Haplogroup E-M35

Haplogroup E-M35
Haplogroup E-M35
Possible time of origin	34,800 BP
Coalescence age	23,900 BP
Possible place of origin	Eastern Africa
Ancestor	E-M215
Descendants	E-V68, E-Z827
Defining mutations	M35

Last updated February 04, 2026

E-M35, also known as E1b1b1-M35, is a human Y-chromosome DNA haplogroup. E-M35 has two basal branches, E-V68 and E-Z827. E-V68 and E-Z827 are primarily distributed in North Africa and the Horn of Africa, and occur at lower frequencies in the Middle East, Europe, and Southern Africa.

Origins

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. E1b1bRoute.png — 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.

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) and estimated its origin to be in Horn of Africa, where the node separating the E-V38 and E-M215 branches occurs about 47,500 years ago (95% CI: 41.3–56.8 ka).^[2] E-M35 was dated by Batini in 2015 to between 15,400 and 20,500 years ago.^[8]

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 of 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. (2008) 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.^[9] Two more E-M78 have been found in the Neolithic Sopot and Lengyel cultures too.^[10]

Concerning E-M35 in Europe 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 North Africa to southwestern Europe, via a maritime route (see below.)

Ancient DNA

According to Lazaridis et al. (2016), Natufian skeletal remains from the ancient Levant predominantly carried the Y-DNA haplogroup E. Of the five Natufian specimens analysed for paternal lineages, one belonged to CT, one to E-M35, one to E-M215, one to E-Z830.^[11] Haplogroup E-M35 was also found the dominant marker among fossils from the ensuing Pre-Pottery Neolithic B culture, with the E-M78 and E-Z830 subclades observed in multiple PPNB specimens (~50%).^[12]

Following Fregel et al. (2017), two Neolithic farmers from the North African site of Ifri n Ammar ou Moussa belonged to haplogroup E-L19. The majority of their genomes traced back to Natufian and PPN ancestry , indicating a recent intrusion into North Africa from the Levant. Their presence is likely linked to the spread of pastoral technologies from the Levant into North Africa.^[13]

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 all belonged to the E1b1b1a1 (M78) subclade, with one skeleton bearing the E1b1b1a1b1 parent lineage to E-V13.^[14] Martiniano et al. (2022) later reassigned all the Taforalt samples to haplogroup E-M78 and none to E-L618, the predecessor to E-V13.^[15]

Haplogroup E-M35 has been identified among both Amorite and Akkadian remains, found in the sites of Megiddo,^[16] Ebla, and Alalakh.^[17] It is also attested among later Semitic populations, such as the Phoenicians who settled throughout the Mediterranean region, where it appears to have been the most prevalent lineage.^[18] Additionally, analyses of medieval Andalusian remains indicate that individuals examined for paternal DNA overwhelmingly belonged to the E-M81 subclade.^[19]^[20]^[21]

In Egypt, haplogroup E-V1515 has been identified in the remains of an elite Egyptian individual from the site of Nuerat, dating to the Old Kingdom period,^[22] while haplogroup E-V22 has been detected in an ancient Egyptian mummy excavated at the Abusir el-Meleq archaeological site in Middle Egypt, dated to between the late New Kingdom and the Roman era.^[23]

Distribution

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.^[24] Amongst populations with an Afro-Asiatic speaking history, a significant proportion of Palestinians and Jewish male lineages are E-M35.^[25] Haplogroup E-M35, which accounts for approximately 18%^[3] to 20%^[26]^[27] of 18% of Palestinians and Ashkenazi and 8.6%^[28] to 30%^[3] of Sephardi Y-chromosomes, appears to be one of the major founding lineages of the Palestinian and the Jewish population.^[29]^{[Note 3]}

The following table only includes sample populations with more than 1% E-M215 men with all known subclades as of June 2015. It contains the E-V1515 clade defined by Trombetta et al. 2015, and all the E1b1b subclades distributed below the Sahara (E-V42, E-M293, E-V92, E-V6), which were identified as E-M35 basal clades in a former phylogeny.^[2]

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
Zriba Arabs	31	Tunisia	AA/Semitic													100
Reguibate Arabs	20	Algeria	AA/Semitic													80
Kairouan Arabs	49	Tunisia	AA/Semitic													89.6
Jerba Arabs	47	Tunisia	AA/Semitic													93.6
Moroccan Arabs	55	Morocco	AA/Semitic	15.9	0.0	0.0	0.0	0.0	0.0	7.3	0.0	0.0	0.0	32.7	0.0	30.9	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Asni Berbers	54	Morocco	AA/Berber	85.2	0.0	0.0	0.0	0.0	0.0	3.7	0.0	0.0	0.0	0.0	1.9	79.6	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Bouhria Berbers	67	Morocco	AA/Berber	79.1	0.0	0.0	0.0	0.0	1.5	0.0	0.0	0.0	0.0	0.0	0.0	77.6	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Middle Atlas Berbers	69	Morocco	AA/Berber	81.2	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	10.1	0.0	71.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Marrakech Berbers	27	Morocco	AA/Berber	92.6	0.0	0.0	0.0	0.0	0.0	3.7	3.7	0.0	0.0	0.0	3.7	77.8	0.0	3.7	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Souss Berbers	34	Morocco	AA/Berber	79.4	2.9	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	76.5	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Ouarzazate Berbers	31	Morocco	AA/Berber	54.8	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	54.8	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Mozabite Berbers	67	Algeria	AA/Berber	89.6	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	1.5	0.0	86.6	0.0	1.5	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Tunisian Jews	10	Tunisia	AA/Semitic	20.0	0.0	0.0	10.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	10.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Libyan Arabs	10	Libya	AA/Semitic	20.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	20.0	0.0	30.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Libyan Jews	23	Libya	AA/Semitic	26.1	0.0	0.0	0.0	0.0	4.3	0.0	0.0	0.0	0.0	4.3	0.0	0.0	0.0	17.4	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Northern Egyptians	49	Egypt	AA/Semitic	20.9	0.0	0.0	0.0	0.0	2.0	16.3	4.1	2.0	0.0	0.0	0.0	4.1	4.1	10.2	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Egyptian Berbers from Siwa	93	Egypt	AA/Semitic	18.3	0.0	0.0	0.0	0.0	0.0	0.0	2.2	0.0	0.0	4.3	2.2	1.1	0.0	2.2	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	6.5	0.0
Egyptians from Baharia	41	Egypt	AA/Semitic	56.1	0.0	0.0	0.0	0.0	2.4	22.0	14.6	0.0	0.0	2.4	7.3	4.9	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	2.4	0.0
Egyptians from Gurna Oasis	34	Egypt	AA/Semitic	17.6	0.0	5.9	0.0	0.0	0.0	0.0	8.8	2.9	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Southern Egyptians	47	Egypt	AA/Semitic	78.7	0.0	0.0	0.0	0.0	0.0	0.0	74.5	0.0	0.0	0.0	0.0	0.0	2.1	2.1	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Western/Central Africa
Mandenka	16	Senegal	NC/Mande	6.3	0.0	0.0	0.0	0.0	0.0	0.0	6.3	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Tuareg	22	Niger	AA/Berber	13.6	0.0	0.0	0.0	0.0	0.0	0.0	4.5	0.0	0.0	0.0	0.0	9.1	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Daba	29	Cameroon (North)	AA/Chadic	3.4	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	3.4	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Guidar	9	Cameroon (North)	AA/Chadic	11.1	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	11.1	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Mandara	82	Cameroon (North)	AA/Chadic	2.4	0.0	0.0	0.0	0.0	0.0	0.0	1.2	0.0	0.0	0.0	1.2	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Shuwa Arabs	5	Cameroon (North)	AA/Semitic	20.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	20.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Fulbe from Cameroon	76	Cameroon (North)	NC/Atlantic	1.3	1.3	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Moundang	21	Cameroon (North)	NC/Adamawa	4.8	0.0	0.0	0.0	0.0	0.0	0.0	0.0	4.8	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Eastern Africa
Tigre	5	Eritrea	AA/Semitic	100.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	60.0	0.0	0.0	0.0	0.0	0.0	0.0	20.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	20.0	0.0
Nara	15	Eritrea	NS/Sudanic	60.0	0.0	0.0	0.0	0.0	0.0	6.7	0.0	13.3	0.0	0.0	0.0	0.0	0.0	0.0	13.3	6.7	0.0	0.0	0.0	0.0	0.0	13.3	0.0	6.7	0.0
Cunama	20	Eritrea	NS/Cunama	65.0	0.0	0.0	0.0	0.0	0.0	5.0	0.0	20.0	0.0	0.0	0.0	0.0	0.0	5.0	0.0	0.0	5.0	0.0	0.0	0.0	0.0	15.0	10.0	5.0	0.0
Saho	94	Eritrea	AA/Cushitic	98.9	0.0	0.0	0.0	1.1	0.0	88.3	0.0	0.0	0.0	0.0	0.0	0.0	0.0	1.1	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	8.5	0.0
Tigrai	32	Eritrea/Ethiopia	AA/Semitic	71.9	0.0	0.0	0.0	0.0	0.0	3.1	3.1	21.9	0.0	0.0	0.0	0.0	0.0	0.0	0.0	3.1	3.1	0.0	0.0	0.0	0.0	0.0	31.3	6.3	0.0
Afar	25	Djibouti	AA/Cushitic	60.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	4.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	56.0	0.0
Somali	40	Djibouti	AA/Cushitic	25.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	25.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Ethiopian Jews	22	Ethiopia	AA/Cushitic	31.8	0.0	0.0	0.0	0.0	0.0	0.0	0.0	9.1	0.0	0.0	0.0	0.0	0.0	13.6	0.0	0.0	0.0	0.0	0.0	0.0	0.0	9.1	0.0	0.0	0.0
Amhara	82	Ethiopia	AA/Semitic	45.1	0.0	0.0	0.0	0.0	0.0	2.4	0.0	11.0	0.0	0.0	0.0	0.0	0.0	13.4	0.0	0.0	2.4	0.0	1.2	0.0	0.0	1.2	0.0	8.5	4.9
Oromo	62	Ethiopia	AA/Cushitic	53.2	0.0	0.0	0.0	0.0	0.0	0.0	0.0	22.6	0.0	0.0	0.0	0.0	0.0	4.8	0.0	0.0	17.7	0.0	0.0	1.6	0.0	3.2	0.0	1.6	1.6
Wolayta	12	Ethiopia	AA/Omotic	58.3	0.0	0.0	0.0	0.0	0.0	8.3	0.0	8.3	0.0	0.0	0.0	0.0	0.0	8.3	0.0	0.0	8.3	0.0	0.0	8.3	0.0	0.0	0.0	16.7	0.0
Somali	12	Ethiopia	AA/Cushitic	50.0	0.0	0.0	0.0	0.0	0.0	8.3	0.0	25.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	16.7	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Gurage	7	Ethiopia	AA/Semitic	42.9	0.0	0.0	0.0	0.0	0.0	28.6	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	14.3	0.0
Somali	5	Somalia	AA/Cushitic	100.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	80.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	20.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Turkana	6	Kenya	NS/Sudanic	50.0	0.0	0.0	0.0	0.0	0.0	33.3	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	16.7	0.0	0.0	0.0	0.0	0.0
Borana	9	Kenya	AA/Cushitic	77.8	0.0	0.0	0.0	0.0	0.0	0.0	0.0	66.7	0.0	0.0	11.1	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Somali	6	Kenya	AA/Cushitic	100.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	66.7	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	16.7	0.0	0.0	0.0	0.0	0.0	0.0	16.7	0.0
Nilotic Western Kenya	11	Kenya	NS/Sudanic	45.5	0.0	0.0	0.0	0.0	0.0	9.1	9.1	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	18.2	0.0	0.0	0.0	9.1	0.0
Luhya	51	Kenya	NC/Bantu	9.8	0.0	0.0	0.0	0.0	0.0	0.0	0.0	5.9	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	3.9	0.0	0.0	0.0	0.0	0.0
Other Bantu	17	Kenya	NC/Bantu	11.8	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	11.8	0.0	0.0	0.0	0.0	0.0
Kikuyu	9	Kenya	NC/Bantu	11.1	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	11.1	0.0	0.0	0.0	0.0	0.0
Maasai	45	Kenya	NS/Sudanic	37.8	0.0	0.0	0.0	0.0	0.0	6.7	0.0	6.7	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	24.4	0.0	0.0	0.0	0.0	0.0
Tutsi	9	Burundi	NC/Bantu	22.2	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	22.2	0.0	0.0	0.0	0.0	0.0
Southern Africa
!Kung	64	Angola	KS	10.9	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	9.4	1.6	0.0	0.0	0.0	0.0
Khwe	26	Namibia	KS	30.8	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	30.8	0.0	0.0	0.0	0.0	0.0
Bantu	8	South Africa	NC/Bantu	12.5	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	12.5	0.0	0.0	0.0	0.0	0.0
Europe
Northern Portuguese	50	Portugal	IE	10.0	0.0	0.0	0.0	0.0	4.0	0.0	0.0	0.0	0.0	0.0	0.0	4.0	0.0	0.0	2.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Southern Portuguese	49	Portugal	IE	16.3	0.0	0.0	0.0	0.0	4.1	0.0	0.0	0.0	0.0	0.0	0.0	12.2	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Pasiegos from Cantabria	56	Spain	IE	42.9	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	1.8	41.1	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Asturians	90	Spain	IE	12.2	0.0	0.0	0.0	0.0	5.6	4.4	0.0	0.0	0.0	0.0	0.0	2.2	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Southern Spaniards	62	Spain	IE	6.5	0.0	0.0	0.0	0.0	0.0	3.2	0.0	0.0	0.0	0.0	1.6	1.6	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Spanish Basques	55	Spain	Basque	3.6	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	3.6	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
French	85	France	IE	8.2	0.0	0.0	0.0	0.0	3.5	0.0	1.2	0.0	0.0	0.0	0.0	3.5	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
French Basques	16	France	Basque	6.3	0.0	0.0	0.0	0.0	0.0	0.0	6.3	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Corsicans	140	France	IE	6.4	0.0	0.0	0.0	0.0	4.3	0.0	0.0	0.0	0.0	0.0	0.7	0.0	0.0	1.4	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Danish	35	Denmark	IE	2.9	0.0	0.0	0.0	0.0	2.9	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Germans	77	Germany	IE	3.9	0.0	0.0	0.0	0.0	3.9	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Northern Italians	80	Italy	IE	11.3	0.0	0.0	0.0	0.0	6.3	2.5	0.0	0.0	0.0	0.0	0.0	1.3	0.0	1.3	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Central Italians	356	Italy	IE	12.9	0.0	0.0	0.0	0.0	5.3	2.0	0.3	0.0	0.0	0.3	0.3	0.8	0.0	3.9	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Southern Italians	141	Italy	IE	15.6	0.7	0.0	0.0	0.0	8.5	1.4	0.7	0.0	0.0	0.0	0.0	1.4	0.0	2.8	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Sicilians	153	Italy	IE	20.3	0.0	0.0	0.0	0.0	7.2	4.6	0.7	0.0	0.0	0.7	0.0	0.7	0.0	6.5	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Sardinians	374	Italy	IE	8.3	0.8	0.0	0.0	0.3	1.1	0.8	0.3	0.0	0.0	1.1	0.3	0.3	0.0	3.5	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Polish	40	Poland	IE	2.5	0.0	0.0	0.0	0.0	2.5	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Slovenians	104	Slovenia	IE	2.9	0.0	0.0	0.0	0.0	2.9	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Estonians	74	Estonia	U	5.4	0.0	0.0	0.0	0.0	4.1	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Hungarians	106	Hungary	U	10.4	0.0	0.0	0.0	0.0	9.4	0.0	0.0	0.0	0.0	0.0	0.0	0.9	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Romanians	30	Romania	IE	26.7	0.0	0.0	0.0	0.0	26.7	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Macedonians	99	Macedonia	IE	18.2	0.0	0.0	0.0	0.0	18.2	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Continental Greeks	32	Greece	IE	28.1	0.0	0.0	0.0	0.0	25.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	3.1	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Bulgarians	112	Bulgaria	IE	22.3	0.0	0.0	0.0	0.0	21.4	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.9	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Sephardic Bulgarians	20	Bulgaria	IE	5.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	5.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Albanians	21	Albania	IE	33.3	0.0	0.0	0.0	0.0	33.3	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Near East
Sephardic Turkish	19	Turkey	A	10.5	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	5.3	0.0	5.3	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Istanbul Turkish	35	Turkey	A	17.1	0.0	0.0	0.0	0.0	2.9	5.7	0.0	0.0	0.0	0.0	0.0	5.7	0.0	2.9	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Southwestern Turkish	40	Turkey	A	7.5	0.0	0.0	0.0	0.0	2.5	0.0	0.0	0.0	0.0	0.0	0.0	2.5	0.0	2.5	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Northeastern Turkish	41	Turkey	A	2.4	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	2.4	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Central Anatolian	61	Turkey	A	9.8	0.0	0.0	0.0	0.0	4.9	0.0	1.6	0.0	0.0	0.0	0.0	0.0	0.0	3.3	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Southeastern Turkish	24	Turkey	A	8.3	0.0	0.0	0.0	0.0	4.2	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	4.2	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Erzurum Turkish	25	Turkey	A	12.0	0.0	0.0	0.0	0.0	0.0	0.0	4.0	0.0	0.0	0.0	0.0	0.0	0.0	8.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Turkish Cypriots	46	Turkey	A	23.9	0.0	0.0	0.0	0.0	10.9	2.2	0.0	0.0	0.0	0.0	0.0	8.7	0.0	2.2	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Bedouins	28	Israel	AA/Semitic	14.3	0.0	0.0	0.0	0.0	0.0	3.6	0.0	0.0	0.0	0.0	0.0	3.6	0.0	7.1	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Druze Arabs	28	Israel	AA/Semitic	14.3	0.0	0.0	0.0	0.0	10.7	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	3.6	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Palestinians	29	Israel	AA/Semitic	13.8	0.0	0.0	0.0	0.0	3.4	6.9	0.0	0.0	0.0	0.0	0.0	0.0	0.0	3.4	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Emiratis	41	United Arab Emirates	AA/Semitic	7.3	0.0	0.0	0.0	0.0	0.0	2.4	0.0	0.0	0.0	0.0	0.0	0.0	0.0	4.9	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Omanites	13	Oman	AA/Semitic	15.4	0.0	0.0	0.0	0.0	0.0	7.7	0.0	0.0	0.0	0.0	0.0	0.0	0.0	7.7	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0
Yemenites	94	Yemen	AA/Semitic	14.9	0.0	0.0	0.0	0.0	0.0	0.0	2.1	3.2	0.0	0.0	0.0	1.1	0.0	7.4	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	0.0	1.1

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.^[4]^[30] 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.^[31] 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. 1 Turkmen individual from Jawzjan with a subclade defining mutation is referred to as E-M35*.^[32] 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.^[2]

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.

— Trombetta et al. (2011)

TMRCA of the major nodes in E-M35

TMRCA (kya)	Trombetta 2015	YFull
E-M215	39	35,4
*E-M35	25	23,9
**E-V68	20	20
***E-M78	15	13
**E-Z827	?	23,6
***E-V257/L19	?	13,9
****E-M81	?	2,7
***E-Z830	20	19
****E-M34	?	15
****E-V1515	19	?

Subclades

E-V68 (E1b1b1a)

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",^[5] and all of Europe.^[33] The European distribution has a frequency peak centered in parts of the Balkans (up to almost 50% in some areas)^[3]^[34] 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.^[2]

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.^[2]^[33]

E-M78 (E1b1b1a1) North Africa, Horn of Africa, West Asia, Europe (formerly "E1b1b1a").
- E-M78* Found in Morocco, southern Portugal, southern Spain and Iran (Tehran and Semnan provinces).
- E-V1477 Found in Tunisian Jews.
- E-V1083
  - E-V1083* Found only in Eritrea (1.1%) and Sardinia (0.3%).
  - E-V13 This is the most common subclade of E-M215 found in Europe. It is especially common in the Balkans.
  - E-V22. Concentrated in Northeast Africa and the Near East. Peaks among the Saho.
- E-V1129
  - E-V12. Found in Egypt, Sudan, and Chad other places. Has an important subclade
    - E-V12* Most common lineage among Southern Egyptians (74.5%).
    - E-V32. Very common among Somalis, Tigre and Oromos.
  - E-V264
    - E-V259 Found in North Cameroon.
    - E-V65 Associated with North Africa, but also found in Sicily and also found in continental Italy.
- E-M521 Not mentioned by Trombetta et al.2015. Found in two individuals in Greece by Battaglia et al. (2008) and in one individual from the Eastern Alpine region of Italy by Coia et al. (2013)

E-Z827 (E1b1b1b)

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 Horn of Africa, the Near East and Europe.

E-L19 (E1b1b1b1)

To date, only two immediate downstream subclades of E-L19 have been recognized: E-PF2431 and E-M81.

E-M81

E-M81 is the dominant subclade of E-L19, accounting for more than 99% of its carriers. It has three main branches: E-M165, carried exclusively by a German and an Israeli;^[35] E-Y596059, carried by a Syrian from Damascus and an Iraqi Al Anbar; and E-CTS4236, carried exclusively by a Syrian from Quneitra, nearly all branches of E-M81 today derives from E-CTS4236.^[36] It is thought to have originated in the Near East,^[37]^[38]^[39]^[40]^[41]^[42] roughly 13,000 years ago, with subsequent expansions into Europe and North Africa occurring either about 1,000 years ago via Arab populations^[43] or around 3,000 years ago through Phoenician movements.^[44]

This distribution has been usually explained as a consequence of a westward expansion from the Near East and this event probably occurred in recent times, possibly about 2 kya [22]. We calculated the forensic indexes including only the E-M81 subjects and we could observe an even more enhanced clinal pattern (Moroccan DC = 0.87; Libyan DC = 0.96; the values for the Egyptian populations cannot be considered informative because our collection included only four E-M81 chromosomes from this country) (Supplementary Table 12).^[45]

This haplogroup has been identified in a variety of historically significant populations, including the ancient Phoenicians from Malaka and Selinunte,^[44]^[44] Medieval Arab communities from Al-Andalus and the Emirate of Sicily,^[43]^[46] as well as among the indigenous population of the Canary Islands.^[47]

**Ancient samples carrying the E-M81 lineage**
ID	Language	Culture	Country	Location	Date	Study
I19190	Semitic	Phoenician	Italy	Selinunte	700-400BC	Ringbauer at al. 2025^[48]
I3682	Semitic	Punico-Roman	Spain	Lucena	300-400AD	Carrion et al. 2024^[49]
I21367	Semitic	Punico-Roman	Spain	Malaga	416-700AD	Carrion et al. 2024^[49]
GUN012	Unknown	Guanche	Spain	Tenerife	593-660 AD	Rodriguez-Varela et al. 2017^[50]
SP7089	Semitic	Andalusian	France	Narbonne	637-765 AD	Gleize et al. 2016^[21]
SP7080	Semitic	Andalusian	France	Narbonne	684-761 AD	Gleize et al. 2016^[21]
78	Semitic	Andalusian	Spain	Pamplona	680-780 AD	de Miguel bánez 2016^[51]
107	Semitic	Andalusian	Spain	Pamplona	680-780 AD	de Miguel Ibanez 2016^[51]
GUN011	Unknown	Guanche	Spain	Tenerife	704-887 AD	Rodriguez-Varela et al. 2017^[50]
GOG23	Semitic	Andalusian	Spain	Vall Uixo	706-888 AD	Oteo-Garcia et al. 2024^[52]
CAN02	Unknown	Guanche	Spain	La Gomera	774-994 AD	Serrano et al. 2023^[53]
ST2933	Indo-European	Flemish	Belgium	Sint Truiden	1020-1160 AD	Beneker et al. 2025^[54]
CAN048	Unknown	Guanche	Spain	Tenerife	1028-1162 AD	Serrano et al. 2023^[53]
GUN002	Unknown	Guanche	Spain	Tenerife	1031-1159 AD	Rodriguez-Varela et al. 2017^[50]
SGBN3	Semitic	Andalusian	Italy	Segesta	1050-1215 AD	Monnereau et al. 2024^[55]
SGBN1	Semitic	Andalusian	Italy	Segesta	1050-1220 AD	Monnereau et al. 2024^[55]
112644	Semitic	Andalusian	Spain	Sagunto	1100-1300 AD	Olalde et al. 2019^[56]
112649	Semitic	Andalusian	Spain	Sagunto	1100-1300 AD	Olalde et al. 2019^[56]
CAN012	Unknown	Guanche	Spain	Gran Canaria	1169-1265 AD	Serrano et al. 2023^[53]
PT23221	Semitic	Andalusian	Portugal	Santarem	1200-1300 AD	Roca-Rada et al. 2024^[57]
CAN023	Unknown	Guanche	Spain	Gran Canaria	1221-1278 AD	Serrano et al. 2023^[53]
CAN041	Unknown	Guanche	Spain	Tenerife	1228-1297 AD	Serrano et al. 2023^[53]
CAN001	Unknown	Guanche	Spain	Tenerife	1272-1388 AD	Serrano et al. 2023^[53]
CAN030	Unknown	Guanche	Spain	La Gomera	1288-1396 AD	Serrano et al. 2023^[53]
CAN025	Unknown	Guanche	Spain	Gran Canaria	1303-1410 AD	Serrano et al. 2023^[53]
CAN043	Unknown	Guanche	Spain	Gran Canaria	1322-1437 AD	Serrano et al. 2023^[53]
CAN042	Unknown	Guanche	Spain	Tenerife	1327-1440 AD	Serrano et al. 2023^[53]
CAN045	Unknown	Guanche	Spain	Gran Canaria	1444-1625 AD	Serrano et al,.2023^[53]
ELW036	Indo-European	German	Germany	Ellwangen	1486-1627 AD	Immel et al. 2021^[58]
GOG59	Indo-European	Andalusian	Spain	Vall Uixo	1501-1600 AD	Oteo-Garcia et al. 2024^[20]
PT22190	Indo-European	Portuguese	Spain	Aveiro	1701-1800 AD	Roca-Rada et al. 2024^[57]

E-PF2431

E-PF2431 is a minor subclade of E-L19 mostly found in Europe, with smaller but notable frequencies in the Middle East/North Africa and sub-Saharan Africa. This haplogroup has been identified among Phoenicians from Motya^[59], ancient Romans from Pompeii,^[60] and Medieval Arabs.^[49]

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,^[61] 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.^[62]^[63]^[64]^[65]

E-M123

E-M123 is mostly known for its major subclade E-M34, which dominates this clade.^{[Note 6]}

E-V1515

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.^[2]

E-M293

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 et al. (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.^[6]

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.

E-V42

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.^[66]

E-V6

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

E-V92

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.

Phylogenetics

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
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.

α Jobling & Tyler-Smith (2000) and Kaladjieva (2001)
β Underhill (2000)
γ Hammer (2001)
δ Karafet (2001)
ε Semino (2000)
ζ Su (1999)
η Capelli (2001)

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)^[67]^[68]^[69]

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)^[70]
    - E-V257/L19 (L19, V257) – E1b1b1b1^[70]
      - 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^[70]
      - 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.

Notes

↑ Semino et al. (2004): "Both phylogeography and microsatellite variance suggest that E-P2 and its derivative, E-M35, probably originated in eastern Africa."
↑ "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)
↑ "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)
↑ 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".
↑ 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"
↑ As of 11 November 2008 for example, the E-M35 phylogeny project ^{[ permanent dead link ]} had records of four E-M123* tests, compared to 93 test results with E-M34.

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↑ ^[permanent dead link ]
↑ "E-M35 Project Data". Archived from the original on 2015-09-24. Retrieved 2012-04-01.
↑ "E-M35 Project Data". Archived from the original on 2015-09-24. Retrieved 2012-04-01.
↑ "E-M35 Project Data". Archived from the original on 2015-09-24. Retrieved 2012-04-01.
↑ "E-M35 Project Data". Archived from the original on 2015-09-24. Retrieved 2012-04-01.
↑ "E-M35 Project Data". Archived from the original on 2015-09-24. Retrieved 2013-01-14.
↑ ISOGG (2011)
↑ Karafet et al. (2008)
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Sources for conversion tables

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[vanOven2014-77] 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.

[ISOGG2015-78] International Society of Genetic Genealogy (ISOGG; 2015), Y-DNA Haplogroup Tree 2015. (Access date: 1 February 2015.)

[A0-T-79] Haplogroup A0-T is also known as A-L1085 (and previously as A0'1'2'3'4).

[A1-80] Haplogroup A1 is also known as A1'2'3'4.

[F-Y27277-81] F-Y27277, sometimes known as F2'4, is both the parent clade of F2 and F4 and a child of F-M89.

[LT-82] Haplogroup LT (L298/P326) is also known as Haplogroup K1.

[K2-83] 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.

[K2b-84] Haplogroup K2b (M1221/P331/PF5911) is also known as Haplogroup MPS.

[K2b1-85] Haplogroup K2b1 (P397/P399) is also known as Haplogroup MS, but has a broader and more complex internal structure.

[P-86] Haplogroup P (P295) is also klnown as K2b2.

[K-M2313-87] 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)

[M-88] Haplogroup M, as of 2017, is also known as K2b1b. (Previously the name Haplogroup M was assigned to K2b1d.)

[S-89] Haplogroup S, as of 2017, is also known as K2b1a. (Previously the name Haplogroup S was assigned to K2b1a4.)

[4] Semino et al. (2004): "Both phylogeography and microsatellite variance suggest that E-P2 and its derivative, E-M35, probably originated in eastern Africa."

[12] "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)

[32] "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)

[38] 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".

[39] 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"

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[36]

[37]

[38]

[39]

[40]

[41]

[42]

[43]

[44]

[45]

[46]

[47]

[48]

[49]

[50]

[51]

[52]

[53]

[54]

[55]

[56]

[57]

[58]

[59]

[60]

[61]

[62]

[63]

[64]

[65]

[Note 6]

[6]

[66]

[67]

[68]

[69]

[70]

[χ 1]

[χ 2]

[χ 3]

[χ 4]

[χ 5]

[χ 6]

[χ 7]

[χ 8]

[χ 9]

[χ 10]

[χ 11]

[χ 12]

[χ 13]