Haplogroup A-L1085

Haplogroup A-L1085
Possible time of origin 140,000 YBP,[1] 125,000 - 382,000 YBP [2]
Possible place of origin Central-Northwest Africa[1]
Ancestor Homo Y-MRCA
Descendants Haplogroup A-V148, A-P305
Highest frequencies Namibia (Tsumkwe San, Nama) 60-70%
Southern Sudan (Dinka, Shilluk, Nuer) 33%-61.5%
Ethiopia (Beta Israel ) 41%-46%

In human population genetics, Haplogroup A-L1085 is the lineage of almost all human males alive today. As a deep branch on the homo tree, it has two functions in population genetics. The first is as an inclusive parent branch. The second is a paragroup A-L1085*.

Origin

Haplogroup A is common among Khoisan people.

Many proposals for haplogroup A-L1085's origin suggest it was associated with the ancestral population of Southern Africa's hunter-gatherers. This is because haplogroup A-L1085 lineages are frequent among the San people.

However, the A-L1085 lineages of Southern Africa are subclades of A lineages found in other parts of Africa. This suggests that A-L1085 lineages arrived in Southern Africa from elsewhere.[3] The two most basal lineages of Haplogroup A-L1085, A-V148 and A-P305, have been detected in West Africa, Northwest Africa and Central Africa. Cruciani et al. 2011 suggests that these lineages may have emerged somewhere in between Central and Northwest Africa, though such an interpretation is still preliminary due to the incomplete geographic coverage of African y-chromosomes.[1]

Initial studies reported that Haplogroup A-L1085 lineages emerged around 60,000 years ago which was significantly more recent than TMRCA for mitochondrial DNA lineages which coalesce to between 150-200kya. Cruciani et al. 2011 with major restructuring of branches pushed back the root of the Y-chromosome tree to 142,000 years ago.[1]

In November 2012, a new study by Scozzari et al. reinforced "the hypothesis of an origin in the north-western quadrant of the African continent for the A1b haplogroup, and, together with recent findings of ancient Y-Chromosome lineages in central-western Africa, provide new evidence regarding the geographical origin of human MSY diversity".[4]

Distribution

Paragroup A-P305*

Haplogroup A-P305* is largely restricted to parts of Africa, though a handful of cases have been reported in Europe and Western Asia. The clade achieves its highest modern frequencies in the Bushmen hunter-gatherer populations of Southern Africa, followed closely by many Nilotic groups in Eastern Africa. However, haplogroup A's oldest sub-clades are exclusively found in Central-Northwest Africa, where it, and consequently Y-chromosomal Adam, is believed to have originated about 140,000 years ago.[1] The clade has also been observed at notable frequencies in certain populations in Ethiopia, as well as some Pygmy groups in Central Africa.

Haplogroup A-L1085 is less common among Niger–Congo speakers, who largely belong to the E1b1a clade. Haplogroup E in general is believed to have originated in Northeast Africa,[5] and was later introduced to West Africa from where it spread around 5,000 years ago to Central, Southern and Southeastern Africa with the Bantu expansion.[6][7] According to Wood et al. (2005) and Rosa et al. (2007), such relatively recent population movements from West Africa changed the pre-existing population Y chromosomal diversity in Central, Southern and Southeastern Africa, replacing the previous haplogroups in these areas with the now dominant E1b1a lineages. Traces of ancestral inhabitants, however, can be observed today in these regions via the presence of the Y DNA haplogroups A-M91 and B-M60 that are common in certain relict populations, such as the Mbuti Pygmies and the Khoisan.[8][9][10]

Haplogroup A frequencies
Africa
Study population Freq.
(in %)
[9] Tsumkwe San (Namibia) 66%
[9] Nama (Namibia) 64
[11] Dinka (Sudan) 62
[11] Shilluk (Sudan) 53
[11] Nuba (Sudan) 46
[12] Khoisan 44
[13][14] Ethiopian Jews 41
[9][13] !Kung/Sekele ~40
[11] Borgu (Sudan) 35
[11] Nuer (Sudan) 33
[11] Fur (Sudan) 31
[9] Maasai (Kenya) 27
[15] Nara (Eritrea) 20
[11] Masalit (Sudan) 19
[9][16] Amhara (Ethiopia) ~16
[12] Ethiopians 14
[17] Bantu (Kenya) 14
[9] Mandara (Cameroon) 14
[11] Hausa (Sudan) 13
[13] Khwe (South Africa) 12
[13] Fulbe (Cameroon) 12
[9] Dama (Namibia) 11
[16] Oromo (Ethiopia) 10
[15] Kunama (Eritrea) 10
[9] South Semitic (Ethiopia) 10
[17] Arabs (Egypt) 3

In a composite sample of 3551 African men, Haplogroup A had a frequency of 5.4%.[18] The highest frequencies of haplogroup A have been reported among the Khoisan of Southern Africa, Beta Israel, and Nilo-Saharans from Sudan.

Africa —Central

Haplogroup A-M13 has been observed in populations of northern Cameroon (2/9 = 22% Tupuri,[9] 4/28 = 14% Mandara,[9] 2/17 = 12% Fulbe[13]) and eastern DRC (2/9 = 22% Alur,[9] 1/18 = 6% Hema,[9] 1/47 = 2% Mbuti[9]).

Haplogroup A-M91(xA-M31,A-M6,A-M32) has been observed in the Bakola people of southern Cameroon (3/33 = 9%).[9]

Without testing for any subclade, haplogroup A-L1085 has been observed in samples of several populations of Gabon, including 9% (3/33) of a sample of Baka, 3% (1/36) of a sample of Ndumu, 2% (1/46) of a sample of Duma, 2% (1/57) of a sample of Nzebi, and 2% (1/60) of a sample of Tsogo.[7]

Africa —Eastern

Haplogroup A-M13 is common among the Southern Sudanese (53%),[11] especially the Dinka Sudanese (61.5%).[19] Haplogroup A-M13 also has been observed in another sample of a South Sudanese population at a frequency of 45% (18/40), including 1/40 A-M171.[12] Haplogroup A also has been reported in 14.6% (7/48) of an Amhara sample,[16] 10.3% (8/78) of an Oromo sample,[16] 13.6% (12/88) of another sample from Ethiopia,[12] and 41% of a sample of the Beta Israel (Cruciani et al. 2002), and important percentages are also shared by Bantus in Kenya (14%, Luis et al. 2004) and Iraqw in Tanzania (3/43 = 7.0% (Luis et al. 2004) to 1/6 = 17% (Knight et al. 2003)).

Africa —Northern

The subclade A1 has been observed in Moroccan Berbers, while the subclade A-M13 has been observed in approximately 3% of Egyptian males.

Africa —Southern

One study has found haplogroup A in samples of various Khoisan-speaking tribes with frequency ranging from 10% to 70%.[9] Surprisingly, this particular haplogroup was not found in a sample of the Hadzabe from Tanzania, a population traditionally considered an ancient remnant of Khoisans due to the presence of click consonants in their language.

Eurasia

Haplogroup A has been observed as A1 in European men in England. As A-M13, it has been observed with low frequency in Asia Minor, the Middle East, and some Mediterranean islands, among Aegean Turks, Sardinians, Palestinians, Jordanians, Yemenites, and Omanis. Without testing for any subclade, haplogroup A has been observed in a sample of Greeks from Mitilini on the Aegean island of Lesvos[20] and in samples of Portuguese from southern Portugal, central Portugal, and Madeira.[21] The authors of one study have reported finding what appears to be haplogroup A in 3.1% (2/65) of a sample of Cypriots,[22] though they have not definitively excluded the possibility that either of these individuals may belong to haplogroup B or haplogroup C.

Subclades

Diversion of haplogroup A (Y-DNA) and its descendants.

A-P305

A-P305 is found in Bakola Pygmies (South Cameroon) at 8.3% and Berbers from Algeria at 1.5%[1] and in Ghana.[4]

A-M31

The subclade A-M31 has been found in approximately 2.8% (8/282) of a pool of seven samples of various ethnic groups in Guinea-Bissau, especially among the Papel-Manjaco-Mancanha (5/64 = 7.8%).[8] An earlier study, Gonçalves et al. 2003, reported finding A-M31 in 5.1% (14/276) of a sample from Guinea-Bissau and in 0.5% (1/201) of a pair of samples from Cabo Verde.[23] The authors of another study have reported finding haplogroup A-M31 in 5% (2/39) of a sample of Mandinka from Senegambia and 2% (1/55) of a sample of Dogon from Mali.[9] Haplogroup A-M31 also has been found in 3% (2/64) of a sample of Berbers from Morocco[13] and 2.3% (1/44) of a sample of unspecified ethnic affiliation from Mali.[12]

At least seven men with ancestral origins in Yorkshire, England, and sharing the distinctive surname Revis, have been identified as belonging to subclade A-M31. News reports suggested that the men were phenotypically "European" and unaware ypicallyunaware of any African ancestry. Subsequent research suggested that they shared a common patrilineal ancestor in the 18th century.[18]

A-M6

A-M6 (formerly A2) is typically found among Khoisan peoples. The authors of one study have reported finding haplogroup A-M6(xA-P28) in 28% (8/29) of a sample of Tsumkwe San and 16% (5/32) of a sample of !Kung/Sekele, and haplogroup A-P28 in 17% (5/29) of a sample of Tsumkwe San, 9% (3/32) of a sample of !Kung/Sekele, 9% (1/11) of a sample of Nama, and 6% (1/18) of a sample of Dama.[9] The authors of another study have reported finding haplogroup A-M6 in 15.4% (6/39) of a sample of Khoisan males, including 5/39 A-M6(xA-M114) and 1/39 A-M114.[12]

A-M32

The clade A-M32 (formerly A3) contains the most populous branches of haplogroup A-L1085 and is mainly found in Eastern Africa and Southern Africa.

A-M28

The subclade A-M28 (formerly A3a) has only been rarely observed in the Horn of Africa. In 5% (1/20) of a mixed sample of speakers of South Semitic languages from Ethiopia,[9] 1.1% (1/88) of a sample of Ethiopians,[12] and 0.5% (1/201) in Somalis.[24]

A-M51

The subclade A-M51 (formerly A3b1) occurs most frequently among Khoisan peoples (6/11 = 55% Nama,[9] 11/39 = 28% Khoisan,[12] 7/32 = 22% !Kung/Sekele,[9] 6/29 = 21% Tsumkwe San,[9] 1/18 = 6% Dama[9]). However, it also has been found with lower frequency among Bantu peoples of Southern Africa, including 2/28 = 7% Sotho–Tswana,[9] 3/53 = 6% non-Khoisan Southern Africans,[12] 4/80 = 5% Xhosa,[9] and 1/29 = 3% Zulu.[9]

A-M13

The subclade A-M13 (formerly A3b2) that is commonly found in East Africa and northern Cameroon is different from those found in the Khoisan samples and only remotely related to them. This finding suggests an ancient divergence.

In Sudan, haplogroup A-M13 has been found in 28/53 = 52.8% of Southern Sudanese, 13/28 = 46.4% of the Nuba of central Sudan, 25/90 = 27.8% of Western Sudanese, 4/32 = 12.5% of local Hausa people, and 5/216 = 2.3% of Northern Sudanese.[25]

In Ethiopia, one study has reported finding haplogroup A-M13 in 14.6% (7/48) of a sample of Amhara and 10.3% (8/78) of a sample of Oromo.[16] Another study has reported finding haplogroup A-M118 in 6.8% (6/88) and haplogroup A-M13(xA-M171, A-M118) in 5.7% (5/88) of a mixed sample of Ethiopians, amounting to a total of 12.5% (11/88) A-M13.[12]

Haplogroup A-M13 also has been observed occasionally outside of Central and Eastern Africa, as in the Aegean Region of Turkey (2/30 = 6.7%[26]), Yemenite Jews (1/20 = 5%[14]), Egypt (4/147 = 2.7%,[17] 3/92 = 3.3%[9]), Palestinian Arabs (2/143 = 1.4%[27]), Sardinia (1/77 = 1.3%,[28] 1/22 = 4.5%[12]), the capital of Jordan, Amman (1/101=1%[29]), and Oman (1/121 = 0.8%[17]).

Phylogenetics

Phylogenetic history

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
A-M317I1A1–H1AA1A1A1A1aA1A1A1aA1aA1aA1aA1a
A-M627I23–H1AA2*A2A2A2A2A2A2A2A2A2A1b1a1a
A-M11427I23–H1AA2aA2aA2aA2aA2aA2aA2aA2aA2aA2aA1b1a1a1a
A-P2827I24–H1AA2bA2bA2bA2bA2bA2bA2bA2bA2bA2bA1b1a1a1b
A-M32********A3A3A3A3A3A3A3A3A3A1b1b
A-M287I1A1–H1AA3aA3aA3aA3aA3aA3aA3aA3aA3aA3aA1b1b1
A-M517I1A1–H1AA3b1A3b1A3b1A3b1A3b1A3b1A3b1A3b1A3b1A3b1A1b1b2a
A-M137I1A2Eu1H1AA3b2*A3b2A3b2A3b2A3b2A3b2A3b2A3b2A3b2A3b2A1b1b2b
A-M1717I1A2Eu1H1AA3b2aA3b2aA3b2aA3b2aA3b2aA3b2aA3b2aA3b2aA3b2aA3b2aremoved
A-M1187I1A2Eu1H1AA3b2bA3b2bA3b2bA3b2bA3b2bA3b2bA3b2bA3b2bA3b2bA3b2bA1b1b2b1

Original research publications

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

Cruciani 2011

The revised y-chromosome family tree by Cruciani et al. 2011 compared with the family tree from Karafet et al. 2008. A1b is now known as A0 according to ISOGG.

A major shift in understanding of the Haplogroup A tree came with the publication of (Cruciani 2011). Initial sequencing of the human y-chromosome suggested that first split in the Y-Chromosome family tree occurred with the M91 mutation that separated Haplogroup A from Haplogroup BT.[30] However, it is now known that deepest split in the Y-chromosome tree is found between two previously reported subclades of Haplogroup A, rather than between Haplogroup A and Haplogroup BT. Subclades A1b and A1a-T now descend directly from the root of the tree. The rearrangement of the Y-chromosome family tree implies that lineages classified as Haplogroup A do not necessarily form a monophyletic clade.[1] Haplogroup A therefore refers to a collection of lineages that do not possess the markers that define Haplogroup BT, though many lineages within haplogroup A are only very distantly related.

The M91 and P97 mutations distinguish Haplogroup A from Haplogroup BT . Within Haplogroup A chromosomes, the M91 marker consists of a stretch of 8 T nucleobase units. In Haplogroup BT and chimpanzee chromosomes, this marker consists of 9 T nucleobase units. This pattern suggested that the 9T stretch of Haplogroup BT was the ancestral version and that Haplogroup A was formed by the deletion of one nucleobase.[1][30]

But according to Cruciani et al. 2011, the region surrounding the M91 marker is a mutational hotspot prone to recurrent mutations. It is therefore possible that the 8T stretch of Haplogroup A may be the ancestral state of M91 and the 9T of Haplogroup BT may be the derived state that arose by an insertion of 1T. This would explain why subclades A1b and A1a-T, the deepest branches of Haplogroup A, both possess the 8T stretch. Furthermore, Cruciani et al. 2011 determined that the P97 marker, which is also used to identify haplogroup A, possessed the ancestral state in haplogroup A but the derived state in Haplogroup BT.[1]

Phylogenetic trees

This phylogenetic tree of haplogroup subclades is based on the Y-Chromosome Consortium (YCC) Tree,[31] the ISOGG Y-DNA Haplogroup Tree,[6] and subsequent published research.

Y-chromosomal Adam

See also

Genetics

Y-DNA A subclades

  • A-M114
  • A-M118
  • A-M13
  • A-M171
  • A-M28
  • A-M31
  • A-M32
  • A-M51
  • A-M6
  • A-P28

Y-DNA backbone tree

Phylogenetic tree of human Y-chromosome DNA haplogroups [χ 1][χ 2]
"Y-chromosomal Adam"
A00 A0-T [χ 3]
A0 A1 [χ 4]
A1a A1b
A1b1 BT
B CT
DE CF
D E C F
F1  F2  F3  GHIJK
G HIJK
IJK H
IJ   K
I J    LT [χ 5]  K2
L T [χ 6] NO [χ 7] K2b [χ 8]     K2c  K2d  K2e [χ 9]
N   O   K2b1 [χ 10]     P
K2b1a[χ 11]     K2b1b K2b1c      M     P1 P2
K2b1a1   K2b1a2   K2b1a3 S [χ 12] Q   R
  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 LT (L298/P326) is also known as Haplogroup K1.
  6. ↑ Between 2002 and 2008, Haplogroup T (M184) was known as "Haplogroup K2" – that name has since been re-assigned to K-M526, the sibling of Haplogroup LT.
  7. ↑ Haplogroup NO (M214) is also known as Haplogroup K2a (although the present Haplogroup K2e was also previously known as "K2a").
  8. ↑ Haplogroup K2b (M1221/P331/PF5911) is also known as Haplogroup MPS.
  9. ↑ Haplogroup K2e (K-M147) was previously known as "Haplogroup X" and "K2a" (but is a sibling subclade of the present K2a, also known as Haplogroup NO).
  10. ↑ Haplogroup K2b1 (P397/P399) is similar to the former Haplogroup MS, but has a broader and more complex internal structure.
  11. ↑ Haplogroup K2b1a has also been known as Haplogroup S-P405.
  12. ↑ Haplogroup S (S-M230), also known as K2b1a4, was previously known as Haplogroup K5.

References

  1. 1 2 3 4 5 6 7 8 9 Cruciani F, Trombetta B, Massaia A, Destro-Bisol G, Sellitto D, Scozzari R (Jun 2011). "A revised root for the human Y chromosomal phylogenetic tree: the origin of patrilineal diversity in Africa". American Journal of Human Genetics. 88 (6): 814–8. doi:10.1016/j.ajhg.2011.05.002. PMC 3113241Freely accessible. PMID 21601174.
  2. ↑ Mendez FL, Krahn T, Schrack B, Krahn AM, Veeramah KR, Woerner AE, Fomine FL, Bradman N, Thomas MG, Karafet TM, Hammer MF (Mar 2013). "An African American paternal lineage adds an extremely ancient root to the human Y chromosome phylogenetic tree". American Journal of Human Genetics. 92 (3): 454–9. doi:10.1016/j.ajhg.2013.02.002. PMC 3591855Freely accessible. PMID 23453668.
  3. ↑ Batini C, Ferri G, Destro-Bisol G, Brisighelli F, Luiselli D, Sánchez-Diz P, Rocha J, Simonson T, Brehm A, Montano V, Elwali NE, Spedini G, D'Amato ME, Myres N, Ebbesen P, Comas D, Capelli C (Sep 2011). "Signatures of the preagricultural peopling processes in sub-Saharan Africa as revealed by the phylogeography of early Y chromosome lineages" (PDF). Molecular Biology and Evolution. 28 (9): 2603–13. doi:10.1093/molbev/msr089. PMID 21478374.
  4. 1 2 Scozzari R, Massaia A, D'Atanasio E, Myres NM, Perego UA, Trombetta B, Cruciani F (2012). "Molecular dissection of the basal clades in the human Y chromosome phylogenetic tree". PLOS ONE. 7 (11): e49170. doi:10.1371/journal.pone.0049170. PMC 3492319Freely accessible. PMID 23145109.
  5. ↑ Abu-Amero KK, Hellani A, González AM, Larruga JM, Cabrera VM, Underhill PA (2009). "Saudi Arabian Y-Chromosome diversity and its relationship with nearby regions". BMC Genetics. 10 (59): 59. doi:10.1186/1471-2156-10-59. PMC 2759955Freely accessible. PMID 19772609.
  6. 1 2 International Society of Genetic Genealogy. "Y-DNA Haplogroup Tree". Retrieved 2012. Check date values in: |access-date= (help)
  7. 1 2 Berniell-Lee G, Calafell F, Bosch E, Heyer E, Sica L, Mouguiama-Daouda P, van der Veen L, Hombert JM, Quintana-Murci L, Comas D (Jul 2009). "Genetic and demographic implications of the Bantu expansion: insights from human paternal lineages" (Epub). Molecular Biology and Evolution. 26 (7): 1581–9. doi:10.1093/molbev/msp069. PMID 19369595.
  8. 1 2 Rosa A, Ornelas C, Jobling MA, Brehm A, Villems R (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 1976131Freely accessible. PMID 17662131.
  9. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 Wood ET, Stover DA, Ehret C, Destro-Bisol G, Spedini G, McLeod H, Louie L, Bamshad M, Strassmann BI, Soodyall H, Hammer MF (Jul 2005). "Contrasting patterns of Y chromosome and mtDNA variation in Africa: evidence for sex-biased demographic processes". European Journal of Human Genetics. 13 (7): 867–76. doi:10.1038/sj.ejhg.5201408. PMID 15856073. cf. Appendix A: Y Chromosome Haplotype Frequencies
  10. ↑ Underhill PA, 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 (Pt 1): 43–62. doi:10.1046/j.1469-1809.2001.6510043.x. PMID 11415522.
  11. 1 2 3 4 5 6 7 8 9 28/53 (Dinka, Nuer, and Shilluk), Hassan HY, Underhill PA, Cavalli-Sforza LL, Ibrahim ME (Nov 2008). "Y-chromosome variation among Sudanese: restricted gene flow, concordance with language, geography, and history" (PDF). American Journal of Physical Anthropology. 137 (3): 316–23. doi:10.1002/ajpa.20876. PMID 18618658.
  12. 1 2 3 4 5 6 7 8 9 10 11 Underhill PA, Shen P, Lin AA, Jin L, Passarino G, Yang WH, Kauffman E, Bonné-Tamir B, Bertranpetit J, Francalacci P, Ibrahim M, Jenkins T, Kidd JR, Mehdi SQ, Seielstad MT, Wells RS, Piazza A, Davis RW, Feldman MW, Cavalli-Sforza LL, Oefner PJ (Nov 2000). "Y chromosome sequence variation and the history of human populations". Nature Genetics. 26 (3): 358–61. doi:10.1038/81685. PMID 11062480.
  13. 1 2 3 4 5 6 Cruciani F, Santolamazza P, Shen P, Macaulay V, Moral P, Olckers A, Modiano D, Holmes S, Destro-Bisol G, Coia V, Wallace DC, Oefner PJ, Torroni A, Cavalli-Sforza LL, Scozzari R, Underhill PA (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): 1197–214. doi:10.1086/340257. PMC 447595Freely accessible. PMID 11910562.
  14. 1 2 Shen P, Lavi T, Kivisild T, Chou V, Sengun D, Gefel D, Shpirer I, Woolf E, Hillel J, Feldman MW, Oefner PJ (Sep 2004). "Reconstruction of patrilineages and matrilineages of Samaritans and other Israeli populations from Y-chromosome and mitochondrial DNA sequence variation". Human Mutation. 24 (3): 248–60. doi:10.1002/humu.20077. PMID 15300852.
  15. 1 2 Cruciani F, Trombetta B, Sellitto D, Massaia A, Destro-Bisol G, Watson E, Beraud Colomb E, Dugoujon JM, Moral P, Scozzari R (Jul 2010). "Human Y chromosome haplogroup R-V88: a paternal genetic record of early mid Holocene trans-Saharan connections and the spread of Chadic languages". European Journal of Human Genetics. 18 (7): 800–7. doi:10.1038/ejhg.2009.231. PMC 2987365Freely accessible. PMID 20051990.
  16. 1 2 3 4 5 Semino O, Santachiara-Benerecetti AS, Falaschi F, Cavalli-Sforza LL, Underhill PA (Jan 2002). "Ethiopians and Khoisan share the deepest clades of the human Y-chromosome phylogeny". American Journal of Human Genetics. 70 (1): 265–8. doi:10.1086/338306. PMC 384897Freely accessible. PMID 11719903.
  17. 1 2 3 4 Luis JR, Rowold DJ, Regueiro M, Caeiro B, Cinnioğlu C, Roseman C, Underhill PA, Cavalli-Sforza LL, Herrera RJ (Mar 2004). "The Levant versus the Horn of Africa: evidence for bidirectional corridors of human migrations". American Journal of Human Genetics. 74 (3): 532–44. doi:10.1086/382286. PMC 1182266Freely accessible. PMID 14973781.
  18. 1 2 King TE, Parkin EJ, Swinfield G, Cruciani F, Scozzari R, Rosa A, Lim SK, Xue Y, Tyler-Smith C, Jobling MA (Mar 2007). "Africans in Yorkshire? The deepest-rooting clade of the Y phylogeny within an English genealogy". European Journal of Human Genetics. 15 (3): 288–93. doi:10.1038/sj.ejhg.5201771. PMC 2590664Freely accessible. PMID 17245408.
    News article: "Yorkshire clan linked to Africa". BBC News. 2007-01-24. Retrieved 2007-01-27.
  19. ↑ 16/26, Hassan et al. 2008
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  21. ↑ Gonçalves R, Freitas A, Branco M, Rosa A, Fernandes AT, Zhivotovsky LA, Underhill PA, Kivisild T, Brehm A (Jul 2005). "Y-chromosome lineages from Portugal, Madeira and Açores record elements of Sephardim and Berber ancestry". Annals of Human Genetics. 69 (Pt 4): 443–54. doi:10.1111/j.1529-8817.2005.00161.x. PMID 15996172.
  22. ↑ Capelli C, 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 (Mar 2006). "Population structure in the Mediterranean basin: a Y chromosome perspective". Annals of Human Genetics. 70 (Pt 2): 207–25. doi:10.1111/j.1529-8817.2005.00224.x. PMID 16626331.
  23. ↑ Gonçalves R, Rosa A, Freitas A, Fernandes A, Kivisild T, Villems R, Brehm A (Nov 2003). "Y-chromosome lineages in Cabo Verde Islands witness the diverse geographic origin of its first male settlers". Human Genetics. 113 (6): 467–72. doi:10.1007/s00439-003-1007-4. PMID 12942365.
  24. ↑ Abu-Amero KK, Hellani A, González AM, Larruga JM, Cabrera VM, Underhill PA (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 2759955Freely accessible. PMID 19772609.
  25. ↑ Hisham Y. Hassan et al. (2008). "Southern Sudanese" includes 26 Dinka, 15 Shilluk, and 12 Nuer. "Western Sudanese" includes 26 Borgu, 32 Masalit, and 32 Fur. "Northern Sudanese" includes 39 Nubians, 42 Beja, 33 Copts, 50 Gaalien, 28 Meseria, and 24 Arakien.
  26. ↑ Cengiz Cinnioglu, Roy King, Toomas Kivisild et al. 2004 "Excavating Y-chromosome haplotype strata in Anatolia," Human Genetics 114 : 127–148. doi:10.1007/s00439-003-1031-4 PMID 14586639
  27. ↑ Nebel A, Filon D, Brinkmann B, Majumder PP, Faerman M, Oppenheim A (Nov 2001). "The Y chromosome pool of Jews as part of the genetic landscape of the Middle East". American Journal of Human Genetics. 69 (5): 1095–112. doi:10.1086/324070. PMC 1274378Freely accessible. PMID 11573163.
  28. ↑ Semino O, Passarino G, Oefner PJ, Lin AA, Arbuzova S, Beckman LE, De Benedictis G, Francalacci P, Kouvatsi A, Limborska S, Marcikiae M, Mika A, Mika B, Primorac D, Santachiara-Benerecetti AS, Cavalli-Sforza LL, Underhill PA (Nov 2000). "The genetic legacy of Paleolithic Homo sapiens sapiens in extant Europeans: a Y chromosome perspective". Science. 290 (5494): 1155–9. doi:10.1126/science.290.5494.1155. PMID 11073453.
  29. ↑ Flores C, Maca-Meyer N, Larruga JM, Cabrera VM, Karadsheh N, Gonzalez AM (2005). "Isolates in a corridor of migrations: a high-resolution analysis of Y-chromosome variation in Jordan". Journal of Human Genetics. 50 (9): 435–41. doi:10.1007/s10038-005-0274-4. PMID 16142507.
  30. 1 2 Karafet TM, Mendez FL, Meilerman MB, Underhill PA, Zegura SL, Hammer MF (May 2008). "New binary polymorphisms reshape and increase resolution of the human Y chromosomal haplogroup tree". Genome Research. 18 (5): 830–8. doi:10.1101/gr.7172008. PMC 2336805Freely accessible. PMID 18385274.
  31. ↑ Krahn, Thomas. "YCC Tree". Houston, Texas: FTDNA. Retrieved 16 May 2011.
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