E-M34 mainly in Central Portugal (4 people out of 102 tested there) with one more person found in the ]. Strikingly, {{Harvcoltxt|Flores et al.|2005}} found 14 out of 45 men tested in the ] area of Jordan to be M34 positive (31.1%), while in the capital ] there were only 4 out of 101. {{Harvcoltxt|Cadenas et al.|2007}} found 8.1% of 62 men tested in ] were positive for M34, compared to much lower levels in ] (1.4%) and the ] (3.1%). {{Harvcoltxt|Arredi et al.|2004}} found 2 Tunisians, 2 Algerians, and 7 Egyptians in their study of 275 men in Northern Africa.


{{columns-list|colwidth=22em|
'''E-M123 in Jews.''' Looking beyond simple regional concentrations, E1b1b1c (E-M123) is also quite common among both ] and ] ], accounting for over 10% of all male lines.<ref>{{Harvcoltxt|Semino et al.|2004}}) </ref> {{Harvcoltxt|Coffman-Levy|2005}} wrote that:
*]
* ]
* ]
* Haplogroup E-M215
* ]
* ]
* ]
* ]
* ]
* ]
* ]
* ]
* ]
* ]
* ]
* ]
* ]
* ]
* ]
}}


===Y-DNA backbone tree===
{{Quotation|...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 ({{Harvcoltxt|Semino et al.|2004}}). According to {{Harvcoltxt|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 ({{Harvcoltxt|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.}}


{{Y-DNA}}
:'''Sub Clades of E1b1b1c1 (E-M34):'''
::*'''E1b1b1c1a.''' Defined by SNP mutation M84, with M136 defining a sub-clade, E1b1b1c1a1 as of October 2008<ref name=isogg/>. The based on testing so far (in January 2009) that E-M84 is dominant in 6 out of the 8 clusters of E-M34 which that project identifies.
::*'''E1b1b1c1b.''' Defined by SNP mutation M290. {{Harvcoltxt|Shen et al.|2004}} found 1 ] exemplar.


==Notes==
===E1b1b1d (E-M281)===


{{reflist|group="Note"}}
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 ], but Cruciani et al. (2004) found no examples.


==References==
===E1b1b1e (E-V6)===
This sub-clade of E-M35 is defined by V6. {{Harvcoltxt|Cruciani et al.|2004}} (Table 1) identified a significant presence of these lineages in ], and also some in the neighboring ] population. Amongst the Ethiopian and Somali samples, the highest were 14.7% amongst the Ethiopian ], and 16.7% amongst the Ethiopian ]. One man in Kenya was also observed with the V6 mutation.


{{Reflist|30em}}
===E1b1b1f (E-P72)===


==Bibliography==
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 {{Harvcoltxt|Henn et al.|2008}}, which associated it with the spread of ] from ] into ]. So far high levels have been found in specific ethnic groups in Tanzania and Southern Africa. Highest were the ] (43%), ] (Kxoe) (31%), ] (28%), and ] (24%). Henn et al. (2008) in their study also found two Bantu-speaking Kenyan males with the M293 mutation.<ref>{{Harvcoltxt|Henn et al.|2008|page=10695}}</ref>

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

<blockquote>
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.
</blockquote>

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 and 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 ]). So the phylogenetic clade name came to be E1b1b1g in late October 2008.<ref>For example the company ] changed its webpages on or about 22 October and the relevant ] was changed on 23 October</ref>

==See also==

*]
*]
*]
*]
*]
*]
*]
*]
{{Y-DNA}}

==Notes==
{{reflist|2}}

==References==
{{Refbegin}} {{Refbegin}}
*{{Citation|author=Adams et al.|title=The Genetic Legacy of Religious Diversity and Intolerance: Paternal Lineages of Christians, Jews, and Muslims in the Iberian Peninsula| journal=The American Journal of Human Genetics | year=2008 |doi=10.1016/j.ajhg.2008.11.007 | volume=83|url=http://www.cell.com/AJHG/abstract/S0002-9297%2808%2900592-2|pages=725}} * {{Citation|last1=Adams|first1=Susan M|title=The Genetic Legacy of Religious Diversity and Intolerance: Paternal Lineages of Christians, Jews, and Muslims in the Iberian Peninsula|journal=The American Journal of Human Genetics|year=2008|doi=10.1016/j.ajhg.2008.11.007|volume=83|url= |pages=725–36|pmid=19061982|pmc=2668061|last2=Bosch|first2=Elena|last3=Balaresque|first3=Patricia L.|last4=Ballereau|first4=Stéphane J.|last5=Lee|first5=Andrew C.|last6=Arroyo|first6=Eduardo|last7=López-Parra|first7=Ana M.|last8=Aler|first8=Mercedes|last9=Grifo|first9=Marina S. Gisbert|last10=Brion|first10=Maria|last11=Carracedo|first11=Angel|last12=Lavinha|first12=João|last13=Martínez-Jarreta|first13=Begoña|last14=Quintana-Murci|first14=Lluis|last15=Picornell|first15=Antònia|last16=Ramon|first16=Misericordia|last17=Skorecki|first17=Karl|last18=Behar|first18=Doron M.|last19=Calafell|first19=Francesc|last20=Jobling|first20=Mark A.|issue=6|display-authors=8}}
* {{Citation|last1=Alvarez|year=2009|title=Y-chromosome variation in South Iberia: Insights into the North African contribution|doi=10.1002/ajhb.20888|volume=21|issue=3|pages=407–409|pmid=19213004|last2=Santos|first2=Cristina|last3=Montiel|first3=Rafael|last4=Caeiro|first4=Blazquez|last5=Baali|first5=Abdellatif|last6=Dugoujon|first6=Jean-Michel|last7=Aluja|first7=Maria Pilar|journal=American Journal of Human Biology|s2cid=7041905|doi-access=free}}
*{{Citation
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*{{Citation | author=Behar et al. | year=2003 | title=Multiple Origins of Ashkenazi Levites: Y Chromosome Evidence for Both Near Eastern and European Ancestries | periodical=Am. J. Hum. Genet. | volume=73 | pages=768–779 | id=PMID 13680527 | url=http://www.journals.uchicago.edu/AJHG/journal/issues/v73n4/40097/40097.html | doi=10.1086/378506 | format={{dead link|date=April 2009}} &ndash; <sup></sup>}}. Also at http://www.ucl.ac.uk/tcga/tcgapdf/Behar-AJHG-03.pdf and http://www.familytreedna.com/pdf/400971.pdf
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*{{Citation
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{{Refend}} {{Refend}}


===Sources for conversion tables===
== External links ==
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{{refbegin|2}}
]
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]
{{refend}}
]
]


] {{DEFAULTSORT:Haplogroup E1b1b (Y-Dna)}}
]
]

Latest revision as of 10:07, 28 December 2024

Human Y-chromosome DNA haplogroup "E3b" redirects here. For the Pennsylvania Railroad locomotive, see PRR E3b.
Haplogroup
  • E-M215
  • E1b1b
Geographic distribution of the haplogroup E1b1b
Possible time of origin47,500—22,400 BP
Coalescence age34,800 BP
Possible place of originNortheast Africa
AncestorE-P2
Descendants
Defining mutationsM215

E-M215 or E1b1b, formerly known as E3b, is a major human Y-chromosome DNA haplogroup. E-M215 has two basal branches, E-M35 and E-M281. E-M35 is primarily distributed in North Africa and the Horn of Africa, and occurs at moderate frequencies in the Middle East, Europe, and Southern Africa. E-M281 occurs at a low frequency in Ethiopia.

Origins

The origins of E-M215 were dated by Cruciani in 2007 to about 22,400 years ago in East Africa.

E1b1b1 origins map
E1b1b1 origins map

Ancient DNA

According to Lazaridis et al. (2016), Natufian skeletal remains from the ancient Levant predominantly carried the Y-DNA haplogroup E1b1b. Of the five Natufian specimens analyzed for paternal lineages, three belonged to the E1b1b1b2(xE1b1b1b2a, E1b1b1b2b), E1b1(xE1b1a1, E1b1b1b1) and E1b1b1b2(xE1b1b1b2a, E1b1b1b2b) subclades (60%). Haplogroup E1b1b was also found at moderate frequencies among fossils from the ensuing Pre-Pottery Neolithic B culture, with the E1b1b1 and E1b1b1b2(xE1b1b1b2a, E1b1b1b2b) subclades observed in two of seven PPNB specimens (~29%). The scientists suggest that the Levantine early farmers may have spread southward into East Africa, bringing along Western Eurasian and Basal Eurasian ancestral components separate from that which would arrive later in North Africa.

Additionally, haplogroup E1b1b1 has been found in an ancient Egyptian mummy excavated at the Abusir el-Meleq archaeological site in Middle Egypt, which dates from a period between the late New Kingdom and the Roman era. Fossils at the Iberomaurusian site of Ifri N'Amr Ou Moussa in Morocco, which have been dated to around 5,000 BCE, also carried haplotypes related to the E1b1b1b1a (E-M81) subclade. These ancient individuals bore an autochthonous Maghrebi genomic component that peaks among modern North Africans, indicating that they were ancestral to populations in the area. The E1b1b haplogroup has likewise been observed in ancient Guanche fossils excavated in Gran Canaria and Tenerife on the Canary Islands, which have been radiocarbon-dated to between the 7th and 11th centuries CE. The clade-bearing individuals that were analysed for paternal DNA were inhumed at the Tenerife site, with all of these specimens found to belong to the E1b1b1b1a1 or E-M183 subclade (3/3; 100%).

Loosdrecht et al. (2018) analysed genome-wide data from seven ancient Iberomaurusian individuals from the Grotte des Pigeons near Taforalt in eastern Morocco. The fossils were directly dated to between 15,100 and 13,900 calibrated years before present. The scientists found that five male specimens with sufficient nuclear DNA preservation belonged to the E1b1b1a1 (M78) subclade, with one skeleton bearing the E1b1b1a1b1 parent lineage to E-V13, another male specimen belonged to E1b1b (M215*).

Distribution

In Africa, E-M215 is distributed in highest frequencies in the Horn of Africa and North Africa, specifically in the countries Somalia and Morocco, whence it has in recent millennia expanded as far south as South Africa, and northwards into Western Asia and Europe (especially the Mediterranean and the Balkans). E-M281 has been found in Ethiopia.

Almost all E-M215 men are also in E-M35. In 2004, M215 was found to be older than M35 when individuals were found who have the M215 mutation, but do not have M35 mutation. In 2013, Di Cristofaro et al. (2013) found one individual in Khorasan, North-East Iran to be positive for M215 but negative for M35.

E-M215 and E-M35 are quite common among Afroasiatic speakers. The linguistic group and carriers of E-M35 lineage have a high probability to have arisen and dispersed together from the Afroasiatic Urheimat. Amongst populations with an Afro-Asiatic speaking history, a significant proportion of Jewish male lineages are E-M35. Haplogroup E-M35, which accounts for approximately 18% to 20% of Ashkenazi and 8.6% to 30% of Sephardi Y-chromosomes, appears to be one of the major founding lineages of the Jewish population.

E-M215 association with endurance

Moran et al. (2004) observed that among Y-DNA (paternal) clades borne by elite endurance athletes in Ethiopia, the haplogroup E3b1 was negatively correlated with elite athletic endurance performance, whereas the haplogroups E*, E3*, K*(xP), and J*(xJ2) were significantly more frequent among the elite endurance athletes.

Subclades

E-M35

Main article: Haplogroup E-M35

Haplogroup E-M35 is a subclade of E-M215.

E-M281

Main article: Haplogroup E-M281

Haplogroup E-M281 is a subclade of E-M215.

Phylogenetics

Phylogenetic history

Main article: Conversion table for Y chromosome haplogroups

Prior to 2002, there were in academic literature at least seven naming systems for the Y-Chromosome phylogenetic tree. This led to considerable confusion. In 2002, the major research groups came together and formed the Y-Chromosome Consortium (YCC). They published a joint paper that created a single new tree that all agreed to use. Later, a group of citizen scientists with an interest in population genetics and genetic genealogy formed a working group to create an amateur tree aiming at being above all timely. The table below brings together all of these works at the point of the landmark 2002 YCC Tree. This allows a researcher reviewing older published literature to quickly move between nomenclatures.

YCC 2002/2008 (Shorthand) (α) (β) (γ) (δ) (ε) (ζ) (η) YCC 2002 (Longhand) YCC 2005 (Longhand) YCC 2008 (Longhand) YCC 2010r (Longhand) ISOGG 2006 ISOGG 2007 ISOGG 2008 ISOGG 2009 ISOGG 2010 ISOGG 2011 ISOGG 2012
E-P29 21 III 3A 13 Eu3 H2 B E* E E E E E E E E E E
E-M33 21 III 3A 13 Eu3 H2 B E1* E1 E1a E1a E1 E1 E1a E1a E1a E1a E1a
E-M44 21 III 3A 13 Eu3 H2 B E1a E1a E1a1 E1a1 E1a E1a E1a1 E1a1 E1a1 E1a1 E1a1
E-M75 21 III 3A 13 Eu3 H2 B E2a E2 E2 E2 E2 E2 E2 E2 E2 E2 E2
E-M54 21 III 3A 13 Eu3 H2 B E2b E2b E2b E2b1 - - - - - - -
E-P2 25 III 4 14 Eu3 H2 B E3* E3 E1b E1b1 E3 E3 E1b1 E1b1 E1b1 E1b1 E1b1
E-M2 8 III 5 15 Eu2 H2 B E3a* E3a E1b1 E1b1a E3a E3a E1b1a E1b1a E1b1a E1b1a1 E1b1a1
E-M58 8 III 5 15 Eu2 H2 B E3a1 E3a1 E1b1a1 E1b1a1 E3a1 E3a1 E1b1a1 E1b1a1 E1b1a1 E1b1a1a1a E1b1a1a1a
E-M116.2 8 III 5 15 Eu2 H2 B E3a2 E3a2 E1b1a2 E1b1a2 E3a2 E3a2 E1b1a2 E1b1a2 E1ba12 removed removed
E-M149 8 III 5 15 Eu2 H2 B E3a3 E3a3 E1b1a3 E1b1a3 E3a3 E3a3 E1b1a3 E1b1a3 E1b1a3 E1b1a1a1c E1b1a1a1c
E-M154 8 III 5 15 Eu2 H2 B E3a4 E3a4 E1b1a4 E1b1a4 E3a4 E3a4 E1b1a4 E1b1a4 E1b1a4 E1b1a1a1g1c E1b1a1a1g1c
E-M155 8 III 5 15 Eu2 H2 B E3a5 E3a5 E1b1a5 E1b1a5 E3a5 E3a5 E1b1a5 E1b1a5 E1b1a5 E1b1a1a1d E1b1a1a1d
E-M10 8 III 5 15 Eu2 H2 B E3a6 E3a6 E1b1a6 E1b1a6 E3a6 E3a6 E1b1a6 E1b1a6 E1b1a6 E1b1a1a1e E1b1a1a1e
E-M35 25 III 4 14 Eu4 H2 B E3b* E3b E1b1b1 E1b1b1 E3b1 E3b1 E1b1b1 E1b1b1 E1b1b1 removed removed
E-M78 25 III 4 14 Eu4 H2 B E3b1* E3b1 E1b1b1a E1b1b1a1 E3b1a E3b1a E1b1b1a E1b1b1a E1b1b1a E1b1b1a1 E1b1b1a1
E-M148 25 III 4 14 Eu4 H2 B E3b1a E3b1a E1b1b1a3a E1b1b1a1c1 E3b1a3a E3b1a3a E1b1b1a3a E1b1b1a3a E1b1b1a3a E1b1b1a1c1 E1b1b1a1c1
E-M81 25 III 4 14 Eu4 H2 B E3b2* E3b2 E1b1b1b E1b1b1b1 E3b1b E3b1b E1b1b1b E1b1b1b E1b1b1b E1b1b1b1 E1b1b1b1a
E-M107 25 III 4 14 Eu4 H2 B E3b2a E3b2a E1b1b1b1 E1b1b1b1a E3b1b1 E3b1b1 E1b1b1b1 E1b1b1b1 E1b1b1b1 E1b1b1b1a E1b1b1b1a1
E-M165 25 III 4 14 Eu4 H2 B E3b2b E3b2b E1b1b1b2 E1b1b1b1b1 E3b1b2 E3b1b2 E1b1b1b2a E1b1b1b2a E1b1b1b2a E1b1b1b2a E1b1b1b1a2a
E-M123 25 III 4 14 Eu4 H2 B E3b3* E3b3 E1b1b1c E1b1b1c E3b1c E3b1c E1b1b1c E1b1b1c E1b1b1c E1b1b1c E1b1b1b2a
E-M34 25 III 4 14 Eu4 H2 B E3b3a* E3b3a E1b1b1c1 E1b1b1c1 E3b1c1 E3b1c1 E1b1b1c1 E1b1b1c1 E1b1b1c1 E1b1b1c1 E1b1b1b2a1
E-M136 25 III 4 14 Eu4 H2 B E3ba1 E3b3a1 E1b1b1c1a E1b1b1c1a1 E3b1c1a E3b1c1a E1b1b1c1a1 E1b1b1c1a1 E1b1b1c1a1 E1b1b1c1a1 E1b1b1b2a1a1

Research publications

The following research teams per their publications were represented in the creation of the YCC Tree.

Discussion

E-M215 and E1b1b1 are the currently accepted names found in the proposals of the Y Chromosome Consortium (YCC), for the clades defined by mutation M215 and M35 respectively, which can also be referred to as E-M215 and E-M35. The nomenclature E3b (E-M215) and E3b1 (E-M35) respectively were the YCC defined names used to designate the same haplogroups in older literature with E-M35 branching as a separate subclade of E-M215 in 2004. Prior to 2002 these haplogroups were not designated in a consistent way, and nor was their relationship to other related clades within haplogroup E and haplogroup DE. But in non-standard or older terminologies, E-M215 is for example approximately the same as "haplotype V", still used in publications such as Gérard et al. (2006).

Phylogenetic trees

Cladogram with the main subclades:

E1b1b (M215) 
E1b1b1 (M35)
E1b1b1a (V68)
E-M78

E-V12

E-V65

E-V13

E-V22

E-V2729

E1b1b1b (Z827)

E-M81

E-M123

E1b1b2 (M281)

The following phylogenetic tree is based on the YCC 2008 tree and subsequent published research as summarized by ISOGG. It includes all known subclades as of June 2015 (Trombetta et al. 2015)

  • E-M215 (E1b1b)
    • E-M215*. Rare or non-existent.
    • E-M35 (E1b1b1)
      • E-V68 (E1b1b1a)
        • E-V2009. Found in individuals in Sardinia and Morocco.
        • E-M78 (E1b1b1a1). North Africa, Horn of Africa, West Asia, Sicily. (Formerly "E1b1b1a".)
          • E-M78*
          • E-V1477. Found in Tunisian Jews.
          • E-V1083.
            • E-V1083*. Found only in Eritrea (1.1%) and Sardinia (0.3%).
            • E-V13
            • E-V22
          • E-V1129
            • E-V12
              • E-V12*
              • E-V32
            • E-V264
              • E-V259. Found in North Cameroon.
              • E-V65
                • E-CTS194
      • E-Z827 (E1b1b1b)
        • E-V257/L19 (L19, V257) – E1b1b1b1
          • E-PF2431
          • E-M81 (M81)
            • E-PF2546
              • E-PF2546*
              • E-CTS12227
                • E-MZ11
                  • E-MZ12
              • E-A929
                • E-Z5009
                  • E-Z5009*
                  • E-Z5010
                  • E-Z5013
                    • E-Z5013*
                    • E-A1152
                • E-A2227
                  • E-A428
                  • E-MZ16
                • E-PF6794
                  • E-PF6794*
                  • E-PF6789
                    • E-MZ21
                    • E-MZ23
                    • E-MZ80
                • E-A930
                • E-Z2198/E-MZ46
                  • E-A601
                  • E-L351
        • E-Z830 (Z830) – E1b1b1b2
          • E-M123 (M123)
            • E-M34 (M34)
              • E-M84 (M84)
                • E-M136 (M136)
              • E-M290 (M290)
              • E-V23 (V23)
              • E-L791 (L791,L792)
          • E-V1515. E-V1515 and its subclades are mainly restricted to eastern Africa.
            • E-V1515*
            • E-V1486
              • E-V1486*
              • E-V2881
                • E-V2881*
                • E-V1792
                • E-V92
              • E-M293 (M293)
                • E-M293*
                • E-P72 (P72)
                • E-V3065*
            • E-V1700
              • E-V42 (V42)
              • E-V1785
                • E-V1785*
                • E-V6 (V6)
      • E-V16/E-M281 (E1b1b2). Rare. Found in individuals in Ethiopia, Yemen and Saudi Arabia.

See also

Genetics

Y-DNA E subclades

Y-DNA backbone tree

Phylogenetic tree of human Y-chromosome DNA haplogroups
This article needs to be updated. Please help update this article to reflect recent events or newly available information. (February 2021)
"Y-chromosomal Adam"
A00 A0-T 
A0 A1 
A1a A1b
A1b1 BT
B CT
DE CF
D E C F
F1  F-Y27277   F3  GHIJK
G HIJK
IJK H
IJ K
I   J     LT        K2 
I1   I2  J1   J2  L     T  K2e K2d K2c K2b   K2a
K2b1    P  K-M2313 
S   M     P1   NO1
P1c P1b P1a N O
R Q
Footnotes
  1. Van Oven M, Van Geystelen A, Kayser M, Decorte R, Larmuseau HD (2014). "Seeing the wood for the trees: a minimal reference phylogeny for the human Y chromosome". Human Mutation. 35 (2): 187–91. doi:10.1002/humu.22468. PMID 24166809. S2CID 23291764.
  2. International Society of Genetic Genealogy (ISOGG; 2015), Y-DNA Haplogroup Tree 2015. (Access date: 1 February 2015.)
  3. Haplogroup A0-T is also known as A-L1085 (and previously as A0'1'2'3'4).
  4. Haplogroup A1 is also known as A1'2'3'4.
  5. F-Y27277, sometimes known as F2'4, is both the parent clade of F2 and F4 and a child of F-M89.
  6. Haplogroup LT (L298/P326) is also known as Haplogroup K1.
  7. Between 2002 and 2008, Haplogroup T-M184 was known as "Haplogroup K2". That name has since been re-assigned to K-M526, the sibling of Haplogroup LT.
  8. Haplogroup K2b (M1221/P331/PF5911) is also known as Haplogroup MPS.
  9. Haplogroup K2b1 (P397/P399) is also known as Haplogroup MS, but has a broader and more complex internal structure.
  10. Haplogroup P (P295) is also klnown as K2b2.
  11. K-M2313*, which as yet has no phylogenetic name, has been documented in two living individuals, who have ethnic ties to India and South East Asia. In addition, K-Y28299, which appears to be a primary branch of K-M2313, has been found in three living individuals from India. See: Poznik op. cit.; YFull YTree v5.08, 2017, "K-M2335", and; PhyloTree, 2017, "Details of the Y-SNP markers included in the minimal Y tree" (Access date of these pages: 9 December 2017)
  12. Haplogroup S, as of 2017, is also known as K2b1a. (Previously the name Haplogroup S was assigned to K2b1a4.)
  13. Haplogroup M, as of 2017, is also known as K2b1b. (Previously the name Haplogroup M was assigned to K2b1d.)

Notes

  1. 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 E-M215. Semino et al. (2004): "This inference is further supported by the presence of additional Hg E lineal diversification and by the highest frequency of E-P2* and E-M35* in the same region. The distribution of E-P2* appears limited to eastern African peoples. The E-M35* lineage shows its highest frequency (19.2%) in the Ethiopian Oromo but with a wider distribution range than E-P2*." For E-M215 Cruciani et al. (2007) reduced their estimate to 22,400 from 25,600 in Cruciani et al. (2004), re-calibrating the same data.
  2. "Paragroup E-M35 * 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." Behar et al. (2004)

References

  1. ^ Trombetta 2015.
  2. Haber M, Jones AL, Connel BA, Asan, Arciero E, Huanming Y, Thomas MG, Xue Y, Tyler-Smith C (June 2019). "A Rare Deep-Rooting D0 African Y-chromosomal Haplogroup and its Implications for the Expansion of Modern Humans Out of Africa". Genetics. 212 (4): 1421–1428. doi:10.1534/genetics.119.302368. PMC 6707464. PMID 31196864.
  3. ^ Cruciani et al. (2007)
  4. "E-M215 YTree".
  5. Cruciani et al. (2004).
  6. Schuenemann, Verena J.; et al. (2017). "Ancient Egyptian mummy genomes suggest an increase of Sub-Saharan African ancestry in post-Roman periods". Nature Communications. 8: 15694. Bibcode:2017NatCo...815694S. doi:10.1038/ncomms15694. PMC 5459999. PMID 28556824.
  7. Fregel; et al. (2018). "Ancient genomes from North Africa evidence prehistoric migrations to the Maghreb from both the Levant and Europe". bioRxiv 10.1101/191569.
  8. Rodrı́guez-Varela; et al. (2017). "Genomic Analyses of Pre-European Conquest Human Remains from the Canary Islands Reveal Close Affinity to Modern North Africans". Current Biology. 27 (1–7): 3396–3402.e5. Bibcode:2017CBio...27E3396R. doi:10.1016/j.cub.2017.09.059. hdl:2164/13526. PMID 29107554.
  9. Van De Loosdrecht, Marieke; Bouzouggar, Abdeljalil; Humphrey, Louise; Posth, Cosimo; Barton, Nick; Aximu-Petri, Ayinuer; Nickel, Birgit; Nagel, Sarah; Talbi, El Hassan; El Hajraoui, Mohammed Abdeljalil; Amzazi, Saaïd; Hublin, Jean-Jacques; Pääbo, Svante; Schiffels, Stephan; Meyer, Matthias; Haak, Wolfgang; Jeong, Choongwon; Krause, Johannes (4 May 2018). "Pleistocene North African genomes link Near Eastern and sub-Saharan African human populations". Science. 360 (6388): 548–552. Bibcode:2018Sci...360..548V. doi:10.1126/science.aar8380. PMID 29545507. S2CID 206666517.
  10. ^ Cruciani et al. (2004)
  11. ^ Semino et al. (2004)
  12. Rosser et al. (2000)
  13. Firasat et al. (2006)
  14. Di Cristofaro, Julie; et al. (October 18, 2013). "Afghan Hindu Kush: Where Eurasian Sub-Continent Gene Flows Converge". PLOS ONE. 8 (10): e76748. Bibcode:2013PLoSO...876748D. doi:10.1371/journal.pone.0076748. ISSN 1932-6203. OCLC 5534533323. PMC 3799995. PMID 24204668. S2CID 16455960.
  15. Ehret, Keita & Newman (2004); Keita & Boyce (2005); Keita (2008).
  16. Behar et al. (2003)
  17. Behar et al. (2004)
  18. Shen et al. (2004)
  19. Adams et al. (2008)
  20. Nebel et al. (2001)
  21. ^ Moran, Colin N.; et al. (2004). "Y chromosome haplogroups of elite Ethiopian endurance runners". Human Genetics. 115 (6): 492–7. doi:10.1007/s00439-004-1202-y. PMID 15503146. S2CID 13960753. Retrieved 6 February 2017.
  22. ^ Karafet et al. (2008)
  23. ^ Y Chromosome Consortium "YCC" (2002)
  24. ISOGG (2011)
  25. ^ ISOGG 2015

Bibliography

Sources for conversion tables

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