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Haplogroup E-M215

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(Redirected fromHaplogroup E1b1b)

Human Y-chromosome DNA haplogroup

"E3b" redirects here. For the Pennsylvania Railroad locomotive, seePRR E3b.

Haplogroup E-M215 E1b1b
Geographic distribution of the haplogroup E1b1b
Possible time of origin	47,500—22,400 BP^[1]^[2]^[3]
Coalescence age	34,800 BP^[4]
Possible place of origin	East Africa^[5]^[1]
Ancestor	E-P2
Descendants	E-M35 E-M281
Defining mutations	M215

E-M215 orE1b1b, formerly known asE3b, is a majorhuman Y-chromosome DNA haplogroup. E-M215 has two basal branches,E-M35 and E-M281. E-M35 is primarily distributed inNorth Africa and theHorn of Africa, and occurs at moderate frequencies in theMiddle East,Europe, andSouthern Africa. E-M281 occurs at a low frequency inEthiopia.

Origins

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The origins of E-M215 were dated by Cruciani in 2007 to about 22,400 years ago inEast Africa.^[3]^{[Note 1]}

Ancient DNA

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According to Lazaridis et al. (2016),Natufian skeletal remains from the ancientLevant predominantly carried the Y-DNA haplogroup E1b1b. Of the five Natufian specimens analyzed for paternal lineages, three belonged to the E1b1b1b2(xE1b1b1b2a, E1b1b1b2b), E1b1(xE1b1a1, E1b1b1b1) and E1b1b1b2(xE1b1b1b2a, E1b1b1b2b) subclades (60%). Haplogroup E1b1b was also found at moderate frequencies among fossils from the ensuingPre-Pottery Neolithic B culture, with the E1b1b1 and E1b1b1b2(xE1b1b1b2a, E1b1b1b2b) subclades observed in two of seven PPNB specimens (~29%). The scientists suggest that the Levantine early farmers may have spread southward into East Africa, bringing along Western Eurasian andBasal Eurasian ancestral components separate from that which would arrive later in North Africa.

Additionally, haplogroup E1b1b1 has been found in anancient Egyptian mummy excavated at theAbusir el-Meleq archaeological site in Middle Egypt, which dates from a period between the lateNew Kingdom and theRoman era.^[6] Fossils at theIberomaurusian site ofIfri N'Amr Ou Moussa inMorocco, which have been dated to around 5,000 BCE, also carried haplotypes related to the E1b1b1b1a (E-M81) subclade. These ancient individuals bore an autochthonous Maghrebi genomic component that peaks among modernNorth Africans, indicating that they were ancestral to populations in the area.^[7] The E1b1b haplogroup has likewise been observed in ancientGuanche fossils excavated inGran Canaria andTenerife on theCanary Islands, which have been radiocarbon-dated to between the 7th and 11th centuries CE. The clade-bearing individuals that were analysed for paternal DNA were inhumed at the Tenerife site, with all of these specimens found to belong to the E1b1b1b1a1 or E-M183 subclade (3/3; 100%).^[8]

Loosdrecht et al. (2018) analysed genome-wide data from seven ancientIberomaurusian individuals from the Grotte des Pigeons nearTaforalt in eastern Morocco. The fossils were directly dated to between 15,100 and 13,900 calibrated yearsbefore present. The scientists found that five male specimens with sufficient nuclear DNA preservation belonged to the E1b1b1a1 (M78) subclade, with one skeleton bearing the E1b1b1a1b1 parent lineage to E-V13, another male specimen belonged to E1b1b (M215*).^[9] Martiniano et al. (2022) later reassigned all the Taforalt samples to haplogroup E-M78 and none to E-L618, the predecessor to E-V13.^[10]

Distribution

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In Africa, E-M215 is distributed in highest frequencies in theHorn of Africa andNorth Africa, specifically in the countriesSomalia andMorocco, whence it has in recent millennia expanded as far south asSouth Africa, and northwards intoWestern Asia andEurope (especially theMediterranean and theBalkans).^[11]^[12]^[13]^[14] E-M281 has been found inEthiopia.^[12]

Almost all E-M215 men are also in E-M35. In 2004, M215 was found to be older than M35 when individuals were found who have the M215 mutation, but do not have M35 mutation.^[11]In 2013, Di Cristofaro et al. (2013) found one individual in Khorasan, North-East Iran to be positive for M215 but negative for M35.^[15]

E-M215 and E-M35 are quite common amongAfroasiatic speakers. The linguistic group and carriers of E-M35 lineage have a high probability to have arisen and dispersed together from theAfroasiatic Urheimat.^[16] Amongst populations with an Afro-Asiatic speaking history, a significant proportion ofJewish male lineages are E-M35.^[17] Haplogroup E-M35, which accounts for approximately 18%^[12] to 20%^[18]^[19] ofAshkenazi and 8.6%^[20] to 30%^[12] ofSephardi Y-chromosomes, appears to be one of the major founding lineages of the Jewish population.^[21]^{[Note 2]}

E-M215 association with endurance

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Moran et al. (2004) observed that among Y-DNA (paternal) clades borne by elite endurance athletes in Ethiopia, the haplogroup E3b1 was negatively correlated with elite athletic endurance performance,^[22] whereas the haplogroupsE*,E3*,K*(xP),^[22] andJ*(xJ2) were significantly more frequent among the elite endurance athletes.^[22]

Subclades

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E-M35

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Main article:Haplogroup E-M35

Haplogroup E-M35 is a subclade of E-M215.

E-M281

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Main article:Haplogroup E-M281

Haplogroup E-M281 is a subclade of E-M215.

Phylogenetics

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Phylogenetic history

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Main article:Conversion table for Y chromosome haplogroups

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 inpopulation 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

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

αJobling & Tyler-Smith (2000) andKaladjieva (2001)
βUnderhill (2000)
γHammer (2001)
δKarafet (2001)
εSemino (2000)
ζSu (1999)
ηCapelli (2001)

Discussion

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E-M215 andE1b1b1 are the currently accepted names found in the proposals of theY Chromosome Consortium (YCC), for the clades defined by mutation M215 and M35 respectively, which can also be referred to as E-M215 and E-M35.^[23] The nomenclatureE3b (E-M215) andE3b1 (E-M35) respectively were the YCC defined names used to designate the samehaplogroups in older literature with E-M35 branching as a separatesubclade of E-M215 in 2004.^[11] Prior to 2002 these haplogroups were not designated in a consistent way, and nor was their relationship to other relatedclades withinhaplogroup E andhaplogroup DE. But in non-standard or older terminologies, E-M215 is for example approximately the same as "haplotype V", still used in publications such asGérard et al. (2006).^[24]

Phylogenetic trees

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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 followingphylogenetic 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)^[25]^[23]^[24]

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)^[26]
    - E-V257/L19 (L19, V257) – E1b1b1b1^[26]
      - 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^[26]
      - 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

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^Cruciani et al. (2004): "Several observations point to eastern Africa as the homeland for haplogroup E3b—that is, it had (1) the highest number of different E3b clades (table 1), (2) a high frequency of this haplogroup and a high microsatellite diversity, and, finally, (3) the exclusive presence of the undifferentiated E3b*paragroup." As mentioned above, "E3b" is the old name for E-M215.Semino et al. (2004): "This inference is further supported by the presence of additional Hg E lineal diversification and by the highest frequency of E-P2* and E-M35* in the same region. The distribution of E-P2* appears limited to eastern African peoples. The E-M35* lineage shows its highest frequency (19.2%) in the Ethiopian Oromo but with a wider distribution range than E-P2*." For E-M215Cruciani et al. (2007) reduced their estimate to 22,400 from 25,600 inCruciani et al. (2004), re-calibrating the same data.
^"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)

References

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^^a ^bTrombetta 2015.
^Haber M, Jones AL, Connel BA, Asan, Arciero E, Huanming Y, Thomas MG, Xue Y, Tyler-Smith C (June 2019)."A Rare Deep-Rooting D0 African Y-chromosomal Haplogroup and its Implications for the Expansion of Modern Humans Out of Africa".Genetics.212 (4):1421–1428.doi:10.1534/genetics.119.302368.PMC 6707464.PMID 31196864.
^^a ^bCruciani et al. (2007)
^"E-M215 YTree".
^Cruciani et al. (2004).
^Schuenemann, Verena J.; et al. (2017)."Ancient Egyptian mummy genomes suggest an increase of Sub-Saharan African ancestry in post-Roman periods".Nature Communications.8 15694.Bibcode:2017NatCo...815694S.doi:10.1038/ncomms15694.PMC 5459999.PMID 28556824.
^Fregel; et al. (2018)."Ancient genomes from North Africa evidence prehistoric migrations to the Maghreb from both the Levant and Europe".Proceedings of the National Academy of Sciences.115 (26):6774–6779.bioRxiv 10.1101/191569.doi:10.1073/pnas.1800851115.PMID 29895688.
^Rodrı́guez-Varela; et al. (2017)."Genomic Analyses of Pre-European Conquest Human Remains from the Canary Islands Reveal Close Affinity to Modern North Africans".Current Biology.27 (1–7): 3396–3402.e5.Bibcode:2017CBio...27E3396R.doi:10.1016/j.cub.2017.09.059.hdl:2164/13526.PMID 29107554.
^Van De Loosdrecht, Marieke; Bouzouggar, Abdeljalil; Humphrey, Louise; Posth, Cosimo; Barton, Nick; Aximu-Petri, Ayinuer; Nickel, Birgit; Nagel, Sarah; Talbi, El Hassan; El Hajraoui, Mohammed Abdeljalil; Amzazi, Saaïd; Hublin, Jean-Jacques; Pääbo, Svante; Schiffels, Stephan; Meyer, Matthias; Haak, Wolfgang; Jeong, Choongwon; Krause, Johannes (4 May 2018)."Pleistocene North African genomes link Near Eastern and sub-Saharan African human populations".Science.360 (6388):548–552.Bibcode:2018Sci...360..548V.doi:10.1126/science.aar8380.PMID 29545507.S2CID 206666517.
^Martiniano, Rui; De Sanctis, Bianca; Hallast, Pille; Durbin, Richard (February 2022)."Placing Ancient DNA Sequences into Reference Phylogenies".Molecular Biology and Evolution.39 (2) msac017.doi:10.1093/molbev/msac017.PMC 8857924.PMID 35084493.
^^a ^b ^cCruciani et al. (2004)
^^a ^b ^c ^dSemino et al. (2004)
^Rosser et al. (2000)
^Firasat et al. (2006)
^Di Cristofaro, Julie; et al. (October 18, 2013)."Afghan Hindu Kush: Where Eurasian Sub-Continent Gene Flows Converge".PLOS ONE.8 (10) e76748.Bibcode:2013PLoSO...876748D.doi:10.1371/journal.pone.0076748.ISSN 1932-6203.OCLC 5534533323.PMC 3799995.PMID 24204668.S2CID 16455960.
^Ehret, Keita & Newman (2004);Keita & Boyce (2005);Keita (2008).
^Behar et al. (2003)
^Behar et al. (2004)
^Shen et al. (2004)
^Adams et al. (2008)
^Nebel et al. (2001)
^^a ^b ^cMoran, Colin N.; et al. (2004)."Y chromosome haplogroups of elite Ethiopian endurance runners".Human Genetics.115 (6):492–7.doi:10.1007/s00439-004-1202-y.PMID 15503146.S2CID 13960753. Retrieved6 February 2017.
^^a ^bKarafet et al. (2008)
^^a ^bY Chromosome Consortium "YCC" (2002)
^ISOGG (2011)
^^a ^b ^cISOGG 2015

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