E   M40/SRY4064/SRY8299, M96, P29, P150, P152, P154, P155, P156, P162, P168, P169, P170, P171, P172, P173, P174, P175, P176
�       E*   -
�       E1   P147
�      �       E1*   -
�      �       E1a   M33, M132
�      �       �       E1a*   -
�      �       �       E1a1   M44
�      �       �       E1a2   P110
�      �       E1b   P177
�      �       �       E1b*   -
�      �       �       E1b1   DYS391p, P2/PN2, P179, P180, P181
�      �       �      �       E1b1*   -
�      �       �      �       E1b1a   DYS271/M2/SY81, M180/P88, P1/PN1, P46, P182, P189, P211, P293
�      �       �      �       �       E1b1a*   -
�      �       �      �       �       E1b1a1   M58
�      �       �      �       �       E1b1a2   M116.2
�      �       �      �       �       E1b1a3   M149
�      �       �      �       �       E1b1a4   M154
�      �       �      �       �       E1b1a5   M155
�      �       �      �       �       E1b1a6   M10, M66, M156, M195
�      �       �      �       �       E1b1a7   M191/P86, U186, P253/U247
�      �       �      �       �      �       E1b1a7*   -
�      �       �      �       �      �       E1b1a7a   P252/U174
�      �       �      �       �      �       �       E1b1a7a*   -
�      �       �      �       �      �       �       E1b1a7a1   P9.2
�      �       �      �       �      �       �       E1b1a7a2   P115
�      �       �      �       �      �       �       E1b1a7a3   P116
�      �       �      �       �      �       �      �       E1b1a7a3*   -
�      �       �      �       �      �       �      �       E1b1a7a3a    P113
�      �       �      �       �       E1b1a8   U175
�      �       �      �       �      �       E1b1a8*   -
�      �       �      �       �      �       E1b1a8a    P277, P278, U209
�      �       �      �       �      �       �       E1b1a8a*   -
�      �       �      �       �      �       �       E1b1a8a1   U290
�      �       �      �       �      �       �      �       E1b1a8a1*   -
�      �       �      �       �      �       �      �       E1b1a8a1a   U181
�      �       �      �       �      �       �       E1b1a8a2   P59
�      �       �      �       �       E1b1a9   P268, P269
�      �       �      �       E1b1b   M215
�      �       �      �       �       E1b1b*   -
�      �       �      �       �       E1b1b1   M35
�      �       �      �       �      �       E1b1b1*   -
�      �       �      �       �      �       E1b1b1a   L18, M78, V68   (position relative to V65 is uncertain)
�      �       �      �       �      �       �       E1b1b1a*   -
�      �       �      �       �      �       �       E1b1b1a1   V12
�      �       �      �       �      �       �      �       E1b1b1a1*   -
�      �       �      �       �      �       �      �       E1b1b1a1a   M224
�      �       �      �       �      �       �      �       E1b1b1a1b   V32
�      �       �      �       �      �       �       E1b1b1a2   V13, V36
�      �       �      �       �      �       �      �       E1b1b1a2*   -
�      �       �      �       �      �       �      �       E1b1b1a2a   V27
�      �       �      �       �      �       �      �       E1b1b1a2b   P65
�      �       �      �       �      �       �      �       E1b1b1a2c   L17
�      �       �      �       �      �       �       E1b1b1a3   V22
�      �       �      �       �      �       �      �       E1b1b1a3*   -
�      �       �      �       �      �       �      �       E1b1b1a3a   M148
�      �       �      �       �      �       �      �       E1b1b1a3b   V19
�      �       �      �       �      �       �       E1b1b1a4   V65
�      �       �      �       �      �       �       E1b1b1a5   M521
�      �       �      �       �      �       E1b1b1b   M81
�      �       �      �       �      �       �       E1b1b1b*   -
�      �       �      �       �      �       �       E1b1b1b1   M107
�      �       �      �       �      �       �       E1b1b1b2   M183   (formerly E1b1b1b)
�      �       �      �       �      �       �      �       E1b1b1b2a    M165
�      �       �      �       �      �       �      �       E1b1b1b2b    M243   (Position relative to M165 is uncertain)
�      �       �      �       �      �       �      �       E1b1b1b2c    M310   (Position relative to M165 is uncertain)
�      �       �      �       �      �       �      �       E1b1b1b2d    L19   (Position relative to M165 is uncertain)
�      �       �      �       �      �       E1b1b1c   M123
�      �       �      �       �      �       �       E1b1b1c*   -
�      �       �      �       �      �       �       E1b1b1c1   M34
�      �       �      �       �      �       �      �       E1b1b1c1*   -
�      �       �      �       �      �       �      �       E1b1b1c1a   M84
�      �       �      �       �      �       �      �       �       E1b1b1c1a1   M136   (formerly E1b1b1c1a)
�      �       �      �       �      �       �      �       E1b1b1c1b   M290
�      �       �      �       �      �       E1b1b1d   M281
�      �       �      �       �      �       E1b1b1e   V6
�      �       �      �       �      �       E1b1b1f   P72
�      �       �      �       �      �       E1b1b1g   M293
�      �       �      �       E1b1c   M329
�      �       �       E1b2   P75
�       E2   M75, P68
�      �       E2*   -
�      �       E2a   M41/P210
�      �       E2b   M54, M90, M98
�      �       �       E2b*   -
�      �       �       E2b1   M85
�      �       �      �       E2b1*   -
�      �       �      �       E2b1a   M200
�      �       �      �       �       E2b1a*    -
�      �       �      �       �       E2b1a1   P45
�      �       �      �       �       E2b1a2   P258

NOTES:

• Identical SNPs that were discovered separately are listed in alphabetical order, not necessarily in the order of discovery, and separated by "/". Examples: M40/SRY4064/SRY8299, P2/PN2, DYS271/M2/SY81, M180/P88, P1/PN1, M191/P86, P253/U247, P252/U174 and M41/P210.

Y-DNA haplogroup E would appear to have arisen in Northeast Africa based on 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.E1b1 is by far the lineage of greatest geographical distribution. It has two important sub-lineages, E1b1a and E1b1b. E1b1a is an African lineage that probably expanded from northern African to sub-Saharan and equatorial Africa with the Bantu agricultural expansion. E1b1a is the most common lineage among African Americans. E1b1b1 probably evolved either in Northeast Africa or the Near East and then expanded to the west--both north and south of the Mediterranean Sea. Eb1b1 clusters are seen today in Western Europe, Southeast Europe, 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, but note that the Cruciani haplogroup labels are now superseded because of the recently discovered new SNPS that lie closer to the root of the E branch of the Y-haplogroup Tree.

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.

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.
Battaglia et al, Y-chromosomal Evidence of the Cultural Diffusion of Agriculture in Southeast Europe European Journal of Human Genetics, 249. 2008.
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.
Cruciani et al, Recurrent Mutation in SNPs within Y chromosome E3b (E-M215) Haplogroup: A Rebuttal. (abstract) American Journal of Human Biolology. Sep-Oct;20(5):614-6, 2008
. 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. (pdf) European Journal of Human Genetics, 12:855-863, 2004.
Henn et al, Y-chromosomal Evidence of a Pastoralist Migration through Tanzania to Southern Africa. PNAS, vol. 105 no. 31 10693-10698, 2008.
Karafet et al, New Binary Polymorphisms Reshape and Increase Resolution of the Human Y-Chromosomal Haplogroup Tree. Abstract. Genome Research, published online April 2, 2008. Supplementary Material.
King et al, Differential Y-chromosome Anatolian Influences on the Greek and Cretan Neolithic. (abstract) Annals of Human Genetics. 72:205�214. 2008.
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:

Bill Harvey, Denis Savard, Victor Villarreal, FTDNA's E-M35 Project (formerly the E3b Project)
Victor Villareal, The E-M35 Phylogeny Project
E-M35 community, Double Helix Forums
Dennis Garvey, Discussion on E3a (No longer mantained)
Dennis Garvey, Discussion on E3b (No longer mantained)

Corrections/Additions made since 31 December 2007:

• Added Karafet et al (2008) on 4 May 2008.
• SNPs added on 4 May 2008: P9.2, P59, P65, P68, P72, P88, P110, P113, P115, P116, P147, P150, P152, P154, P155, P156, P162, P168, P169, P170, P171, P172, P173, P174, P175, P176, P177, P179, P180, P181, P182, P189, P211, P252, P253, P258, P268, P269, P277, P278, P293.
• SNP M183 moved from E1b1b1b2 to E1b1b1b on 5 May 2008.
• Added Victor Villarreal as a contact expert on 26 September 2008.
• Removed David Wilson as a contact expert on 29 September 2008.
• Changed subclades for M183, M165, M136 and added M293 on 23 October 2008.
• Added Henn et al on 23 October 2008.
• Added L18, M243 and M310 on 21 November 2008.
• Added L17 and L19 on 24 November 2008.
• Additional Resources list updated on 10 December 2008.
• Added M521, V68, Battaglia et al (2008) and Cruciani et al (2008) on 31 December 2008.

Contact Person for Haplogroup E: Victor Villarreal

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