Haplogroup E-M215 (Y-DNA)

"E3b" redirects here. For the Pennsylvania Railroad locomotive, see PRR E3b.
Haplogroup E-M215
Possible time of origin 42,600[1]
Possible place of origin North Africa or Horn of Africa[2][3]
Ancestor Haplogroup E-P2
Descendants Haplogroups E-M35 and E-M281
Defining mutations M215, most often found in conjunction with M35

Human Y-chromosome DNA haplogroup E-M215, also referred to in the literature by other names such as E1b1b and E3b (see further discussion below), is a major Y chromosome haplogroup. It is a division of the macro haplogroup E-M96, which is defined by the single-nucleotide polymorphism (SNP) mutation M215.[4][5][6] In other words, it is one of the major patrilineal lineages of humanity, linking from father-to-son back to a common male-line ancestor ("Y-chromosomal Adam"). It is a subject of discussion and study in genetics as well as genetic genealogy, archaeology, and historical linguistics.

E-M215 has two ancient branches that contain all known modern E-M215, E-M35 and E-M281 subclades. Of the latter two, the only branch that has been confirmed in a native population outside of Ethiopia is E-M35. E-M35 in turn has two known branches, haplogroup E-V68 and haplogroup E-Z827, which contain by far the majority of all modern E-M215 men. E-V68 and E-V257 have been found in highest numbers in North Africa and the Horn of Africa, but also in lower numbers in parts of the Middle East and Europe, and in isolated populations of Southern Africa.


The origins of E-M215 were dated by Cruciani in 2007 to about 22,400 years ago in the Horn of Africa.[7][Note 1] E-M35 was dated by Batini in 2015 to between 15,400 and 20,500 years ago.[8] In June 2015, Trombetta et al. reported a previously unappreciated large difference in the age between haplogroup E-M215 (38.6 kya; 95% CI 31.4-45.9 kya) and its sub-haplogroup E-M35 (25.0 kya; 95% CI 20.0-30.0 kya).[9]

The ancient dispersals of the major E-M35 lineages. The map shows the supposed earliest movements of E-M215 lineages as described in the most recent articles.[2][7][10][11]

All major sub-branches of E-M35 are thought to have originated in the same general area as the parent clade: in North Africa, the Horn of Africa, or nearby areas of the Near East. Some branches of E-M35 are assumed to have left Africa thousands of years ago, whereas others may have arrived from the Near East. For example, Underhill (2002) associates the spread have the haplogroup with the Neolithic Revolution, believing that the structure and regional pattern of E-M35 subclades potentially give "reagents with which to infer specific episodes of population histories associated with the Neolithic agricultural expansion". Battaglia et al. (2007) also estimate that E-M78 (called E1b1b1a1 in that paper) has been in Europe longer than 10,000 years. Accordingly, human remains excavated in a Spanish funeral cave dating from approximately 7,000 years ago were shown to be in this haplogroup.[12] Two more E-M78 have been found in the Neolitich Sopot and Lengyel cultures too.[13]

Concerning European E-M35 within this scheme, Underhill & Kivisild (2007) have remarked that E-M215 seems to represent a late-Pleistocene migration from North Africa to Europe over the Sinai Peninsula in Egypt.[Note 2] While this proposal remains uncontested, it has more recently been proposed by Trombetta et al. (2011) that there is also evidence for additional migration of E-M215 carrying men directly from Africa to southwestern Europe, via a maritime route (see below.)


In Africa, E-M215 is distributed in highest frequencies in the Horn of Africa and North Africa, 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).[2][3][14][15]

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.[2] In 2013, one individual in Khorasan, North-East Iran, was found by Di Cristofaro et al. (2013) 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.[16][17][18] Amongst populations with an Afro-Asiatic speaking history, a significant proportion of Jewish male lineages are E-M35.[19] Haplogroup E-M35, which accounts for approximately 18%[3] to 20%[20][21] of Ashkenazi and 8.6%[22] to 30%[3] of Sephardi Y-chromosomes, appears to be one of the major founding lineages of the Jewish population.[23][Note 3]

The following table only includes sample populations with more than 1% E-M215 men with all known subclades as of June 2015 including the recent clade (E-V1515) which was defined by Trombetta et al. 2015 and includes all the sub-Saharan haplogroups (E-V42, E-M293, E-V92, E-V6) reported as E-M35 basal clades in a previous phylogeny.[9]

Population N Region Language Total E-M215 E-V2009 E-M78* E-V1477 E-V1083* E-V13 E-V22 E-V12* E-V32 E-V259 E-V65 E-V257* E-M81 E-M123* E-M34 E-V1515* E-V1486* E-V2881* E-V1792 E-V92 E-M293* E-V3065 E-V42 E-V1785* E-V6 E-V16
Northern Africa
Moroccan Arabs221MoroccoAA/Semitic70.
Asni Berbers54MoroccoAA/Berber85.
Bouhria Berbers67MoroccoAA/Berber79.
Moyen Atlas Berbers187MoroccoAA/Berber89.
Marrakech Berbers27MoroccoAA/Berber92.
Souss Berbers65MoroccoAA/Berber98.
Ouarzazate Berbers31MoroccoAA/Berber54.
Mozabite Berbers67AlgeriaAA/Berber89.
Tunisian Jews10TunisiaVarious20.
Libyan Arabs10LibyaVarious50.
Libyan Jews23LibyaVarious26.
Northern Egyptians49EgyptAA/Semitic42.
Egyptian Berbers from Siwa93EgyptAA/Semitic18.
Egyptians from Baharia41EgyptAA/Semitic56.
Egyptians from Gurna Oasis34EgyptAA/Semitic17.
Southern Egyptians47EgyptAA/Semitic78.
Western/Central Africa
Daba29Cameroon (North)AA/Chadic3.
Guidar9Cameroon (North)AA/Chadic11.
Mandara82Cameroon (North)AA/Chadic2.
Shuwa Arabs5Cameroon (North)AA/Semitic20.
Fulbe from Cameroon76Cameroon (North)NC/Atlantic1.
Moundang21Cameroon (North)NC/Adamawa4.
Eastern Africa
Ethiopian Jews22EthiopiaAA/Cushitic31.
Nilotic Western Kenya11KenyaNS/Sudanic45.
Other Bantu17KenyaNC/Bantu11.
Southern Africa
Bantu8South AfricaNC/Bantu12.
Northern Portuguese50PortugalIE10.
Southern Portuguese49PortugalIE16.
Pasiegos from Cantabria56SpainIE42.
Southern Spaniards62SpainIE6.
Spanish Basques55SpainBasque3.
French Basques16FranceBasque6.
Northern Italians80ItalyIE11.
Central Italians356ItalyIE12.
Southern Italians141ItalyIE15.
Continental Greeks32GreeceIE28.
Sephardic Bulgarians20BulgariaIE5.
Near East
Sephardic Turkish19TurkeyA10.
Istanbul Turkish35TurkeyA17.
Southwestern Turkish40TurkeyA7.
Northeastern Turkish41TurkeyA2.
Central Anatolian61TurkeyA9.
Southeastern Turkish24TurkeyA8.
Erzurum Turkish25TurkeyA12.
Turkish Cypriots46TurkeyA23.
Druze Arabs28IsraelAA/Semitic14.
Arabs41United Arab EmirateAA/Semitic7.

Subclades of E-M215

Family Tree

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)[4][5][6]

Exceptional cases of men who are M215 positive but M35 negative ("E-M215*") have been discovered so far in two Amharas of Ethiopia and one Yemeni.[2][25] At least some of these men, perhaps all, are known since early 2011 to be in a rare sibling clade to E-M35, known as E-V16 or E-M281.[26] The discovery of M281 was announced by Semino et al. 2002, who found it in two Ethiopian Oromo. Trombetta et al. 2011 found 5 more Ethiopian individuals and an equivalent SNP to M281, V16. It was in the 2011 paper that the family tree position (M215+/M35-) was discovered as described above. The E-M215 derivative, E-M35 is defined by the M35 SNP. Individuals with no known subclade defining mutations are referred to as E-M35*. As of June 2015, there is an increasingly complex tree structure which divides most men in E-M35 into two branches: E-V68 and E-Z827.

The most frequently described subclades are E-M78, a part of E-V68, and E-M81, which is a branch of E-Z827. These two subclades represent the largest proportion of the modern E-M215 population. E-M78 is found over most of the range where E-M215 is found excluding Southern Africa. E-M81 is found mainly in North Africa. E-M123 is less common but widely scattered, with significant populations in specific parts of the Horn of Africa, the Levant, Arabia, Iberia, and Anatolia. A new clade (E-V1515) was defined by Trombetta et al. 2015, which originated about 12 kya (95% CI 8.6-16.4) in eastern Africa where it is currently mainly distributed. This clade includes the E-V42, E-M293, E-V92 and E-V6 subclades, which were identified as E-M35 basal clades in a previous phylogeny.[9]

Within E-M35, there are striking parallels between two haplogroups, E-V68 and E-V257. Both contain a lineage which has been frequently observed in Africa (E-M78 and E-M81, respectively) and a group of undifferentiated chromosomes that are mostly found in southern Europe. An expansion of E-M35 carriers, possibly from the Middle East as proposed by other authors, and split into two branches separated by the geographic barrier of the Mediterranean Sea, would explain this geographic pattern. However, the absence of E-V68* and E-V257* in the Middle East makes a maritime spread between northern Africa and southern Europe a more plausible hypothesis.

TMRCA of the major nodes in E-M215

TMRCA (kya) Trombetta 2015 YFull

E-V68 (E1b1b1a)

Main article: Haplogroup E-V68

E-V68, is dominated by its longer-known subclade E-M78. Three "E-V68*" individuals who are in E-V68 but not E-M78 have been reported in Sardinia, by Trombetta et al. 2011, when announcing the discovery of V68. The authors noted that because E-V68* was not found in the Middle Eastern samples, this appears to be evidence of maritime migration from Africa to southwestern Europe. E-M78 is a commonly occurring subclade, widely distributed in North Africa, the Horn of Africa, West Asia, (the Middle East and Near East) "up to Southern Asia",[7] and all of Europe.[27] The European distribution has a frequency peak centered in parts of the Balkans (up to almost 50% in some areas)[3][28] and Sicily, and declining frequencies evident toward western, central, and northeastern Europe.

Based on genetic STR variance data, Cruciani et al. 2007 suggests that E-M78 originated in the region of Egypt and Libya.[Note 4] about 18,600 years ago (17,300 - 20,000 years ago).[Note 5] Battaglia et al. 2008 describe Egypt as "a hub for the distribution of the various geographically localized M78-related subclades" and, based on archaeological data, they propose that the point of origin of E-M78 (as opposed to later dispersal from Egypt) may have been in a refugium which "existed on the border of present-day Sudan and Egypt, near Lake Nubia, until the onset of a humid phase around 8500 BC. The northward-moving rainfall belts during this period could have also spurred a rapid migration of Mesolithic foragers northwards in Africa, the Levant and ultimately onward to Asia Minor and Europe, where they each eventually differentiated into their regionally distinctive branches". Towards the south, Hassan et al. 2008 also explain evidence that some subclades of E-M78, specifically E-V12 and E-V22, "might have been brought to Sudan from North Africa after the progressive desertification of the Sahara around 6,000-8,000 years ago". And similarly, Cruciani et al. 2007 propose that E-M78 in Ethiopia, Somalia and surrounding areas, back-migrated to this region from the direction of Egypt after acquiring the E-M78 mutation.

Recently, E-M78 was dated by Trombetta et al. 2015. between 20,300 and 14,800 years ago.[9]

Subclades of E-M78

Listed here are the main subclades of M78 as of June 2015. Within the E-M78 subclade, Trombetta et al. 2015 allocated most of the former E-M78* chromosomes to three new distinct branches: E-V1083*, E-V1477 and E-V259. The first is a paragroup sister to clades E-V22 and E-V13. The mutation V1477 defines a new basal branch that has been observed only in one northern African sample. Finally, a sister clade of E-V12 defined by V264 includes E-V65 and V259, a new lineage distributed in central Africa.[9][29]

E-Z827 (E1b1b1b)

Main article: Haplogroup E-Z827

In human genetics, E-Z827, is the name of a major human Y-chromosome DNA haplogroup abundantly found in North Africa, particularly the Maghreb, and to a lesser extent in Northeast Africa, the Near East and Europe.

E-V257/E-L19 (E1b1b1b1)

E-V257* individuals in their samples who were E-V257, but not E-M81. A Borana from Kenya, a Marrakesh Berber, a Corsican, a Sardinian, a southern Spaniard and a Cantabrian. As mentioned above, Trombetta et al. 2011 propose that the absence of E-V257* in the Middle East makes a maritime movement from northern Africa to southern Europe the most plausible hypothesis so far to explain its distribution.


E-M81 is the most common Y-chromosome haplogroup in North Africa, dominated by its subclade E-M183. It is thought to have originated in the area of North Africa 14,200 years ago.[30] This haplogroup reaches a mean frequency of 42% in North Africa, decreasing in frequency from approximately 80% or more in some Moroccan Berber populations, including Saharawis, to approximately 10% to the east of this range in Egypt.[31][32][33] Because of its prevalence among these groups and also others such as Mozabite, Middle Atlas, Kabyle and other Berber groups, it is sometimes referred to as a genetic "Berber marker". Pereira et al. 2010 report high levels amongst Tuareg in two Saharan populations - 77.8% near Gorom-Gorom, in Burkina Faso, and 81.8% from Gosi in Mali. There was a much lower frequency of 11.1% in the vicinity of Tanut in Niger. E-M81 is also quite common among Maghrebi Arabic-speaking groups. It is generally found at frequencies around 45% in coastal cities of North Africa (Oran, Tizi Ouzou, Algiers, Tunis, Sousse).[31][34]

Distribution of E-M81 in select areas of Africa, Asia and Europe

In this key area from Egypt to the Atlantic Ocean, Arredi et al. 2004 report a pattern of decreasing microsatellite haplotype variation (implying greater lineage age in those areas) from East to West, accompanied by a substantial increasing frequency. At the eastern extreme of this core range, Kujanova et al. 2009 found M81 in 28.6% (10 out of 35 men) in el-Hayez in the Libyan Desert in Egypt.

Arredi et al. 2004 believe the pattern of distribution and variance to be consistent with the hypothesis of a post Paleolithic "demic diffusion" from the East. The ancestral lineage of E-M81 in their hypothesis could have been linked with the spread of Neolithic food-producing technologies from the Fertile Crescent via the Nile, although pastoralism rather than agriculture. E-M81 may also have been carried into its currently most common region together with a form of the proto-Afroasiatic language. On the basis of these possible links, the men who brought E-M81 into northwestern Africa may therefore have come from Asia, or they may represent a "local contribution to the North African Neolithic transition". But there is no autochthonous presence of E-M81 in the Near East, indicating that M81 most likely emerged from its parent clade M35 either in North Africa, or possibly as far south as the Horn of Africa.[18]

In Europe, E-M81 is widespread but rare, in the Iberian Peninsula Spain shows an average frequency of 4.3% (49/1140) in the Iberian Peninsula with frequencies reaching 9% in Galicia, 10% in Western Andalusia and Northwest Castile. However this study includes 153 individuals from Majorca, Minorca and Ibiza islands as well as 24 individuals from Gascony which are not in the Iberian Peninsula. Without these 177 individuals, average for Iberian Peninsula is 4.9% (47/963),[35] it is found at comparable levels to E-M78, with an average frequency of around 5%, and in some regions it is more common. Its frequencies are higher in the western half of the peninsula with frequencies reaching 8% in Extremadura and southern Portugal, 4% to 9% in Galicia, 14% in western Andalusia and 10% in northwest Castile and 9% to 17% in Cantabria.[22][36][37][38][39] The highest frequencies of this clade found so far in Europe were observed in the Valles Pasiegos from Cantabria, ranging from 18% (8/45)[39] to 41% (23/56).[2] An average frequency of 8.28% (54/652) has also been reported in the Spanish Canary Islands with frequencies over 10% in the three largest islands of Tenerife (10.68%), Gran Canaria (11.54%) and Fuerteventura (13.33%).[40]

E-M81 is also found in France,[2] 2.70% (15/555) overall with frequencies surpassing 5% in Auvergne (5/89) and Île-de-France (5/91),[41][42] in Sicily (approximately 2% overall, but up to 5% in Piazza Armerina),[43] and in very much lower frequency near Lucera (1.7%), in continental Italy,[38] possibly due to ancient migrations during the Islamic, Roman, and Carthaginian empires. In a 2014 study by Stefania Sarno et al. with 326 samples from Cosenza, Matera, Lecce and 5 Siclian provinces, E-M81 shows an average frequency of 1.5%, but the typical Maghrebin core haplotype 13-14-30-24-9-11-13 has been found in only two out of the five E-M81 individuals. These results, along with the negligible contribution from North-African populations revealed by the admixture-like plot analysis, suggest only a marginal impact of trans-Mediterranean gene flows on the current SSI genetic pool.[43][44]

E-M81 was also found in 2013 at 5.8% in a large sample of 1 204 Sardinians.[45]

As a result of its old world distribution, this subclade is found throughout Latin America, for example 6.1% in Cuba,[46] 5.4% in Brazil (Rio de Janeiro),[Note 6] and among Hispanic men from California and Hawaii 2.4%.[47]

In smaller numbers, E-M81 men can be found in areas in contact with [North Africa, both around the Sahara, in places like Sudan, and around the Mediterranean in places like Lebanon, Turkey, and amongst Sephardi Jews.

There are two recognized subclades of E-M81, although one is much more important than the other.


Underhill 2000 found one example of E-M107 in Mali.


E-M183 is extremely dominant within E-M81. Karafet et al. 2008 first described it as a subclade of E-M81. The known subclades of E-M183 include:

E-Z830 (E1b1b1b2)

This is a recently discovered subclade which has not yet been included in most haplogroup trees, E-Z830 includes the confirmed subclades of E-M123, E-V1515 (E-M293, E-V42, E-V6, E-V92), and E-Z830*, and is a sibling clade to E-L19. Currently, the E-M35 phylogeny project recognizes four distinct clusters of Z830* carriers, two of which are exclusively Jewish in origin. The remaining two are significantly smaller, and include scattered individuals in Germany, Spain, Latin America, Egypt, and Ethiopia.[48][49][50][51]

Main article: Haplogroup E-M123

E-M123 is mostly known for its major subclade E-M34, which dominates this clade.[Note 7]


A new clade (E-V1515) was defined by Trombetta et al. 2015, which originated about 12 kya (95% CI 8.6-16.4) in eastern Africa where it is currently mainly distributed. This clade includes all the sub-Saharan haplogroups (E-V42, E-M293, E-V92, E-V6) reported as E-M35 basal clades in a previous phylogeny.[9]


E-M293 is a subclade of E-V1515. It was first identified by ISOGG as the second clade within E-Z830. It was discovered before E-Z830, being announced in Henn 2008, which associated it with the spread of pastoralism from East Africa into Southern Africa. So far high levels have been found in specific ethnic groups in Tanzania and Southern Africa. Highest were the Datooga (43%), Khwe (Kxoe) (31%), Burunge (28%), and Sandawe peoples (24%). Henn (2008) in their study also found two Bantu-speaking Kenyan males with the M293 mutation.[10]

Other E-M215 subclades are rare in Southern Africa. The authors state "Without information about M293 in the Maasai, Hema, and other populations in Kenya, Sudan, and Ethiopia, we cannot pinpoint the precise geographic source of M293 with greater confidence. However, the available evidence points to present-day Tanzania as an early and important geographic locus of M293 evolution.". They also say that "M293 is only found in sub-Saharan Africa, indicating a separate phylogenetic history for M35.1 * (former) samples further north".

E-P72 appears in Karafet (2008). Trombetta et al. 2011 announced that this is a subclade of E-M293.


Trombetta et al. 2011 announced the discovery of E-V42 in two Beta Israel persons. It was suggested that it may be restricted to the region around Ethiopia. However, further testing by commercial DNA testing companies confirmed many positive results for this subclade in Saudi Arabia, Kuwait and one person in Portugal who has a root from Arabia.[52]


The E-V6 subclade of E-V1515 is defined by V6. Cruciani et al. (2004) identified a significant presence of these lineages in Ethiopia and also some in the neighboring Somalis. Among the Ethiopian and Somali samples, the highest were 14.7% among the Amhara and 16.7% among the Wolayta.

To the south, Tishkoff et al. (2007) identified one V6+ man in a sample of 35 Datooga of Tanzania. And further to the north, Dugoujon et al. (2009) identified another 6 men in a sample of 93 from the Siwa Oasis, which is a Berber population


Trombetta et al. 2011 announced the discovery of E-V92 in two Amharas. Like E-V6 and E-V42 it possibly only exists in the area of Ethiopia.


Phylogenetic history

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

Research publications

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


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.[5] 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.[2] 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).[6]

See also

Wikiquote has quotations related to: Haplogroup E-M215 (Y-DNA)


Y-DNA E Subclades

Y-DNA backbone tree

Phylogenetic tree of human Y-chromosome DNA haplogroups [χ 1][χ 2]
"Y-chromosomal Adam"
A00 A0-T [χ 3]
A0 A1 [χ 4]
A1a A1b
A1b1 BT
F1  F2  F3  GHIJK
IJ   K
I J    LT [χ 5]  K2
L T [χ 6] NO [χ 7] K2b [χ 8]     K2c  K2d  K2e [χ 9]
N   O   K2b1 [χ 10]     P
K2b1a[χ 11]     K2b1b K2b1c      M     P1 P2
K2b1a1   K2b1a2   K2b1a3 S [χ 12] Q   R
  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.
  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 A0'1'2'3'4.
  4. Haplogroup A1 is also known as A1'2'3'4.
  5. Haplogroup LT (L298/P326) is also known as Haplogroup K1.
  6. 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.
  7. Haplogroup NO (M214) is also known as Haplogroup K2a (although the present Haplogroup K2e was also previously known as "K2a").
  8. Haplogroup K2b (M1221/P331/PF5911) is also known as Haplogroup MPS.
  9. Haplogroup K2e (K-M147) was previously known as "Haplogroup X" and "K2a" (but is a sibling subclade of the present K2a, also known as Haplogroup NO).
  10. Haplogroup K2b1 (P397/P399) is similar to the former Haplogroup MS, but has a broader and more complex internal structure.
  11. Haplogroup K2b1a has also been known as Haplogroup S-P405.
  12. Haplogroup S (S-M230), also known as K2b1a4, was previously known as Haplogroup K5.


  1. Cruciani et al. (2004) Semino et al. (2004) 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. "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."Underhill & Kivisild (2007:547)
  3. "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)
  4. Cruciani et al. 2007 use the term Northeastern Africa to refer to Egypt and Libya, as shown in Table 1 of the study. Prior to Cruciani et al. 2007, Semino et al. 2004 East Africa as a possible place of origin of E-M78, based upon Ethiopian testing. This was because of the high frequency and diversity of E-M78 lineages in the region of Ethiopia. However, Cruciani et al. 2007 were able to study more data, including populations from North Africa who were not represented in the Semino et al. 2004 study, and found evidence that the E-M78 lineages which make up a significant proportion of some populations in that region, were relatively young branches (see E-V32 below). They therefore 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". So according to Cruciani et al. 2007 E-M35, the parent clade of E-M78, originated in East Africa, subsequently spread to Northeast Africa, and then there was a "back migration" of E-M215 chromosomes that had acquired the E-M78 mutation. Cruciani et al. 2007 therefore note this as evidence for "a corridor for bidirectional migrations" 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".
  5. 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 ASD method is used, while for a second "ρ method", used as a check, gives 13.7kya with a standard deviation of 2.3kya, but the difference between the two methods is only large for the age estimation of E-M78, not its subclades. The authors state that the big difference is "attributable to the relevant departure from a star-like structure because of repeated founder effects"
  6. (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 9.6% in Portugal, quite similar to the frequency found in Rio de Janeiro (5.4%) among European contributors." Silva et al. 2006
  7. As of 11 November 2008 for example, the E-M35 phylogeny project had records of four E-M123* tests, compared to 93 test results with E-M34.


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  2. 1 2 3 4 5 6 7 8 Cruciani et al. (2004)
  3. 1 2 3 4 5 Semino et al. (2004)
  4. 1 2 ISOGG (2011)
  5. 1 2 3 Karafet et al. (2008)
  6. 1 2 3 Y Chromosome Consortium "YCC" (2002)
  7. 1 2 3 Cruciani et al. (2007)
  8. Large-scale recent expansion of European patrilineages shown by population resequencing, Chiara Batini et al, nature.com, 2015
  9. 1 2 3 4 5 6 Trombetta et al. 2015, Phylogeographic refinement and large scale genotyping of human Y chromosome haplogroup E provide new insights into the dispersal of early pastoralists in the African continent
  10. 1 2 Henn et al. (2008)
  11. Hassan et al. (2008)
  12. Lacan et al. (2011)
  13. "Molecular genetic investigation of the Neolithic population history in the western Carpathian Basin" (PDF).
  14. Rosser et al. (2000)
  15. Firasat et al. (2006)
  16. Ehret et al. (2004)
  17. Keita & Boyce (2005)
  18. 1 2 Keita 2008
  19. Behar et al. (2003)
  20. Behar et al. (2004)
  21. Shen et al. (2004)
  22. 1 2 Adams et al. (2008)
  23. Nebel et al. (2001)
  24. 1 2 3 ISOGG 2015
  25. Cadenas et al. 2007
  26. Trombetta et al. 2011
  27. Cruciani et al. 2006
  28. Peričic et al. 2005
  29. Cruciani et al. 2006
  30. "E-M81 YTree". www.yfull.com. Retrieved 2016-05-23.
  31. 1 2 Arredi et al. 2004
  32. Alvarez et al. 2009
  33. Bosch et al. 2001
  34. Robino et al. 2008
  35. see table.
  36. Flores et al. 2005
  37. Beleza et al. 2006
  38. 1 2 Capelli et al. 2009
  39. 1 2 MacaMaca-Meyer et al. 2003
  40. Fregel et al. 2009, see table
  41. Ramos-Luisa et al. 2009
  42. Only men with French surname were analysed, in order to try to exclude more recent immigrants.
  43. 1 2 Di Gaetano et al. 2009
  44. "An Ancient Mediterranean Melting Pot: Investigating the Uniparental Genetic Structure and Population History of Sicily and Southern Italy". PLoS ONE. 9: e96074. doi:10.1371/journal.pone.0096074. This article contains quotations from this source, which is available under a Creative Commons Attribution 4.0 International (CC BY 4.0) license.
  45. Francalacci et al. (2013), Low-Pass DNA Sequencing of 1200 Sardinians Reconstructs European Y-Chromosome Phylogeny
  46. (8 out of 132), Mendizabal et al. 2008
  47. (7 out of 295), Paracchini et al. 2003
  48. http://www.haplozone.net/e3b/project/cluster/3
  49. http://www.haplozone.net/e3b/project/cluster/4
  50. http://www.haplozone.net/e3b/project/cluster/81
  51. http://www.haplozone.net/e3b/project/cluster/72
  52. http://www.haplozone.net/e3b/project/cluster/60

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