E   M40/SRY4064/SRY8299, M96, P29
�       E*   -
�       E1   M33, M132
�      �       E1*   -
�      �       E1a   M44
�       E2   M75
�      �       E2*   -
�      �       E2a   M41
�      �       E2b   M54, M90, M98
�      �       �       E2b*   -
�      �       �       E2b1   P45
�      �       �       E2b2   M85 (formerly E2b)
�      �       �      �       E2b2*   -
�      �       �      �       E2b2a   M200
�       E3   P2, DYS391p
�      �       E3*   -
�      �       E3a   DYS271/M2/SY81, M180, P1, P46
�      �       �       E3a*   -
�      �       �       E3a1   M58
�      �       �       E3a2   M116.2
�      �       �       E3a3   M149
�      �       �       E3a4   M154
�      �       �       E3a5   M155
�      �       �       E3a6   M10, M66, M156, M195
�      �       �       E3a7   M191, U186, U247
�      �       �      �       E3a7*   -
�      �       �      �       E3a7a   U174
�      �       �       E3a8   U175, U209
�      �       �      �       E3a8*   -
�      �       �      �       E3a8a   U290
�      �       �      �       �       E3a8a*   -
�      �       �      �       �       E3a8a1   U181
�      �       E3b   M215
�      �       �       E3b*   -
�      �       �       E3b1   M35 (formerly in E3b)
�      �       �      �       E3b1*   -
�      �       �      �       E3b1a   M78 (formerly E3b1)
�      �       �      �       �       E3b1a*   -
�      �       �      �       �       E3b1a1   V12
�      �       �      �       �      �       E3b1a1*   -
�      �       �      �       �      �       E3b1a1a    V32 (formerly E3b1a1b)
�      �       �      �       �       E3b1a2   V13, V36
�      �       �      �       �      �       E3b1a2*   -
�      �       �      �       �      �       E3b1a2a   V27
�      �       �      �       �      �       E3b1a2b   M224 (formerly E3b1b, E3b1a2 and E3b1a1a)
�      �       �      �       �       E3b1a3   V22
�      �       �      �       �      �       E3b1a3*   -
�      �       �      �       �      �       E3b1a3a   M148 (formerly E3b1a, then E3b1a2)
�      �       �      �       �      �       E3b1a3b   V19
�      �       �      �       �       E3b1a4    V65
�      �       �      �       E3b1b   M81, M183 (formerly E3b2)
�      �       �      �       �       E3b1b*   -
�      �       �      �       �       E3b1b1   M107 (formerly E3b2a)
�      �       �      �       �       E3b1b2    M165 (formerly E3b2b)
�      �       �      �       E3b1c   M123 (formerly E3b3)
�      �       �      �       �       E3b1c*   -
�      �       �      �       �       E3b1c1   M34 (formerly E3b3a)
�      �       �      �       �      �       E3b1c1*   -
�      �       �      �       �      �       E3b1c1a   M84, M136 (formerly E3b3a1)
�      �       �      �       �      �       E3b1c1b   M290 (formerly E3b3a2)
�      �       �      �       E3b1d   M281 (formerly E3c)
�      �       �      �       E3b1e   V6 (formerly E3e)
�      �       E3c   M329
�       E4   P75

NOTES:

• In E the designation M40/SRY4064/SRY8299 means that these three SNPs are identical but were discovered separately. The listing is alphabetical, not necessarily in the order of discovery.
• In E3a the designation DYS271/M2/SY81 means that these three SNPs are identical but were discovered separately. The listing is alphabetical, not necessarily in the order of discovery.
• M183 relocated from E3b1b2 to E3b1b1 to reflect analysis of more recent results.

Y-DNA haplogroup E probably arose in Northeast Africa, if one looks only at the concentration and variety of E subclades in that area today. But the fact that Haplogroup E is closely linked with Haplogroup D, which is not found in Africa, leaves open the possibility that E first arose in the Near or Middle East and was subsequently carried into Africa by a back migration.Today E* is found predominantly in Ethiopia. E1 and E2 are found in Northeast Africa, but surveys show E1 may actually be more prevalent in Mali than in its presumed region of origin. E4 is a minor subclade. E3 is by far the lineage of greatest geographical distribution. It has two important sub-lineages, E3a and E3b. E3a is an African lineage that probably expanded from northern Africa to sub-Saharan and equatorial Africa with the Bantu agricultural expansion. E3a is the most common lineage among African Americans. E3b probably evolved either in Northeast Africa or the Near East and then expanded to the west both north and south of the Mediterranean Sea. E3b clusters are seen today in Western Europe, the Balkans, the Near East, Northeast Africa and Northwest Africa. The Cruciani articles (references and links below) are indispensable resources for understanding the structure of this complicated haplogroup.

A caution on clade labels: Because knowledge of this branch of the Y-chromosome tree has advanced so quickly in the last few years, different clade labels can be found in current use for the same SNP-determined branch of the tree. For example, it is still common to see E3b1 and E3b2 used to distinguish between the M78 and M81 branches of the tree though greater resolution is now possible. Also, STR-based distinctions in the M78 branch at one time permitted broad distinctions of alpha, beta, gamma and delta clusters. With the new SNPs reported in the 2006 and 2007 Cruciani studies, it has become possible to see that the alpha cluster, which is widely distributed in Europe, is strongly correlated with the V13 SNP that identifies Haplogroup E3b1a2; the beta cluster is strongly correlated with the V65 SNP that identifies Haplogroup E3b1a4; the gamma cluster correlates with the V32 SNP that identifies Haplogroup E3b1a1a; and the delta cluster tends to correlate with the V22 SNP (E3b1a3) though it includes some V12 haplotypes (E3b1a1) as well.

References:

Alonso et al, The Place of the Basques in the European Y-chromosome Diversity Landscape. (available by subscription) European Journal of Human Genetics, 13:1293-1302, 2005.
Behar et al, Contrasting Patterns of Y Chromosome Variation in Ashkenazi Jewish and Host Non-Jewish European Populations. (pdf) Hum Genet 114:354-365, 2004.
Bortolini et al, Y-Chromosome Evidence for Differing Ancient Demographic Histories in the Americas. American Journal of Human Genetics, 73:524�539, (2003).
Capelli et al, Population Structure in the Mediterranean Basin: A Y Chromosome Perspective. (pdf) Annals of Human Genetics, 2005.
Cinnioglu et al, Excavating Y-chromosome Haplotype Strata in Anatolia. (pdf) Human Genetics. 114:127-148, 2004.
Cruciani et al, A Back Migration from Asia to Sub-Saharan Africa is Supported by High-Resolution Analysis of Human Y-Chromosome Haplotypes. (pdf) American Journal of Human Genetics, 70:1197-1214, 2002.
Cruciani et al, Phylogeographic Analysis of Haplogroup E3b (E-M215) Y Chromosomes Reveals Multiple Migratory Events Within and Out of Africa. (pdf) American Journal of Human Genetics, 74:1014-1022, 2004.
Cruciani et al, Molecular Dissection of the Y Chromosome Haplogroup E-M78 (E3b1a): A Posteriori Evaluation of a Microsatellite-Networked-Based Approach Through Six New Biallelic Markers. (pdf) Human Mutation: Mutation in Brief #916, 2006.
Cruciani et al, Tracing Past Human Male Movements in Northern/Eastern Africa and Western Eurasia: New Clues from Y-Chromosomal Haplogroups E-M78 and J-M12. (pdf) Molecular Biology and Evolution 24(6):1300-1311, 2007.
Deng et al, Evolution and Migration History of the Chinese Population Inferred from the Chinese Y-chromosome Evidence. (pdf) Journal of Human Genetics, 49:339-348, 2004.
Flores et al, Reduced Genetic Structure of the Iberian Peninsula Revealed by Y-chromosome Analysis: Implications for Population Demography. (available by subscription) European Journal of Human Genetics, 12:855-863, 2004.
Nasidze et al, MtDNA and Y-chromosome Variation in Kurdish Groups. (abstract) Annals of Human Genetics, 69:401-412, 2005.
Regueiro et al, Iran: Tricontinental Nexus for Y-Chromosome Driven Migration. (abstract) Human Heredity, Vol. 61, No 3, 132-143, 2006.
Semino et al, Ethiopians and Khoisan Share the Deepest Clades of the Human Y-Chromosome Phylogeny. (pdf) American Journal of Human Genetics, 70:265-268, 2002.
Semino et al, 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. (pdf) American Journal of Human Genetics, 74:1023-1034, 2004.
Sengupta et al, Polarity and Temporality of High Resolution Y-chromosome Distributions in India Identify Both Indigenous and Exogenous Expansions and Reveal Minor Genetic Influence of Central Asian Pastoralists. (pdf) American Journal of Human Genetics, 78:202-221, 2006.
Shen et al, Reconstruction of Patrilineages and Matrilineages of Samaritans and other Israeli Populations from Y-Chromosome and Mitochondrial DNA Sequence Variation. (pdf) Human Mutation, 24:248-260, 2004.
Sims et al, Sub-Populations Within the Major European and African Derived Haplogroups R1b3 and E3a Are Differentiated by Previously Phylogenetically Undefined Y-SNPs. Human Mutation: Mutation in Brief #940, Online, 2007.
Valone et al, Y SNP Typing of African-American and Caucasian Samples Using Allele-Specific Hybridization and Primer Extension. (pdf) Journal of Forensic Science, 49:4, July 2004.

Additional Resources:

- E3b Y-DNA Project
Dennis Garvey, Discussion on E3a
Dennis Garvey, Discussion on E3b
Ron Scott, E3b: Frequency Distribution of Extended Haplotypes Gathered from YSearch
Victor Villareal, E3b Project

Corrections/Additions made since 20 December 2006:

• Changed email address for content person on 8 June 2007.
• Added Ron Scott reference in Additional Resources on 8 June 2007.
• Added Bortolini et al (2003) on 8 June 2007.
• Added V36 to E3b1a2; added E3b1a4 with V65; added Cruciani et al (2007) on 17 June 2007.
• Changed last sentence in 2nd paragraph of Haplogroup description on 17 June 2007.
• Added Victor Villareal reference in Additional Resources on 1 July 2007.
• Due to new research by Family Tree DNA, the locations of SNPs V32 and M224 have been changed on 25 July 2007.
Added to notes on 17 October 2007.

Contact Person for Haplogroup E: David Wilson

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