Y-DNA Haplogroup E and its Subclades - 2012
The entire work is identified by the Version Number and date given on the Main Page.   Directions for citing the document are given at the bottom of the Main Page.
Version History     Last revision date for this specific page: 8 November 2012

Because of continuing research, the structure of the Y-DNA Haplogroup Tree changes and ISOGG does its best to keep the tree updated with the latest developments in the field. The viewer may observe other versions of the tree on the Web. Email Alice Fairhurst if the differences need clarification or if you find any broken links on this page.

LINKS:  Main Page   Y-DNA Tree Trunk   SNP Index   Papers/Presentations Cited   Glossary   Listing Criteria
CLADE/SUBCLADE SYMBOLS:  Added  Redefined 
SNP SYMBOLS:  Not on 2011 tree  Confirmed within subclade  Provisional  Private  Investigation 

E   L339, L504, L507, L511, L537, L614, L856, M40/SRY4064/SRY8299, M96, P29, P150, P152,
       P154, P155, P156, P162, P168, P169/Page54, P170, P171, P172, P173, P174, P175, P176
�    E*   -
�    E1   P147
�   �    E1*   -
�   �    E1a   L633, M33, M132
�   �    �    E1a*   -
�   �    �    E1a1   M44
�   �    �    E1a2   P110
�   �    �    E1a3   L94
�   �    �    E1a4   L133/Page74
�   �   ; E1b   P177
�   �    �    E1b*   -
�   �    �    E1b1   DYS391p, P2/PN2, P178, P179, P180, P181
�   �    �   �    E1b1*   -
�   �    �   �    E1b1a   L222.1, TSC0077541/V38, V100
�   �    �   �    �    E1b1a*   -
�   �    �   �    �    E1b1a1   DYS271/M2/SY81, KHS1479188, M291, P1/PN1, P189, P293,
                                           TSC0424987, V43, V95, Z1101, Z1107, Z1116, Z1120, Z1122,
                                           Z1123, Z1124, Z1125, Z1127, Z1130, Z1133, Z1135

�   �    �   �    �   �    E1b1a1*   -
�   �    �   �    �   �    E1b1a1a   L576/TSC0860384
�   �    �   �    �   �    �    E1b1a1a*   -
�   �    �   �    �   �    �    E1b1a1a1   L86.1, M180/P88, P182, Page66, Z1111, Z1132
�   �    �   �    �   �    �   �    E1b1a1a1*   -
�   �    �   �    �   �    �   �    E1b1a1a1a   M58, Page27
�   �    �   �    �   �    �   �    E1b1a1a1b   M116.2
�   �    �   �    �   �    �   �    E1b1a1a1c   M149
�   �    �   �    �   �    �   �    E1b1a1a1d   M155
�   �    �   �    �   �    �   �    E1b1a1a1e   M10, M66, M156, M195
�   �    �   �    �   �    �   �    E1b1a1a1f    L485
�   �    �   �    �   �    �   �    �    E1b1a1a1f*   -
�   �    �   �    �   �    �   �    �    E1b1a1a1f1   L514
�   �    �   �    �   �    �   �    �   �    E1b1a1a1f1*   -
�   �    �   �    �   �    �   �    �   �    E1b1a1a1f1a   M191/P86, U186, P253/U247, TSC1640819/Z1712.1
�   �    �   �    �   �    �   �    �   �    �    E1b1a1a1f1a*   -
�   �    �   �    �   �    �   �    �   �    �    E1b1a1a1f1a1   P252/U174
�   �    �   �    �   �    �   �    �   �    �   �    E1b1a1a1f1a1*   -
�   �    �   �    �   �    �   �    �   �    �   �    E1b1a1a1f1a1a   P9.2
�   �    �   �    �   �    �   �    �   �    �   �    E1b1a1a1f1a1b   P115
�   �    �   �    �   �    �   �    �   �    �   �    E1b1a1a1f1a1c   P116
�   �    �   �    �   �    �   �    �   �    �   �    �    E1b1a1a1f1a1c*   -
�   �     �   �    �   �    �   �    �   �    �   �    �    E1b1a1a1f1a1c1   P113
�   �    �   �    �   �    �   �    �   �    �   �    E1b1a1a1f1a1d   Z1704
�   �    �   �    �   �    �   �    �   �    E1b1a1a1f1b   L515, L516, L517, M263.2
�   �    �   �    �   �    �   �    �   �    �    E1b1a1a1f1b*   -
�   �    �   �    �   �    �   �    �   �    �    E1b1a1a1f1b1   Z1893
�   �    �   �    �   �    �   �    E1b1a1a1g   U175
�   �    �   �    �   �    �   �    �    E1b1a1a1g*   -
�   �    �   �    �   �    �   �    �    E1b1a1a1g1   P277, P278.1, U209
�   �    �   �    �   �    �   �    �   �    E1b1a1a1g1*   -
�   �    �   �    �   �    �   �    �   �    E1b1a1a1g1a   U290
�   �    �   �    �   �    �   �    �   �    �    E1b1a1a1g1a*   -
�   �    �   �    �   �    �   �    �   �    �    E1b1a1a1g1a1   U181
�   �    �   �    �   �    �   �    �   �    �   �    E1b1a1a1g1a1*   -
�   �    �   �    �   �    �   �    �   �    �   �    E1b1a1a1g1a1a   L97
�   �    �   �    �   �    �   �    �   �    �    E1b1a1a1g1a2   Z1725
�   �    �   �    �   �    �   �    �   �    E1b1a1a1g1b   P59
�   �    �   �    �   �    �   �    �   �    E1b1a1a1g1c   M154
�   �    �   �    �   �    �   �    �   �    E1b1a1a1g1d   V39
�   �    �   �    �   �    �   �    E1b1a1a1h   P268, P269
�   �    �   �    �    E1b1a2   M329
�   �    �   �    E1b1b   M215/Page40
�   �    �   �    �    E1b1b*   -
�   �    �   �    �    E1b1b1   L336, M35.1, M243
�   �    �   �    �   �    E1b1b1*   -
�   �    �   �    �   �    E1b1b1a   V68
�   �    �   �    �   �    �    E1b1b1a*   -
�   �    �   �    �   �    �    E1b1b1a1   L18, M78
�   �    �   �    �   �    �   �    E1b1b1a1*   -
�   �    �   �    �   �    �   �    E1b1b1a1a   V12/Z1216
�   �    �   �    �   �    �   �    �    E1b1b1a1a*   -
�   �    �   �    �   �    �   �    �    E1b1b1a1a1   M224
�   �    �   �    �   �    �   �    �    E1b1b1a1a2   V32
�   �    �   �    �   �    �   �    E1b1b1a1b   L142.1, L542, V13, V36,
�   �    �   �    �   �    �   �    �    E1b1b1a1b*   -
�   �    �   �    �   �    �   �    �    E1b1b1a1b1   V27
�   �    �   �    �   �    �   �    �    E1b1b1a1b2   P65_1, P65_2, P65_3
�   �    �   �    �   �    �   �    �    E1b1b1a1b3   L17
�   �    �   �    �   �    �   �    �    E1b1b1a1b4   DYS643-null/L143
�   �    �   �    �   �    �   �    �    E1b1b1a1b5   M35.2
�   �    �   �    �   �    �   �    �    E1b1b1a1b6   L241
�   �    �   �    �   �    �   �    �    E1b1b1a1b7   L250, L251, L252
�   �    �   �    �   �    �   �    �    E1b1b1a1b8   L540
�   �    �   �    �   �    �   �    E1b1b1a1c   V22
�   �    �   �    �   �    �   �    �    E1b1b1a1c*   -
�   �    �   �    �   �    �   �    �    E1b1b1a1c1   M148
�   �    �   �    �   �    �   �    �    E1b1b1a1c2   V19
�   �    �   �    �   �    �   �    E1b1b1a1d   V65
�   �    �   �    �   �    �   �    E1b1b1a1e   M521
�   �    �   �    �   �    E1b1b1b   Z827
�   �    �   �    �   �    �    E1b1b1b*   -
�   �    �   �    �   �    �    E1b1b1b1   L19/V257, L335, M310
�   �    �   �    �   �    �   �    E1b1b1b1*   -
�   �    �   �    �   �    �   �    E1b1b1b1a   M81
�   �    �   �    �   �    �   �    �    E1b1b1b1a*   -
�   �    �   �    �   �    �   �    �    E1b1b1b1a1   M107
�   �    �   �    �   �    �   �    �    E1b1b1b1a2   M183/Page33
�   �    �   �    �   �    �   �    �   �    E1b1b1b1a2*   -
�   �    �   �    �   �    �   �    �   �    E1b1b1b1a2a    M165
�   �    �   �    �   �   �    E1b1b1b2   Z830
�   �    �   �    �   �    �   �    E1b1b1b2*   -
�   �    �   �    �   �    �   �    E1b1b1b2a   M123
�   �    �   �    �   �    �   �    �    E1b1b1b2a*   -
�   �    �   �    �   �    �   �    �    E1b1b1b2a1   M34
�   �    �   �    �   �    �   �    �   �    E1b1b1b2a1*   -
�   �    �   �    �   �    �   �    �   �    E1b1b1b2a1a   M84, L29/Page47
�   �    �   �    �   �    �   �    �   �    �    E1b1b1b2a1a*   -
�   �    �   �    �   �    �   �    �   �    �    E1b1b1b2a1a1   M136
�   �    �   �    �   �    �   �    �   �    E1b1b1b2a1b   M290
�   �    �   �    �   �    �   �    �   �    E1b1b1b2a1c   V23    (V23 position relative to M84 and M290 uncertain)
�   �    �   �    �   �    �   �    �   �    E1b1b1b2a1d   L791, L792
�   �    �   �    �   �    �   �    E1b1b1b2b   M293
�   �    �   �    �   �    �   �    �    E1b1b1b2b*   -
�   �    �   �    �   �    �   �    �    E1b1b1b2b1   P72
�   �    �   �    �   �    �   �    E1b1b1b2c    V42
�   �    �   �    �   �    E1b1b1c   V6
�   �    �   �    �   �    E1b1b1d   V92
�   �    �   �    �    E1b1b2   M281, V16
�   �    �    E1b2   P75
�    E2   M75, P68
�   �    E2*   -
�   �    E2a   M41/P210
�   �    E2b   M54, M90, M98
�   �    �    E2b*   -
�   �    �    E2b1   M85
�   �    �   �    E2b1*   -
�   �    �   �    E2b1a   M200
�   �    �   �    �    E2b1a*    -
�   �    �   �    �    E2b1a1   P45
�   �    �   �    �    E2b1a2   P258

Private SNPs are being removed from the tree and placed in the following category:
Private SNPs - After having been investigated, these SNPs have not met the population distribution criteria for placement on the tree. Either too few confirmed positive testers have been found OR multiple confirmed testers were confined to either a single surname or to a small group of related males.

SNPs under Investigation - Additional testing is needed to confirm adequate positive samples and/or correct placement on the tree.

NOTES:

Y-DNA haplogroup E would appear to have arisen in northern hemisphere of Africa based on the concentration and variety of E subclades in that area today. E1b1 is by far the lineage of greatest geographical distribution. It has two major sub-lineages, E1b1a and E1b1b. E1b1a is a lineage that originated and expanded from West or Central Africa to Eastern Africa and Southern Africa. E1b1b probably evolved either in Northeast Africa or the Near East and then expanded to the west--both north and south of the Mediterranean Sea. E1b1b1 clusters are seen today in Western Europe, Southeast Europe, the Near East, Northeast Africa and Northwest Africa. E2 is possibly of Central African origin and is spread in low frequencies across the continent. 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:

Adams et al, The Genetic Legacy of Religious Diversity and Intolerance: Paternal Lineages of Christians, Jews, and Muslims in the Iberian Peninsula, American Journal of Human Genetics, 83(6): 725-36, 2008.
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).
Bosch et al, Paternal and Maternal Lineages in the Balkans Show a Homogeneous Landscape over Linguistis Barriers except for the Isolated Aromuns. Annals of Human Genetics, 70:459-87, (2006).
Capelli et al, Population Structure in the Mediterranean Basin: A Y Chromosome Perspective. (pdf) Annals of Human Genetics, 2005.
Cadenas et al, Y-chromosome Diversity Characterizes the Gulf of Oman. European Journal of Human Genetics, 16:374-386, 2008.
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. 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.
El Sibai et al, Geographical Structure of the Y-Chromosomal Genetic Landscape of the Levant: A Coastal Inland Contrast, Annals of Human Genetics, 73:568-81, 2009. (abstract)
Firasat et al, Y-chromosomal Evidence for a Limited Greek Contribution to the Pathan Population of Pakistan, European Journal Human Genetics, 15:121+126, 2007.
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.
Gomes et al, Digging Deeper into East African Human Y Chromosone Lineages, Human Genetics, 127(5):603-13, 2010.
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, The Coming of the Greeks to Provence and Corsica: Y-Chromosome Models of Archaic Greek Colonization of the Western Mediterranean, BMC Evolutionary Biology, 11: 69, 2011.
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.
Repping et al, High mutation rates have driven extensive structural polymorphism among human Y chromosomes. Nature Genetics 38, 463 - 467, 2006.
Rozen et al, Remarkably Little Variation in Proteins Encoded by the Y Chromosome's Single-Copy Genes, Implying Effective Purifying Selection. American Journal of Human Genetics. 2009 December 11; 85(6): 923-928.
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.
Trombetta et al, 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), Online, 2011.
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.
Zhao et al, Presence of Three Different Paternal Lineages among North Indians: A Study of 560 Y Chromosomes. (abstract) Annals of Human Biology, 36(1):46-59, 2009.

Additional Resources:

ISOGG Wiki - What you need to know about Genetic Genealogy.
E1a - (M33, M132) Project.
Haplogroup E1b1a (E1b1a1-M2), Roberta Estes.
The African DNA Project (E1b1a), Dr. Ana Oquendo Pabon.
Arabian E Y-DNA Project (E1b1).
E-M35 Project (formerly the E3b Project) (E1b1b1), Bill Harvey, Denis Savard, Victor Villarreal.
E-M35 Project - New SNP Tracker.
The E-M35 Phylogeny Project, Victor Villareal.
Double Helix Forums, E-M35 community.

Corrections/Additions made since 1 January 2012:

Contact Person for Haplogroup E: Aaron R. Brown.

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