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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=]{{sfnp|Cruciani|La_Fratta|Santolamazza|Sellitto|2004}}{{sfn|Trombetta|2015}}
| origin-place = ]<ref>Semino et al. (2004), , '']'', 74: 1023–1034.</ref> or ]<ref name=isogg/>
|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 = E1b1 or E-P2
|ancestor=]
| descendants = E1b1b1 or M35
|descendants={{Hlist|]|E-M281}}
| mutations = M215, and most often also M35
|mutations=M215
}}
|map=E1b1b.png|caption=Geographic distribution of the haplogroup E1b1b}}


In ], '''Haplogroup E1b1b''' (formerly known as '''E3b''', as well as '''Haplogroup 21''' or '''Hg21''' and '''Eu4''') is a ] ], a subgroup of ]. It is defined by the ] (SNP) mutation M215<ref name=isogg/>. E1b1b is found in various forms in the ], ], the ], ], and the ]. '''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 ].


==Origins==
The terminology E1b1b is recent, having being proposed in 2008 by Karafet et. al. Since the 2002 "Y Chromosome Consortium" it had been widely referred to as E3b, and prior to that it was referred to as HG21 (Haplogroup 21).
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>


]
Concerning the defining SNP, almost all men with the mutation M215, also have the mutation M35, which is why M35 was sometimes referred to as the defining SNP. It was in Cruciani et. al.'s 2004 article that M215 was shown to be prior to M35, because there are some lineages which have the M215 mutation, but not M35.


==Ancient DNA==
However, most E1b1b (M215) lineages are also within E1b1b1 (M35). (As with other haplogroup names, the speed of new discoveries is leading many writers to use simpler names which simply refer to the SNP being considered, for example M215 or M35).
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>
==Origins==
According to Cruciani et al. (2004 and 2006), E1b1b and E1b1b1 are believed to have first appeared in the Horn of Africa approximately 26,000 years ago amongst populations that already had lineages with the mutations P2 (also referred to as PN2), as well as DYS391p, P179, P180, and P181<ref name=isogg/>.


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>
East Africa is thought to be the place of origin for haplogroup E1b1b because it has: 1) the highest number of different E1b1b clades, 2) a high frequency of this haplogroup and a high microsatellite diversity, and 3) the exclusive presence of the undifferentiated E1b1b* paragroup<ref>Cruciani et al. (2004), , '']'', 74: 1014-1022.</ref>. However, the fact that Haplogroup E1b1b's parent E clade is closely linked with ], which is not found in Africa, leaves open the possibility that E1b1b first arose in the Near or Middle East and was subsequently carried into Africa by a back migration<ref name=isogg/>.


==Distribution==
While most sub-branches of E1b1b are thought to have originated in Northeast and North Africa, E1b1b1a2 (V13*), the most common sub-clade of E1b1b1a (M78) among Europeans, appears to have been founded in the northern Levant/Anatolia before dispersing from the Balkans around Europe and the Mediterranean area (Cruciani ''et al''. 2007).


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" />
==Subclades==


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/>
Nearly all E1b1b lineages are within E1b1b1 (E-M35).
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>
The most current phylogeny of E1b1b1 (E-M35) includes the ancestral state (E1b1b1*) plus seven branches, which are defined by the following SNPs: M78, M81, M123, M281, V6 and P72<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>.


===E-M215 association with endurance===
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 (E-M215). E1b1b1 is by far the most common sub-clade of E1b1b in Europe, and generally outside of Africa. Together, E-M78 and E-M81 form a very significant part of all male lineages in Northeast and North Africa.


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"/>
A third very significant, but still less well-known, sub-clade of E1b1b1 is E1b1b1c (E-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>.


==Subclades==
A 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 Southern Africa, and is thought by the authors to include the majority of M35 lines which do not have M78, M81 or M123.


===E-M35===
Smaller E1b1b1 sub-clades recognized are defined by the SNP mutations M281, V6, and P72.
{{Main|Haplogroup E-M35}}


Haplogroup E-M35 is a subclade of E-M215.
===E1b1b1a (M78); formerly E3b1a===


===E-M281===
Estimations of age are difficult and vary greatly, but M78's age has been estimated at about 22 thousand years. This means it is not necessarily significantly younger than M215 itself. Like M215 and M35, M78 is thought have occurred in the Horn of Africa (Cruciani et. al. 2004). The most recent estimates are as follows:
{{Main|Haplogroup E-M281}}
<blockquote>
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)
</blockquote>


Haplogroup E-M281 is a subclade of E-M215.
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 M78 in Europe, see below concerning sub-clade V13.


==Phylogenetics==
E1b1b1a has been further divided into subclades by Fulvio Cruciani et. al. (2004, 2006, 2007), on the basis of the following SNP mutations, according to Karafet 2008, and .
*'''E1b1b1a1 (V12).''' The oldest sub-clade, found mainly in Southern ] (arose ca. 15.2 kya). 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 ] profiles.
::*'''E1b1b1a1a (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 (V32).''' This is a sub-clade of V12. It is prevalent in the Horn of Africa among ], ]s, and ]. Before the discovery of V32, (Cruciani et. al. 2004) referred to the same lineages as the "gamma cluster" which is estimated to have arisen ca. 8.5 thousand years ago.
*'''E1b1b1a2 (V13).''' (All known positive tests are also positive for V36.)
::This the the most prevalent clade among ], especially in the ], where the highest concentrations are amongst ]. Also ], ], ], and ] show high concentrations. 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.


===Phylogenetic history===
::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 (Cruciani 2007) - with the latter being considered a genetically isolated community, and therefore of particular interest.
{{main|Conversion table for Y chromosome haplogroups}}
::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 ] expansion. However Cruciani et. al. (2007) suggests that the timing for dispersal of European V13 from the Balkans may be much more recent, indeed no earlier than 5300 years ago, and this might therefore have been associated with immigrations into Europe associated with the Balkan ]. ]
::The ] is frequently also discussed in connection to V13, as a haplogroup with a seemingly very similar distribution and pre-history.
::There are two recognized sub-clades, both of which may be very small:
::*'''E1b1b1a2a.''' Defined by V27.
::*'''E1b1b1a2b.''' Defined by P65.
*'''E1b1b1a3 (V22).''' Prevalent in ] and ], with higher microsatellite variance (0.35 vs. 0.46, respectively) in Egypt. This clade comprises most of those classified in Cruciani et al's earlier (2004) "delta cluster". There are two recognized sub-clades:
::*'''E1b1b1a3a.''' Defined by M148.
::*'''E1b1b1a3b.''' Defined by V19.
*'''E1b1b1a4 (V65).''' This sub-clade, equivalent to the previously classified "beta cluster", is found in high levels in the ] regions of far northern Africa. 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 Northeast African origin for this lineage.


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.
===E1b1b1b (M81); formerly E3b1b, E3b2===
]s and ]]]
'''E1b1b1b (M81).''' The most common Y chromosome haplogroup in ]. Colloquially referred to as the "] marker" for its prevalence among ], ], ] and other ] groups. Also quite common among North African ] groups. Reaches frequencies of up to 80% in the ].


{| class="wikitable"
This haplogroup is also found in significant amounts in the ], Southern ] and ]<ref>Cruciani et al. (2004),</ref>, probably due to ancient migrations during the ], ], and ] empires.
! align="center" style="background:#c63;"|'''YCC 2002/2008 (Shorthand)'''
! align="center" style="background:#f96;"|'''(α)'''
! align="center" style="background:#f96;"|'''(β)'''
! align="center" style="background:#f96;"|'''(γ)'''
| align="center" style="background:#f96;"|'''(δ)'''
| align="center" style="background:#f96;"|'''(ε)'''
| align="center" style="background:#f96;"|'''(ζ)'''
| align="center" style="background:#f96"|'''(η)'''
| align="center" style="background:#f96"|'''YCC 2002 (Longhand)'''
| align="center" style="background:#c96;"|'''YCC 2005 (Longhand)'''
| align="center" style="background:#c96;"|'''YCC 2008 (Longhand)'''
| align="center" style="background:#c96;"|'''YCC 2010r (Longhand)'''
| align="center" style="background:#ff9;"|'''ISOGG 2006'''
| align="center" style="background:#ff9;"|'''ISOGG 2007'''
| align="center" style="background:#ff9;"|'''ISOGG 2008'''
| align="center" style="background:#ff9;"|'''ISOGG 2009'''
| align="center" style="background:#ff9;"|'''ISOGG 2010'''
| align="center" style="background:#ff9;"|'''ISOGG 2011'''
| align="center" style="background:#ff9;"|'''ISOGG 2012'''
|-
| ]||21||III||3A||13||Eu3||H2||B||E*||E||E||E||E||E||E||E||E||E||E
|-
| ]||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
|-
| ]||21||III||3A||13||Eu3||H2||B||E2a||E2||E2||E2||E2||E2||E2||E2||E2||E2||E2
|-
| ]||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
|-
| ]||8||III||5||15||Eu2||H2||B||E3a*||E3a||E1b1||E1b1a||E3a||E3a||E1b1a||E1b1a||E1b1a||E1b1a1||E1b1a1
|-
| ]||8||III||5||15||Eu2||H2||B||E3a1||E3a1||E1b1a1||E1b1a1||E3a1||E3a1||E1b1a1||E1b1a1||E1b1a1||E1b1a1a1a||E1b1a1a1a
|-
| ]||8||III||5||15||Eu2||H2||B||E3a2||E3a2||E1b1a2||E1b1a2||E3a2||E3a2||E1b1a2||E1b1a2||E1ba12||removed||removed
|-
| ]||8||III||5||15||Eu2||H2||B||E3a3||E3a3||E1b1a3||E1b1a3||E3a3||E3a3||E1b1a3||E1b1a3||E1b1a3||E1b1a1a1c||E1b1a1a1c
|-
| ]||8||III||5||15||Eu2||H2||B||E3a4||E3a4||E1b1a4||E1b1a4||E3a4||E3a4||E1b1a4||E1b1a4||E1b1a4||E1b1a1a1g1c||E1b1a1a1g1c
|-
| ]||8||III||5||15||Eu2||H2||B||E3a5||E3a5||E1b1a5||E1b1a5||E3a5||E3a5||E1b1a5||E1b1a5||E1b1a5||E1b1a1a1d||E1b1a1a1d
|-
| ]||8||III||5||15||Eu2||H2||B||E3a6||E3a6||E1b1a6||E1b1a6||E3a6||E3a6||E1b1a6||E1b1a6||E1b1a6||E1b1a1a1e||E1b1a1a1e
|-
| ]||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
|-
| ]||25||III||4||14||Eu4||H2||B||E3b1a||E3b1a||E1b1b1a3a||E1b1b1a1c1||E3b1a3a||E3b1a3a||E1b1b1a3a||E1b1b1a3a||E1b1b1a3a||E1b1b1a1c1||E1b1b1a1c1
|-
| ]||25||III||4||14||Eu4||H2||B||E3b2*||E3b2||E1b1b1b||E1b1b1b1||E3b1b||E3b1b||E1b1b1b||E1b1b1b||E1b1b1b||E1b1b1b1||E1b1b1b1a
|-
| ]||25||III||4||14||Eu4||H2||B||E3b2a||E3b2a||E1b1b1b1||E1b1b1b1a||E3b1b1||E3b1b1||E1b1b1b1||E1b1b1b1||E1b1b1b1||E1b1b1b1a||E1b1b1b1a1
|-
| ]||25||III||4||14||Eu4||H2||B||E3b2b||E3b2b||E1b1b1b2||E1b1b1b1b1||E3b1b2||E3b1b2||E1b1b1b2a||E1b1b1b2a||E1b1b1b2a||E1b1b1b2a||E1b1b1b1a2a
|-
| ]||25||III||4||14||Eu4||H2||B||E3b3*||E3b3||E1b1b1c||E1b1b1c||E3b1c||E3b1c||E1b1b1c||E1b1b1c||E1b1b1c||E1b1b1c||E1b1b1b2a
|-
| ]||25||III||4||14||Eu4||H2||B||E3b3a*||E3b3a||E1b1b1c1||E1b1b1c1||E3b1c1||E3b1c1||E1b1b1c1||E1b1b1c1||E1b1b1c1||E1b1b1c1||E1b1b1b2a1
|-
| ]||25||III||4||14||Eu4||H2||B||E3ba1||E3b3a1||E1b1b1c1a||E1b1b1c1a1||E3b1c1a||E3b1c1a||E1b1b1c1a1||E1b1b1c1a1||E1b1b1c1a1||E1b1b1c1a1||E1b1b1b2a1a1
|-
|}


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


The following research teams per their publications were represented in the creation of the YCC Tree.
There are two recognized sub-clades, although at this time it seems that the majority of M81 is "M81*", meaning that they are not in any of the known sub-clades.
{{columns-list|colwidth=22em|
::*'''E1b1b1b1 (M107).'''
* '''α''' {{harvp|Jobling|Tyler-Smith|2000}} and {{harvp|Kaladjieva|2001}}
::*'''E1b1b1b2 (M165).''' Also shows M183.
* '''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====
===E1b1b1c (M123); formerly E3b1c, E3b3===
This clade is mostly known for its major sub-clade E1b1b1c1 (M34). According to Cruciani et al. (2004), M34 is found at very small frequencies in North Africa and southeastern Europe, and has its highest concentration in ] and the ]. However, because the diversity is apparently low in Ethiopia, the authors suggest that M34 was likely introduced into Ethiopia from the Near East. This article located one M123* individual in Bulgaria after testing 3401 individuals from five continents, and Underhill et. al. (2000) located one individual in Central Asia.


'''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>
Two subclades of E1b1b1c1 are recognized:
::*'''E1b1b1c1a.''' Defined by SNP mutations M84 and M136
::*'''E1b1b1c1b.''' Defined by SNP mutation M290


===E1b1b1d (M281)=== ====Phylogenetic trees====


Cladogram with the main subclades:
This SNP was discussed in Underhill et al. (2000) and Semino et al. (2002), but Cruciani et. al. (2004) found no cases of this SNP.


{{Clade
===E1b1b1e (V6)===
| label1= '''E1b1b''' ('''M215''')&nbsp;
Cruciani et al (2004) identified a significant presence of these lineages in Ethiopia, but apparently not elsewhere.
| 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"/>
===E1b1b1f (P72)===


* '''E-M215''' (E1b1b)
Appears in Karafet et. al. (2008).
** '''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.


===E-M293=== ==See also==


{{wikiquote}}
This sub-clade is associated by the authors who announced it with the spread of ] into ]. See Henn et. al. (2008). The authors referred to this sub-clade with the proposed name E3b1f. But this name was already out of date by the time the article was published because E1b1b1 is the new name for E3b1, the clade defined by SNP M35, and also because the sub-clade under E-M35 with the name "f" had already been proposed in Karafet 2008, for SNP P72.


==See also== ===Genetics===

{{columns-list|colwidth=22em|
*]
*]
*]
*]
*]
*]
*] *]
* ] *]
*]
*]
*]
*]
*]
}}

===Y-DNA E subclades===

{{columns-list|colwidth=22em|
*]
* ]
* ]
* Haplogroup E-M215
* ]
* ]
* ]
* ]
* ]
* ]
* ]
* ]
* ]
* ]
* ]
* ]
* ]
* ]
* ]
}}

===Y-DNA backbone tree===

{{Y-DNA}} {{Y-DNA}}


==Notes== ==Notes==

{{reflist}}
{{reflist|group="Note"}}


==References== ==References==


{{Reflist|30em}}
*Arredi et al. (2004) , '']'', 75: 338–345.

*Cruciani et al. (2004) , '']'', 74: 1014-1022.
==Bibliography==
*Cruciani et al. (2006) , ''Human Mutation''.
*Cruciani et al. (2007) , ''Molecular Biology and Evolution'', 24:1300-1311.
*Henn et. al. (2008) Y-chromosomal evidence of a pastoralist migration through Tanzania to southern Africa. See and .
*Karafet et al, (2008) New Binary Polymorphisms Reshape and Increase Resolution of the Human Y-Chromosomal Haplogroup Tree. . Genome Research, published online April 2, 2008.
*Luis et al. (2004) , '']'', 74: 532-544.
*Peričic et al. (2005) , Molecular Biology and Evolution 2005 22(10):1964-1975
*Rosser et al. (2000) , '']'', 67: 1526-1543.
*Semino et al. (2004) , '']'', 74: 1023–1034.
*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
*Underhill PA, et. al. (2000) Y chromosome sequence variation and the history of human populations. Nat Genet 26:358–361.
*Underhill PA, et. al. (2001)
*YCC (2002) Genome Research, Vol. 12, Issue 2, 339-348, February 2002


{{Refbegin}}
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* {{Citation|last1=Firasat|year=2006|title=Y-chromosomal evidence for a limited Greek contribution to the Pathan population of Pakistan|doi=10.1038/sj.ejhg.5201726|journal=European Journal of Human Genetics|volume=15|pages=121–126|pmid=17047675|pmc=2588664|issue=1|last2=Khaliq|first2=Shagufta|last3=Mohyuddin|first3=Aisha|last4=Papaioannou|first4=Myrto|last5=Tyler-Smith|first5=Chris|last6=Underhill|first6=Peter A|last7=Ayub|first7=Qasim}}
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* {{Citation|last1=Flores|year=2005|title=Isolates in a corridor of migrations: a high-resolution analysis of Y-chromosome variation in Jordan|journal=J Hum Genet|volume=50|pages=435–441|doi=10.1007/s10038-005-0274-4|pmid=16142507|last2=Maca-Meyer|first2=Nicole|last3=Larruga|first3=Jose M.|last4=Cabrera|first4=Vicente M.|last5=Karadsheh|first5=Naif|last6=Gonzalez|first6=Ana M.|issue=9|doi-access=free}}
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* {{Citation|last1=Gonçalves|year=2005|title=Y-chromosome Lineages from Portugal, Madeira and Açores Record Elements of Sephardim and Berber Ancestry|journal=Annals of Human Genetics|volume=69|pages=443–454|url=http://www3.interscience.wiley.com/cgi-bin/fulltext/118682163/PDFSTART|doi=10.1111/j.1529-8817.2005.00161.x|pmid=15996172|first1=R|last2=Freitas|first2=A|last3=Branco|first3=M|last4=Rosa|first4=A|last5=Fernandes|first5=AT|last6=Zhivotovsky|first6=LA|last7=Underhill|first7=PA|last8=Kivisild|first8=T|last9=Brehm|first9=A|issue=Pt 4|hdl=10400.13/3018 |s2cid=3229760|hdl-access=free}}{{dead link|date=May 2022|bot=medic}}{{cbignore|bot=medic}}
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* {{Citation|last=Lancaster|first=Andrew|year=2009|url=http://www.jogg.info/51/files/Lancaster.pdf|journal=Journal of Genetic Genealogy|volume=5|issue=1|title=Y Haplogroups, Archaeological Cultures and Language Families: a Review of the Multidisciplinary Comparisons using the case of E-M35|access-date=2009-06-26|archive-date=2016-05-06|archive-url=https://web.archive.org/web/20160506150956/http://www.jogg.info/51/files/Lancaster.pdf|url-status=dead}}
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===Sources for conversion tables===
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{{refbegin|2}}
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* {{cite journal |last1=Capelli |first1=Cristian |last2=Wilson |first2=James F. |last3=Richards |first3=Martin |last4=Stumpf |first4=Michael P.H. |last5=Gratrix |first5=Fiona |display-authors=4 |title=A Predominantly Indigenous Paternal Heritage for the Austronesian-Speaking Peoples of Insular Southeast Asia and Oceania |journal=The American Journal of Human Genetics |date=February 2001 |volume=68 |issue=2 |pages=432–443 |doi=10.1086/318205 |pmid=11170891 |pmc=1235276 |ref={{harvid|Capelli|2001}}}}
* {{cite journal |last1=Hammer |first1=Michael F. |last2=Karafet |first2=Tatiana M. |last3=Redd |first3=Alan J. |last4=Jarjanazi |first4=Hamdi |last5=Santachiara-Benerecetti |first5=Silvana |display-authors=4 |title=Hierarchical Patterns of Global Human Y-Chromosome Diversity |journal=Molecular Biology and Evolution |date=1 July 2001 |volume=18 |issue=7 |pages=1189–1203 |doi=10.1093/oxfordjournals.molbev.a003906 |pmid=11420360 |ref={{harvid|Hammer|2001}}|doi-access=free }}
* {{cite journal |last1=Kaladjieva |first1=Luba |last2=Calafell |first2=Francesc |last3=Jobling |first3=Mark A |last4=Angelicheva |first4=Dora |last5=de Knijff |first5=Peter |display-authors=4 |title=Patterns of inter- and intra-group genetic diversity in the Vlax Roma as revealed by Y chromosome and mitochondrial DNA lineages |journal=European Journal of Human Genetics |date=February 2001 |volume=9 |issue=2 |pages=97–104 |doi=10.1038/sj.ejhg.5200597 |pmid=11313742 |s2cid=21432405 |ref={{harvid|Kaladjieva|2001}}|doi-access=free }}
* {{cite journal |last1=Karafet |first1=Tatiana |last2=Xu |first2=Liping |last3=Du |first3=Ruofu |last4=Wang |first4=William |last5=Feng |first5=Shi |display-authors=4 |title=Paternal Population History of East Asia: Sources, Patterns, and Microevolutionary Processes |journal=The American Journal of Human Genetics |date=September 2001 |volume=69 |issue=3 |pages=615–628 |doi=10.1086/323299 |pmid=11481588 |pmc=1235490 |ref={{harvid|Karafet|2001}}|doi-access=free }}
* {{citation |last1=Semino |year=2000 |doi=10.1126/science.290.5494.1155 |title=The Genetic Legacy of Paleolithic Homo sapiens sapiens in Extant Europeans: A Y Chromosome Perspective |first1=O. |journal=Science |volume=290 |issue=5494 |pages=1155–9 |pmid=11073453 |last2=Passarino |first2=G |last3=Oefner |first3=PJ |last4=Lin |first4=AA |last5=Arbuzova |first5=S |display-authors=4 |last6=Beckman |bibcode=2000Sci...290.1155S |ref={{harvid|Semino|2000}}}}
* {{cite journal |last1=Su |first1=Bing |last2=Xiao |first2=Junhua |last3=Underhill |first3=Peter |last4=Deka |first4=Ranjan |last5=Zhang |first5=Weiling |display-authors=4 |title=Y-Chromosome Evidence for a Northward Migration of Modern Humans into Eastern Asia during the Last Ice Age |journal=The American Journal of Human Genetics |date=December 1999 |volume=65 |issue=6 |pages=1718–1724 |doi=10.1086/302680 |pmid=10577926 |pmc=1288383 |ref={{harvid|Su|1999}}}}
* {{cite journal |last1=Underhill |first1=Peter A. |last2=Shen |first2=Peidong |last3=Lin |first3=Alice A. |last4=Jin |first4=Li |last5=Passarino |first5=Giuseppe |display-authors=4 |title=Y chromosome sequence variation and the history of human populations |journal=Nature Genetics |date=November 2000 |volume=26 |issue=3 |pages=358–361 |doi=10.1038/81685 |pmid=11062480 |s2cid=12893406 |ref={{harvid|Underhill|2000}}}}
{{refend}}


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

Category:
Haplogroup E-M215: Difference between revisions Add topic