Haplogroup E-V38

From Wikipedia, the free encyclo28 (1): 97–97. doi:10.1002/humu.9469. PMID17154278.  
Luis, J.R.; D.J.Rowold (March 2004). “The Levant versus the Horn of Africa: Evidence for Bidirectional Corridors of Human Migrations”. The American Society of Human Genetics 74 (3): 532–544. doi:10.1086/382286. PMC 1182266. PMID 14973781. Retrieved 2010-09-20

Published on Mar 2, 2013

Spencer Wells talks to a news team in Tucson, Arizona about their genealogy results. Haplogroup J, E, R1b (are discussed) mtDNA N descendants –



mtDNA results come from historical persons whose mitochondrial DNA has been tested; it identifies direct maternal ancestry, which is just one line out of many. ]

plogroup E-V38/ E3a/ E1b1a
Possible time of origin approx 25,000-30,000 years BP[1]
Possible place of origin West Africa or East Africa[2]
Ancestor E-P2
Descendants E-M2, E-M329
Defining mutations L222.1, V38, V100

In human genetics, Haplogroup E-V38 is a human Y-chromosome DNA haplogroup. It is the phylogenetic term for the series of unique sequence variants on the human Y-chromosome. It is often found in African males and their descendants and is heritably passed in lineage from father to son. Geneticists study these variants in populations to find the evolutionary lineage to a common male human ancestor. It can also be referred to in phylogenetic nomenclature by names such as E1b1a (although the exact definition of phylogenetic names can vary over time).

E-V38 has two basal branches, E-M329 and E-M2, the former is almost exclusively found in Ethiopia. The E-M2 branches are the predominant lineage in Western Africa, Central Africa, Southern Africa, and the southern parts of Eastern Africa. E-M2 has several subclades, however many members are included in either E-L485 or E-U175.


The discovery of two SNPs (V38 and V100) by Trombetta et al. (2011) significantly redefined the E-V38 phylogenetic tree. This led the authors to suggest that E-V38 may have originated in East Africa. V38 joins the West African-affiliated E-M2 and the northern East African-affiliated E-M329 with an earlier common ancestor who, like E-P2, may have also originated in East Africa.[2] It is possible that soon after the evolution of E-V38, trans-Saharan migrants carried the E-V38 marker to Northern Africa and Central Africa and/or West Africa where the more common E-M2 marker later arose and became prolific within the last 20,000-30,000 years.[1][3]

The downstreams SNP E-M180 possibly originated on the moist south-central Saharan savannah/grassland of northern West Africa during the early Holocene period. Much of the population that carried E-M2 retreated to southern West Africa with the drying of the Sahara. These later people migrated from Southeastern Nigeria and Cameroon ~8.0 kya to Central Africa, East Africa, and Southern Africa causing or following the Bantu expansion.[4][5][6] According to Wood et al. (2005) and Rosa et al. (2007), such population movements from West Africa changed the pre-existing population Y chromosomal diversity in Western, Central, Southern and southern East Africa, replacing the previous haplogroups frequencies in these areas with the now dominant E1b1a1 lineages. Traces of earlier inhabitants, however, can be observed today in these regions via the presence of the Y DNA haplogroups A1a, A1b, A2, A3, and B-M60 that are common in certain populations, such as the Mbuti and Khoisan.[1][7][8]


E-V38 reaches frequencies of over 80% in many parts of West Africa, Central Africa, East Africa as well as Southern Africa.[9] This haplogroup’s frequency and diversity are highest in the West Africa region. Within Africa, E-V38 displays a west-to-east as well as a south-to-north clinal distribution. In other words, the frequency of the haplogroup decreases as one moves from western and southern Africa toward the eastern and northern parts of the continent.[10]

Incidence of E-V38
Population group frequency References
Bamileke 96%-100% [10][11]
Ewe 97% [7]
Ga 97% [7]
Yoruba 93.1% [12]
Tutsi 85% [10]
Fante 84% [7]
Mandinka 79%-87% [1][7]
Ovambo 82% [7]
Senegalese 81% [13]
Ganda 77% [7]
Bijagós 76% [1]
Balanta 73% [1]
Fula 73% [1]
Herero 71% [7]
Nalú 71% [1]

Populations on the northern fringes of West Africa, central Eastern Africa and Madagascar have tested at more moderate frequencies.

Incidence of E-V38
Population group frequency References
Tuareg from Tânout, Niger 44.4% (8/18 subjects) [14]
Tuareg from Gorom, Burkina Faso 16.6% (3/18) [14]
Tuareg from Gossi, Mali 9.1% (1/9) [14]
Cape Verdeans 15.9% (32/201) [15]
Maasai 15.4% (4/26) [7]
Luo 66% (6/9) [7]
Iraqw 11.11% (1/9) [7]
Comoros 23.46% (69/294) [16]
Merina people (also called Highlanders) 44% (4/9) [17]
Antandroy 69.6% (32/46) [17]
Antanosy 48.9% (23/47) [17]
Antaisaka 37.5% (3/8) [17]

E-V38 is found at low to moderate frequencies in North Africa, and northern East Africa. The some of the lineages found in these areas are possibly due to the Bantu expansion or other migrations.[10][18] The E-M2 marker that appeared in North African samples may stem from recent acquisitions.[10] However, the discovery in 2011 of the E-V38 marker that predates E-M2 has led Trombetta et al. to suggest that E-V38 may have originated in East Africa (please refer to the Origins section for the details).

Incidence of E-V38
Population group frequency References
Tuareg from Al Awaynat and Tahala, Libya 46.5% (20/43) [Note 1] [19]
Oran, Algeria 8.6% (8/93) [20]
Berbers, southern and north-central Morocco 9.5% (6/63) [21][Note 2]
MoroccanArabs 6.8% (3/44) [21]
Saharawis 3.5% (1/29) [21]
Egyptians 8.33% (3/36), 1.4% (2/147), and (0/73) [10][22][23]
Tunisians 1.4% (2/148) [23]
SudaneseHausa 12.5% (4/32) [24]
Somalis 1.5% (3/201) [18]
Ethiopians 3.4% (3/88) [25]
Oromo 2.6% (2/78) [13]
Amhara 0% (0/48)[Note 3] [13]

Outside of Africa, E-V38 has been found at low frequencies. In Eurasia, the clade has primarily been found in West Asia. There have also been reported a few isolated incidents of E-V38 in Southern European populations in Croatia, Malta, Spain and Portugal.[26][27][28][29]

Incidence of E-V38 in Eurasia
Population group frequency References
Saudi Arabians 7.6% (12/157) [30]
Omanis 6.6% (8/121) [10]
Emiratis 5.5% (9/164) [31]
Yemenis 4.8% (3/62) [31]
Majorcans 3.2% (2/62) [29]
Qataris 4.2% (3/72) [31]
Southern Iranians 1.7% (2/117) [32]
Iraqis 1.4% (2/139) [33]
Pakistanis 1.4% (9/638) [34]
Istanbul, Turkey 1.2% (1/81) [35]

The Trans-Atlantic slave trade brought people to North America, Central America and South America including the Caribbean. Consequently, the haplogroup is often observed in the United States populations in men who self-identify as African Americans.[9] It has also been observed in a number of populations in Mexico, the Caribbean, Central America, and South America among people of African descent.

Incidence of E-V38 in populations of the Americas
Population group frequency References
U.S. Americans 7.7-7.9% [Note 4] [9]
Cubans 9.8% (13/132) [36]
Dominicans 7.69% (2/26) [37]
Puerto Ricans 19.23% (5/26) [37]
Nicaraguans 5.5% (9/165) [38]
Several populations of Colombians 6.18% (69/1116) [39]
Alagoas, Brazil 4.45% (11/247) [40]
Bahia, Brazil 19% (19/100) [41]



African spatial distribution of haplogroup E3a-M2. Rosa et al. (2007)

E1b1a1 is defined by markers DYS271/M2/SY81, M291, P1/PN1, P189, P293, V43, and V95.


E1b1a1a1 is commonly defined by M180/P88. The basal subclade is quite regularly observed in V38+ samples.


E1b1a1a1a is defined by marker M58. 5% (2/37) of the town Singa-Rimaïbé, Burkina Faso tested positive for E-M58.[11] 15% (10/69) of Hutus in Rwanda tested positive for M58.[10] Three South Africans tested positive for this marker.[8] One Carioca from Rio de Janeiro, Brazil tested positive for the M58 SNP.[42] The place of origin and age is unreported.


E1b1a1a1b is defined by M116.2, a private marker. A single carrier was found in Mali.[8][Note 5]


E1b1a1a1c is defined by private marker M149. This marker was found in a single South African.[8]


E1b1a1a1d is defined by a private marker M155. It is known from a single carrier in Mali.[8]


E1b1a1a1e is defined by markers M10, M66, M156 and M195. Wairak people in Tanzania tested 4.6% (2/43) positive for E-M10.[10] E-M10 was found in a single person of the Lissongo group in the Central African Republic and two members in a “Mixed” population from the Adamawa region.[8]


E1b1a1a1f is defined by L485. The basal node E-L485* appears to be somewhat uncommon but has not been sufficiently tested in large populations. The ancestral L485 SNP (along with several of its subclades) was very recently discovered. Some of these SNPs have little or no published population data and/or have yet to receive nomenclature recognition by the YCC.

  • E1b1a1a1f1 is defined by marker L514. This SNP is currently without population study data outside of the 1000 Genomes Project.
  • E1b1a1a1f1a (YCC E1b1a7) is defined by marker M191/P86. Filippo et al. (2011) studied a number of African populations that were E-M2 positive and found the basal E-M191/P86 (without E-P252/U174) in a population of Gur speakers in Burkina Faso.[43] Montano et al. (2011) found similar sparse distribution of E-M191* in Nigeria, Gabon, Cameroon and Congo.[6] M191/P86 positive samples occurred in tested populations of Annang (38.3%), Ibibio (45.6%), Efik (45%), and Igbo (54.3%) living in Nigeria, West Africa.[44] E-M191/P86 appears in varying frequencies in Central and Southern Africa but almost all are also positive for P252/U174. Bantu-speaking South Africans (89/343) tested 25.9% positive and Khoe-San speaking South Africans tested 7.7% (14/183) positive for this SNP.[45] It also appears commonly in Africans living in the Americas. A population in Rio de Janeiro, Brazil tested 9.2% (12/130) positive.[42] 34.9% (29/83) of American Haplogroup E men tested positive for M191.[9]
Veeramah et al. (2010) studies of the recombining portions of M191 positive Y chromosomes suggest that this lineage has “diffusely spread with multiple high frequency haplotypes implying a longer evolutionary period since this haplogroup arose”.[44] The subclade E1b1a1a1f1a appears to express opposite clinal distributions to E1b1a1* in the West African Savanna region. Haplogroup E1b1a1a1f1a (E-M191) has a frequency of 23% in Cameroon (where it represents 42% of haplotypes carrying the DYS271 mutation or E-M2), 13% in Burkina Faso (16% of haplotypes carrying the M2/DYS271 mutation) and only 1% in Senegal.[13] Similarly, while E1b1a reaches its highest frequency of 81% in Senegal, only 1 of the 139 Senegalese that were tested showed M191/P86.[13] In other words, as one moves to West Africa from western Central Africa, the less subclade E1b1a1f is found. “A possible explanation might be that haplotype 24 chromosomes [E-M2*] were already present across the Sudanese belt when the M191 mutation, which defines haplotype 22, arose in central western Africa. Only then would a later demic expansion have brought haplotype 22 chromosomes from central western to western Africa, giving rise to the opposite clinal distributions of haplotypes 22 and 24.”[11]
  • E1b1a1a1f1a1 (YCC E1b1a7a) is defined by P252/U174. It appears to be the most common subclade of E-L485. It is believed to have originated near western Central Africa. It is rarely found in the most western portions of West Africa. Montano et al. (2011) found this subclade very prevalent in Nigeria and Gabon.[6] Filippo et al. (2011) estimated a tMRCA of ~4.2 kya from sample of Yoruba population positive for the SNP.[43]
  • E1b1a1a1f1a1b (YCC E1b1a7a2) is defined by P115. This subclade has only been observed amongst Fang people of Central Africa.[6]
  • E1b1a1a1f1a1c (YCC E1b1a7a3) is defined by P116. Montano et al. (2011) observed this SNP only in Gabon and a Bassa population from Cameroon.[6]
  • E1b1a1a1f1a1d is defined by Z1704. This subclade has been observed across Africa. The 1000 Genomes Project Consortium found this SNP in Yoruba Nigerian, three Kenyan Luhyas and one African descent Puerto Rican.[46]
  • E1b1a1a1f1b is defined by markers L515, L516, L517, and M263.2. This subclade was found by the researchers of Y-Chromosome Genome Comparison Project using data from the commercial bioinformatics company 23andme.[47]


E1b1a1a1g (YCC E1b1a8) is defined by marker U175. The basal E-U175* is extremely rare. Montano et al. (2011) only found one out of 505 tested African subjects who was U175 positive but negative for U209.[6] Brucato et al. found similarly low frequencies of basal E-U175* in subjects in the Ivory Coast and Benin. Veeramah et al. (2010) found U175 in tested Annang (45.3%), Ibibio (37%), Efik (33.3%), and Igbo (25.3%) but did not test for U209.[44]

The supposed “Bantu haplotype” found in E-U175 carriers is “present at appreciable frequencies in other Niger–Congo languages speaking peoples as far west as Guinea-Bissau“.[44] This is the modal haplotype of STR markers that is common in carriers of E-U175.[Note 6]

E-U175 haplotype DYS19 DYS388 DYS390 DYS391 DYS392 DYS393
15 12 21 10 11 13

E1b1a1a1g has several subclades.

  • E1b1a1a1g1 (YCC E1b1a8a) is defined by U209. It is the most prominent subclade of U175. This subclade has very high frequencies of over fifty percentages in Cameroonian populations of Bassa and Bakaka, possibly indicating place of origin. However, E-U209 is widely found at lower frequencies in West and Central African countries surrounding Cameroon and Gabon.[6] Brucato et al. (2010) found the SNP in a populations of Ahizi (in Ivory Coast) 38.8% (19/49), Yacouba (Ivory Coast) 27.5% (11/40), and Beninese 6.5% (5/77) respectively.[48]
  • E1b1a1a1g1a (YCC E1b1a8a1) is defined by U290. The Montano et al. (2011) study of U290 showed a lower frequency in Nigeria (11.7%) and western Central Africa than basal node U209. The highest population frequency rate in that study was 57.7% (13/26) in Ewondo in Cameroon.[6] 32.5% (27/83) of American Haplogroup E men tested by Sims et al. (2007) were positive for this SNP.[9]
  • E1b1a1a1g1a2 is defined by Z1725. This marker has been observed by The 1000 Genomes Project Consortium in Yoruba Nigerians and Luhya Kenyans.[46]
  • E1b1a1a1g1c (YCC E1b1a4) is defined by M154. A Bamilike population tested 31.3% (15/48) for the marker. Bakaka speakers from Cameroon tested 8%.[11] An Ovimbundu test population found this SNP at 14% (14/100).[49] Members of this subclade have also been found in South Africa.[8][45]
  • E1b1a1a1g1d is defined by V39. Trombetta et al. first published this SNP in 2011 but gave little population data about it.[2] It is only known to have been found in an African population.


E1b1a1a1h is defined by markers P268 and P269. It was first reported in a person from the Gambia.[50]


E1b1a2 is defined by the SNP mutation M329.[Note 7] The majority of the few cases so far observed have been found in East Africa. Semino et al. (2004) found 2 Ethiopian Oromo in a study of >2400 individuals, including 78 Oromo.[51] Cadenas et al. (2007) found 1 case in Qatar, out of 72 people tested there in that study.[31]


Phylogenetic history[edit]

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[edit]

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


This phylogenetic tree of haplogroup subclades is based on the Y-Chromosome Consortium (YCC) 2008 Tree,[50] the ISOGG Y-DNA Haplogroup E Tree,[4] and subsequent published research.

  • E1b1a (L222.1, V38, V100)
    • E1b1a1 (DYS271/M2/SY81, M291, P1/PN1, P189, P293, V43, V95, Z1101, Z1107, Z1116, Z1120, Z1122, Z1123, Z1124, Z1125, Z1127, Z1130, Z1133) [Note 8]
      • E1b1a1a (L576)
        • E1b1a1a1 (L86.1, L88.3, M180/P88, PAGES00066, P182, Z1111, Z1112)
          • E1b1a1a1a (M58, PAGES00027)
          • E1b1a1a1b (M116.2)
          • E1b1a1a1c (M149)
          • E1b1a1a1d (M155)
          • E1b1a1a1e (M10, M66, M156, M195)
          • E1b1a1a1f (L485)
            • E1b1a1a1f1 (L514)
              • E1b1a1a1f1a (M191/P86, P253/U247, U186, Z1712, rs9786041)
                • E1b1a1a1f1a1 (P252/U174)
                  • E1b1a1a1f1a1a (P9.2)
                  • E1b1a1a1f1a1b (P115)
                  • E1b1a1a1f1a1c (P116)
                    • E1b1a1a1f1a1c1 (P113)
                  • E1b1a1a1f1a1d (Z1704)
                  • (L372)
              • E1b1a1a1f1b (L515, L516, L517, M263.2)
                • E1b1a1a1f1b1 (Z1893)
                  • (Z1894)
          • E1b1a1a1g (U175)
            • E1b1a1a1g1 (L220.3, L652, P277, P278.1, U209, rs7067329, rs7474403, rs7893016)
              • E1b1a1a1g1a (U290)
                • E1b1a1a1g1a1 (U181)
                  • E1b1a1a1g1a1a (L97)
                • E1b1a1a1g1a2 (Z1725)
                • (L649, L650, L651)
              • E1b1a1a1g1b (P59)
              • E1b1a1a1g1c (M154)
              • E1b1a1a1g1d (V39)
              • (L609, L611)
          • E1b1a1a1h (P268, P269)
    • E1b1a2 (M329)
Evolutionary tree of human Y-chromosome DNA (Y-DNA) haplogroups[n 1][n 2]
Y-chromosomal Adam
A00 A0-T [n 3]
A0 A1[n 4]
A1* A1a A1b
A1b* A1b1 BT
I J LT (K1) K2
L T MPS (K2b) NO (K2a) K2c K2d K2e[n 5]
  1. Jump up ^ 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. Jump up ^ International Society of Genetic Genealogy (ISOGG; 2015), Y-DNA Haplogroup Tree 2015. (Access date: 1 February 2015.)
  3. Jump up ^ Haplogroup A0-T is also known as A0’1’2’3’4.
  4. Jump up ^ Haplogroup A1 is also known as A1’2’3’4.
  5. Jump up ^ Haplogroup K2e (K-M147) was previously known as ex-K2a and/or “Haplogroup X”.

See also[edit]

Ancient Genetics[edit]

According to a genetic study in December 2012, Egyptian Monarch Ramesses III belonged to Y-DNA haplogroup E1b1a, a YDNA haplogroup mainly found in Africa with a possible source of origin in East Africa.[52]


Y-DNA E subclades[edit]

Y-DNA backbone tree[edit]

Evolutionary tree of human Y-chromosome DNA (Y-DNA) haplogroups[n 1][n 2]
Y-chromosomal Adam
A00 A0-T [n 3]
A0 A1[n 4]
A1* A1a A1b
A1b* A1b1 BT
I J LT (K1) K2
L T MPS (K2b) NO (K2a) K2c K2d K2e[n 5]
  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 K2e (K-M147) was previously known as ex-K2a and/or “Haplogroup X”.


  1. All were positive for U175.
  2. The publication refers to E-V38 as H22.
  3. Five of the 48 Amhara tested positive for P2 but were negative for M215,M2 and M329. They were not tested for V38 or L222.
  4. E-V38 is approximately 7.7-7.9% of total US male population.
  5. he publication transposes M116.2 with M116.1 in Table 1.
  6. The YCAII STR marker value of 19-19 is also usually indicative of U175.
  7. It was formerly known as E1b1c.
  8. YS271/M2/SY81, P1/PN1, P189, P293, and M291 appear to form E1b1a1*. L576 forms a subclade immediately after the previously mentioned SNPs. L576 gave rise to a deeper subclade of M180/P88, P182, L88.3, L86, and PAGES0006. From this subclade, all the major subclades (i.e. E-U175 and E-L485) of E1b1a evolved. The exact position of V43 and V95 within these three subclades and E1b1a1a1b (M116.2), E1b1a1a1c (M149), and E1b1a1a1d (M155) remains uncertain.


  1. , Alexandra; Carolina Ornelas; Mark A Jobling; António Brehm; Richard Villems (27 July 2007). “Y-chromosomal diversity in the population of Guinea-Bissau: a multiethnic perspective”. BMC Evolutionary Biology 7: 124. doi:10.1186/1471-2148-7-124. PMC 1976131. PMID 17662131. 
  2. Tmbetta, Beniamino; Fulvio Cruciani; Daniele Sellitto; Rosaria Scozzari (6 January 2011). MacAulay, Vincent, ed. “A New Topology of the Human Y Chromosome Haplogroup E1b1 (E-P2) Revealed through the Use of Newly Characterized Binary Polymorphisms”. PLoS ONE 6 (1): e16073. doi:10.1371/journal.pone.0016073. PMC 3017091. PMID 21253605. Retrieved 7 January 2010. 
  3. Semino, Ornella; Chiara Magri, Giorgia Benuzzi, Alice A. Lin, Nadia Al-Zahery, Vincenza Battaglia, Liliana Maccioni, Costas Triantaphyllidis, Peidong Shen, Peter J. Oefner, Lev A. Zhivotovsky, Roy King, Antonio Torroni, L. Luca Cavalli-Sforza, Peter A. Underhill, and A. Silvana Santachiara-Benerecetti (1 May 2004). “Origin, Diffusion, and Differentiation of Y-Chromosome Haplogroups E and J: Inferences on the Neolithization of Europe and Later Migratory Events in the Mediterranean Area”. American Journal of Human Genetics 74 (5): 1023–1034. doi:10.1086/386295. PMC 1181965. PMID 15069642. 
  4. International Society of Genetic Genealogy (3 February 2010). “Y-DNA Haplogroup E and its Subclades – 2010”. Retrieved 17 December 2010. 
  5. Jump up ^ Adams, Jonathan. “Africa During the Last 150,000 Years”. Retrieved 26 January 2011. 
  6. Montano ia; Gianmarco Ferri; Veronica Marcari; Chiara Batini; Okorie Anyaele; Giovanni Destro-Bisol; David Comas (1 July 2011). “The Bantu expansion revisited a new analysis of Y chromosome variation in Central Western Africa”. Molecular Ecology 20 (13): 2693–2708. doi:10.1111/j.1365-294X.2011.05130.x. PMID 21627702. 
  7. Wood, Elizabeth T; Daryn A Stover, Christopher Ehret, Giovanni Destro-Bisol, Gabriella Spedini, Howard McLeod, Leslie Louie, Mike Bamshad, Beverly I Strassmann, Himla Soodyall and Michael F Hammer (27 Apr 2005). “Contrasting patterns of Y chromosome and mtDNA variation in Africa: evidence for sex-biased demographic processes”. European Journal of Human Genetics 13 (7): 867–876. doi:10.1038/sj.ejhg.5201408. PMID 15856073. Retrieved 2010-10-20. 
  8. Underhill, P.A; Passarino G, Lin AA, Shen P, Mirazón Lahr M, Foley RA, Oefner PJ, Cavalli-Sforza LL. (Jan 2001). “The phylogeography of Y chromosome binary haplotypes and the origins of modern human populations”. Annals of Human Genetics 65 (1): 43–62. doi:10.1046/j.1469-1809.2001.6510043.x. PMID 11415522. 
  9. Sims, Lynn M.; Dennis Garvey; Jack Ballantyne (Jan 2007). “Sub-Populations Within the Major European and African Derived Haplogroups R1b3 and E3a Are Differentiated by Previously Phylogenetically Undefined Y-SNPs”. Human Mutation 28 (1): 97–97. doi:10.1002/humu.9469. PMID 17154278. carterhlc
  10. Luis, J.R.; D.J.Rowold (March 2004). “The Levant versus the Horn of Africa: Evidence for Bidirectional Corridors of Human Migrations”. The American Society of Human Genetics 74 (3): 532–544. doi:10.1086/382286. PMC 1182266. PMID 14973781. Retrieved 2010-09-20. 
  11. ^ Jump up to: a b c d Cruciani, Fulvio; Piero Santolamazza, Peidong Shen, Vincent Macaulay, Pedro Moral, Antonel Olckers, David Modiano, Susan Holmes, Giovanni Destro-Bisol, Valentina Coia, Douglas C. Wallace, Peter J. Oefner, Antonio Torroni, L. Luca Cavalli-Sforza, Rosaria Scozzari, and Peter A. Underhill (May 2002). “A back migration from Asia to sub-Saharan Africa is supported by high-resolution analysis of human Y-chromosome haplotypes”. American Journal of Human Genetics 70 (5): 1197–1214. doi:10.1086/340257. PMC 447595. PMID 11910562. 
  12. Jump up ^ The International Hapmap Consortium, Consortium; The International HapMap Consortium (27 October 2005). “A haplotype map of the human genome”. Nature 437 (7063): 1299–1320. doi:10.1038/nature04226. PMC 1880871. PMID 16255080. Retrieved 2010-09-22. 
  13. ^ Jump up to: a b c d e Semino O, Santachiara-Benerecetti AS, Falaschi F, Cavalli-Sforza LL, Underhill PA (January 2002). “Ethiopians and Khoisan share the deepest clades of the human Y-chromosome phylogeny”. Am. J. Hum. Genet. 70 (1): 265–268. doi:10.1086/338306. PMC 384897. PMID 11719903. 
  14. Painta, Luísa; Viktor Cerný, María Cerezo, Nuno M Silva, Martin Hájek, Alžbeta Vašíková, Martina Kujanová, Radim Brdicka and Antonio Salas (17 Mar 2010). “Linking the sub-Saharan and West Eurasian gene pools: maternal and paternal heritage of the Tuareg nomads from the African Sahel”. European Journal of Human Genetics 18 (9): 915–923. doi:10.1038/ejhg.2010.21. PMC 2987384. PMID 20234393. Retrieved 2010-09-08. 
  15.  Gonçalves, Rita; Alexandra Rosa; Ana Freitas; Ana Fernandes; Toomas Kivisild; Richard Villems; António Brehm (26 August 2003). “Y-chromosome lineages in Cabo Verde Islands witness the diverse geographic origin of its first male settlers”. Human Genetics 113 (6): 467–472. doi:10.1007/s00439-003-1007-4. PMID 12942365. 
  16.  Msaidie, Said; Axel Ducourneau, Gilles Boetsch, Guy Longepied, Kassim Papa, Claude Allibert, Ali Ahmed Yahaya, Jacques Chiaroni and Michael J Mitchell (January 2011). “Genetic diversity on the Comoros Islands shows early seafaring as major determinant of human biocultural evolution in the Western Indian Ocean”. European Journal of Human Genetics 19 (1): 89–94. doi:10.1038/ejhg.2010.128. PMC 3039498. PMID 20700146 
  17. Tofanelli, Sergio; Stefania Bertoncini; Loredana Castrì; Donata Luiselli; Francesc Calafell; Giuseppe Donati; Giorgio Paoli (17 June 2009). “On the origins and admixture of Malagasy: new evidence from high-resolution analyses of paternal and maternal lineages”. Molecular Biology and Evolution 26 (9): 2109–2124. doi:10.1093/molbev/msp120. PMID 19535740. Retrieved 2010-10-18. 
  18. ^ Jump up to: a b Sanchez, Juan J; Charlotte Hallenberg; Claus Børsting; Alexis Hernandez; Niels Morling (9 March 2005). “High frequencies of Y chromosome lineages characterized by E3b1, DYS19-11, DYS392-12 in Somali males”. European Journal of Human Genetics 13 (6): 856–866. doi:10.1038/sj.ejhg.5201390. PMID 15756297. Retrieved 2009-08-13. 
  19. Jump up ^ Ottoni, Claudio; Maarten H.D. Larmuseau; Nancy Vanderheyden; Cristina Martínez-Labarga; Giuseppina Primativo; Gianfranco Biondi; Ronny Decorte; Olga Rickards (10 Feb 2011). “Deep into the roots of the Libyan Tuareg: A genetic survey of their paternal heritage”. American Journal of Physical Anthropology 145 (1): 118–124. doi:10.1002/ajpa.21473. PMID 21312181. 
  20. Jump up ^ Robino, Carlos; F. Crobu, C. Di Gaetano, A. Bekada, S. Benhamamouch, N. Cerutti, A. Piazza, S. Inturri and C. Torre (31 August 2007). “Analysis of Y-chromosomal SNP haplogroups and STR haplotypes in an Algerian population sample” (EPUB). International Journal of Legal Medicine 122 (3): 251–255. doi:10.1007/s00414-007-0203-5. PMID 17909833.  Cite uses deprecated parameter |coauthors= (help)BOosch, Elena; Francesc Calafell; David Comas; Peter J. Oefner; Peter A. Underhill; Jaume Bertranpetit (April 2001). “High-Resolution Analysis of Human Y-Chromosome Variation Shows a Sharp Discontinuity and Limited Gene Flow between Northwestern Africa and the Iberian Peninsula”. American Journal of Human Genetics 68 (4): 1019–1029. doi:10.1086/319521. PMC 1275654. PMID 11254456. Retrieved 2010-09-22. 
  21. Jump up ^ Karafet, Tatiana M.; .L. Zegura, O. Posukh, L. Osipova, A. Bergen, J. Long, D. Goldman, W. Klitz, S. Harihara, P. de Knijff, V. Wiebe, R.C. Griffiths, A.R. Templeton and M.F. Hammer (March 1999). “Ancestral Asian source(s) of new world Y-chromosome founder haplotypes”. American Journal of Human Genetics 64 (3): 817–831. doi:10.1086/302282. PMC 1377800. PMID 10053017.  Cite uses deprecated parameter |coauthors= (help)
  22. ^ Jump up to: a b Arredi, Barbara; Estella S. Poloni; Silvia Paracchini; Tatiana Zerjal; Dahmani M. Fathallah; Mohamed Makrelouf; Vincenzo L. Pascali; Andrea Novelletto; Chris Tyler-Smith (1 August 2004). “A Predominantly Neolithic Origin for Y-Chromosomal DNA Variation in North Africa”. American Journal of Human Genetics 75 (2): 338–345. doi:10.1086/423147. PMC 1216069. PMID 15202071. 
  23.  Hassan, Hisham Y.; Peter A. Underhill; Luca L. Cavalli-Sforza; Muntaser E. Ibrahim (Nov 2008). “Y-chromosome variation among Sudanese: Restricted gene flow, concordance with language, geography, and history”. American Journal of Physical Anthropology 137 (3): 316–323. doi:10.1002/ajpa.20876. PMID 18618658. 
  24. Jump up ^ Underhill PA; Shen P; Lin AA et al. (November 2000). “Y chromosome sequence variation and the history of human populations”. Nat. Genet. 26 (3): 358–361. doi:10.1038/81685. PMID 11062480.  Cite uses deprecated parameter |author-separator= (help)
  25.  Mrsic, Gordan; Gordan Mrsic (6 March 2012). Branka Grskovic, Andro Vrdoljak, Maja Popovic, Ivica Valpotic, Simun Andelinovic, Vlastimil Stenzl, Edvard Ehler, Ludvik Urban, Gordana Lackovic, Peter Underhill, Dragan Primorac. “Croatian national reference Y-STR haplotype database”. Molecular Biology Reports 39 (7): 15. doi:10.1007/s11033-012-1610-3. PMID 22391654. 
  26. Capelli, Cristian; Redhead N, Romano V, Calì F, Lefranc G, Delague V, Megarbane A, Felice AE, Pascali VL, Neophytou PI, Poulli Z, Novelletto A, Malaspina P, Terrenato L, Berebbi A, Fellous M, Thomas MG, Goldstein DB. (13 March 2005). “Population structure in the Mediterranean basin: a Y chromosome perspective”. Annals of Human Genetics 70 (2): 207–225. doi:10.1111/j.1529-8817.2005.00224.x. PMID 16626331.Flores, Carlos; Nicole Maca-Meyer; Ana M Gonzalez; Peter J Oefner; Peidong Shen; Jose A Perez; Antonio Rojas; Jose M Larruga; Peter A Underhill (28 July 2004). “Reduced genetic structure of the Iberian peninsula revealed by Y-chromosome analysis: implications for population demography”. European Journal of Human Genetics 12 (10): 855–863. doi:10.1038/sj.ejhg.5201225. PMID 15280900. Retrieved 2010-12-29. 
  27. Adams, Susan M.; Elena Bosch, Patricia L. Balaresque, Stéphane J. Ballereau, Andrew C. Lee, Eduardo Arroyo, Ana M. López-Parra, Mercedes Aler, Marina S. Gisbert Grifo, Maria Brion, Angel Carracedo, João Lavinha, Begoña Martínez-Jarreta, Lluis Quintana-Murci, Antònia Picornell, Misericordia Ramon, Karl Skorecki, Doron M. Behar, Francesc Calafell and Mark A. Jobling (4 December 2008). “The Genetic Legacy of Religious Diversity and Intolerance: Paternal Lineages of Christians, Jews, and Muslims in the Iberian Peninsula”. The American Journal of Human Genetics 83 (6): 725–736. doi:10.1016/j.ajhg.2008.11.007. PMC 2668061. PMID 19061982.  Cite uses deprecated parameter |coauthors= (help)
  28. Jump up ^ Abu-Amero, Khaled; Ali Hellani; Ana M González; Jose M Larruga; Vicente M Cabrera; Peter A Underhill (22 September 2009). “Saudi Arabian Y-Chromosome diversity and its relationship with nearby regions”. BMC Genetics 10: 59. doi:10.1186/1471-2156-10-59. PMC 2759955. PMID 19772609. 
  29. ^ Jump up to: a b c d Cadenas, Alicia M.; Lev A Zhivotovsky, Luca L Cavalli-Sforza, Peter A Underhill and Rene J Herrera (10 October 2007). “Y-chromosome diversity characterizes the Gulf of Oman”. European Journal of Human Genetics 16 (3): 374–386. doi:10.1038/sj.ejhg.5201934. PMID 17928816. 
  30. Jump up ^ Regueiro M, Cadenas AM, Gayden T, Underhill PA, Herrera RJ (2006). “Iran: tricontinental nexus for Y-chromosome driven migration”. Hum. Hered. 61 (3): 132–143. doi:10.1159/000093774. PMID 16770078. 
  31. Jump up ^ Al-Zahery, N; O. Semino; G. Benuzzi; C. Magri; G. Passarino; A. Torroni; and A.S. Santachiara-Benerecetti (September 2003). “Y-chromosome and mtDNA polymorphisms in Iraq, a crossroad of the early human dispersal and of post-Neolithic migrations”. Molecular Phylogenetics and Evolution 28 (3): 458–472. doi:10.1016/S1055-7903(03)00039-3. PMID 12927131. 
  32. Jump up ^ Firasat Sadaf; Khaliq Shagufta; Mohyuddin Aisha et al. (2007). “Y-chromosomal evidence for a limited Greek contribution to the Pathan population of Pakistan”. European Journal of Human Genetics 15 (1): 121–126. doi:10.1038/sj.ejhg.5201726. PMC 2588664. PMID 17047675.  Cite uses deprecated parameter |author-separator= (help)
  33. Jump up ^ Cinnioğlu, Cengiz; Roy King, Toomas Kivisild, Ersi Kalfoğlu, Sevil Atasoy, Gianpiero L. Cavalleri, Anita S. Lillie, CharlesC. Roseman, Alice A. Lin and Kristina Prince (January 2004). “Excavating Y-chromosome haplotype strata in Anatolia”. Human Genetics 114 (2): 127–148. doi:10.1007/s00439-003-1031-4. PMID 14586639.  Cite uses deprecated parameter |coauthors= (help)
  34.  Mendizabal, Isabel; Karla Sandoval; Gemma Berniell-Lee; Francesc Calafell; Antonio Salas; Antonio Martínez-Fuentes; David Comas (21 July 2008). “Genetic origin, admixture, and asymmetry in maternal and paternal human lineages in Cuba”. BMC Evolutionary Biology 8: 213. doi:10.1186/1471-2148-8-213. PMC 2492877. PMID 18644108. Retrieved 2011-05-01. ^ Jump up to: a b Bryc, Katarzyna; Christopher Velez; Tatiana Karafet; Andres Moreno-Estrada; Andy Reynolds; Adam Auton; Michael Hammer; Carlos D. Bustamante; Harry Ostrer (6 May 2010). “Genome-wide patterns of population structure and admixture among Hispanic/Latino populations”. Proc Natl Acad Sci U S A 107 (Supplement 2): 8954–8961. doi:10.1073/pnas.0914618107. PMC 3024022. PMID 20445096. 
  35. Jump up ^ Nuñez, Carolina; Miriam Baeta; Cecilia Sosa; Yolanda Casalod; Jianye Ge; Bruce Budowle; Begoña Martinez-Jarreta (December 2010). “Reconstructing the population history of Nicaragua by means of mtDNA, Y-chromosome STRs, and autosomal STR marker”. American Journal of Physical Anthropology 143 (4): 591–600. doi:10.1002/ajpa.21355. PMID 20721944. 

Rojas, Winston; María Victoria Parra, Omer Campo, María Antonieta Caro, Juan Guillermo Lopera, William Arias, Constanza Duque, Andrés Naranjo, Jharley García, Candelaria Vergara, Jaime Lopera, Erick Hernandez, Ana Valencia, Yuri Caicedo, Mauricio Cuartas, Javier Gutiérrez, Sergio López, Andrés Ruiz-Linares, Gabriel Bedoya (September 2010). “Genetic make up and structure of Colombian populations by means of uniparental and biparental DNA markers”. American Journal of Physical Anthropology 143 (1): 13–20. doi:10.1002/ajpa.21270. PMID 20734436.  Cite uses deprecated parameter |coauthors= (help)

  1. Jump up ^ de Azevedo, Dalmo A.; Luiz Antonio F. da Silva, Leonor Gusmãob and Elizeu F. de Carvalho (December 2009). “Analysis of Y chromosome SNPs in Alagoas, Northeastern Brazil”. Forensic Science International: Genetics Supplement Series 2 (1): 421–422. doi:10.1016/j.fsigss.2009.08.166. Retrieved 2010-09-24.  Cite uses deprecated parameter |coauthors= (help)
  2. Jump up ^ Nascimento, Eugênio; Eneida Cerqueira; Eliana Azevedo; Vilma Freitas; Dalmo Azevedo (December 2009). “The Africa male lineages of Bahia’s people—Northeast Brazil: A preliminary SNPs study”. Forensic Science International: Genetics Supplement Series 2 2 (1): 349–350. doi:10.1016/j.fsigss.2009.07.010. Retrieved 2009-12-00. 
  3. ^ Jump up to: a b Hünmeier, Tábita; Cláudia Carvalho; Andrea Rita Marrero; Francisco Mauro Salzano; Sérgio Danilo Junho Pena; Maria Cátira Bortolini (June 2007). “Niger–Congo speaking populations and the formation of the Brazilian gene pool: mtDNA and Y-chromosome data”. American Journal of Physical Anthropology 133 (2): 854–867. doi:10.1002/ajpa.20604. PMID 17427922. 
  4. ^ Jump up to: a b de Filippo, Cesare; Chiara Barbieri, Mark Whitten, Sununguko Wata Mpoloka, Ellen Drofn Gunnarsdóttir, Koen Bostoen, Terry Nyambe, Klaus Beyer, Henning Schreiber, Peter de Knijff, Donata Luiselli, Mark Stoneking, and Brigitte Pakendorf (Mar 28, 2011). “Y-chromosomal variation in Sub-Saharan Africa: insights into the history of Niger–Congo groups”. Molecular Biology and Evolution 28 (3): 1255–1269. doi:10.1093/molbev/msq312. PMC 3561512. PMID 21109585.  Cite uses deprecated parameter |coauthors= (help)
  5. ^ Jump up to: a b c d Veeramah, Krishna R; Bruce A Connell; Naser Ansari Pour; Adam Powell; Christopher A Plaster; David Zeitlyn; Nancy R Mendell; Michael E Weale; Neil Bradman; Mark G Thomas (31 March 2010). “Little genetic differentiation as assessed by uniparental markers in the presence of substantial language variation in peoples of the Cross River region of Nigeria”. BMC Evolutionary Biology 10: 92. doi:10.1186/1471-2148-10-92. PMC 2867817. PMID 20356404. Retrieved 2010-09-16. 
  6. ^ Jump up to: a b Naidoo, Thijessen; Carina M Schlebusch; Heeran Makkan; Pareen Patel; Rajeshree Mahabeer; Johannes C Erasmus; Himla Soodyall (1 September 2010). “Development of a single base extension method to resolve Y chromosome haplogroups in sub-Saharan African populations”. Investigative Genetics 1 (6): 6. doi:10.1186/2041-2223-1-6. PMC 2988483. PMID 21092339. Retrieved 2010-09-23. 
  7. ^ Jump up to: a b Durbin, Richard M.; David L. Altshuler, Gonçalo R. Abecasis, David R. Bentley, Aravinda Chakravarti, Andrew G. Clark,Francis S. Collins, Francisco M. De La Vega, Peter Donnelly, Michael Egholm, Paul Flicek, Stacey B. Gabriel, Richard A. Gibbs, Bartha M. Knoppers, Eric S. Lander, Hans Lehrach, Elaine R. Mardis, Gil A.McVean, Debbie A. Nickerson, Leena Peltonen, Alan J. Schafer, Stephen T. Sherry, Jun Wang, Richard K. Wilson (28 October 2010). “A map of human genome variation from population-scale sequencing”. Nature 467 (7319): 1061–73. doi:10.1038/nature09534. PMC 3042601. PMID 20981092. Retrieved 14 July 2012.  Cite uses deprecated parameter |coauthors= (help)

Reynolds, David; Squecco, Adriano. “Y-Chromosome Genome Comparison”. Retrieved 1 August 2011. 

  1. Jump up ^ Brucato, Nicolas; Olivier Cassar; Laure Tonasso; Patricia Tortevoye; Florence Migot-Nabias; Sabine Plancoulaine; Evelyne Guitard; Georges Larrouy; Antoine Gessain; Jean-Michel Dugoujon (19 October 2010). “The imprint of the Slave Trade in an African American population: mitochondrial DNA, Y chromosome and HTLV-1 analysis in the Noir Marron of French Guiana”. BMC Evolutionary Biology 10: 314. doi:10.1186/1471-2148-10-314. PMC 2973943. PMID 20958967. Retrieved 20 May 2011. 
  2. Jump up ^ Brito, P.; Carvalho, M.; Gomes, V.; Melo, M.M.; Bogas, V.; Balsa, F.; Andrade, L.; Serra, A.; Lopes, V.; Gusmão, L.; Anjos, M.J.; Corte-Real, F. (December 2011). “Y-SNP analysis in an Angola population”. Forensic Science International: Genetics Supplement Series 3 (1): e369–e370. doi:10.1016/j.fsigss.2011.09.046. 
  3. ^ Jump up to: a b Karafet, Tatiana M.; Fernando L. Mendez; Monica B. Meilerman; Peter A. Underhill; Stephen L. Zegura; Michael F. Hammer (May 2008). “New binary polymorphisms reshape and increase resolution of the human Y chromosomal haplogroup tree”. Genome Research 18 (5): 830–838. doi:10.1101/gr.7172008. PMC 2336805. PMID 18385274. 
  4. Jump up ^ Semino; Magri, C; Benuzzi, G; Lin, AA; Al-Zahery, N; Battaglia, V; MacCioni, L; Triantaphyllidis, C et al. (2004). “Origin, Diffusion, and Differentiation of Y-Chromosome Haplogroups E and J: Inferences on the Neolithization of Europe and Later Migratory Events in the Mediterranean Area”. American Journal of Human Genetics 74 (5): 1023–1034. doi:10.1086/386295. PMC 1181965. PMID 15069642  CS1 maint: Explicit use of et al. (link)
  5. Jump up ^ Hawass (2012). “Revisiting the harem conspiracy and death of Ramesses III: anthropological, forensic, radiological, and genetic study”. BMJ 345: e8268. doi:10.1136/bmj.e8268. 

External links[edit]

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About Harold L Carter

Bachelor of Science, Columbia University Masters degree, Ohio State University Undergraduate National Officer, Alpha Phi Alpha Fraternity, Eastern Asst Vice President, when a student at Columbia University Profile Photograph: Mom & Me, when I was a graduate student
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