Haplogroup L-M20

Haplogroup L-M20
Possible time of origin 25,000-30,000 years BP[1]
Possible place of origin South Asia
Ancestor LT
Defining mutations M11, M20, M61, M185, L656, L863, L878, L879 (Krahn & FTDNA 2013)
Highest frequencies South Asians, Burusho, Kalash, Pashtuns, Tamil Kallars, Afshar village, Al-Raqqah, east Balochistan, northern Afghanistan, Chechens, South Tyrol

Haplogroup L-M20 is a human Y-DNA haplogroup. It is most commonly found in populations native to South Asia – especially Afghanistan, Pakistan and South India. The clade also occurs in Tajikistan and Anatolia, as well as at lower frequencies in Iran, the Caucasus and Central Asia.

The subclade L2 (L-L595), while it is extremely rare, is seldom found outside Western Europe.

L-M20 is defined by SNPs M11, M20, M61 and M185.

Phylogenetic tree

There are several confirmed and proposed phylogenetic trees available for haplogroup L-M20. The scientifically accepted one is the Y-Chromosome Consortium (YCC) one published in Karafet 2008 and subsequently updated. A draft tree that shows emerging science is provided by Thomas Krahn at the Genomic Research Center in Houston, Texas. The International Society of Genetic Genealogy (ISOGG) also provides an amateur tree.

This is Thomas Krahn at the Genomic Research Center's Draft tree Proposed Tree for haplogroup L-M20 (Krahn & FTDNA 2013).

Origins

Haplogroup L-M20 is strongly associated with South Asia, especially the Indus Valley area.

It has also been found at low frequencies among populations of Central Asia and South West Asia (including Arabia, Iraq, Syria, Turkey, Lebanon, Egypt, and Yemen) as well as in Southern Europe (especially areas adjoining the Mediterranean Sea).

L-M20 is a descendant haplogroup of haplogroup K-M9, and is believed to have first appeared approximately 30,000 years ago. Gareth Henson, administrator of the Haplogroup T project at FTDNA, has theorized "I think both T and L originated in the Iraq/Iran region...the branches of L all went in different directions (L1 southeast, L2 west and L3 northeast)."

Geographical distribution

Sengupta 2006 discovered three subbranches of haplogroup L: L-M76, L-M317, and L-M357. All three are found mostly in South Asia. In Pakistan, it has highest frequency in Baluchistan.[2] In India, it has higher frequency among Dravidian castes, but is somewhat rarer in Indo-Aryan castes.[3] They make a case for an indigenous origin of L-M76 in India, by arguing that the spatial distributions of both L-M76 HG frequency and associated microsatellite variance show a pattern of spread emanating from southern India. By linking haplogroup L-M76 to the Dravidian speakers, they simultaneously argue for an Indian origin of Dravidian languages (Sengupta 2006).

Preliminary evidence gleaned from non-scientific sources, such as individuals who have had their Y-chromosomes tested by commercial labs (Henson, Hrechdakian & FTDNA 2013), suggests that most European examples of Haplogroup L-M20 might belong to the subclade L-M317, which is, among South Asian populations, generally the rarest of the subclades of Haplogroup L.(Henson, Hrechdakian & FTDNA 2013)

South Asia

India

It has higher frequency among Dravidian castes (ca. 17-19%) but is somewhat rarer in Indo-Aryan castes (ca. 5-6%).[3] It reaches up to 68% in some tribes and castes of Karnataka[4] 38% in some castes and tribes of Gujarat and reaches up to 48% in some castes in Tamil Nadu.[3][4] Earlier studies (e.g. Wells 2001) report a very high frequency (approaching 80%) of Haplogroup L-M20 in Tamil Nadu appear to have been due to extrapolation from data obtained from a sample of 84 Kallars, a Tamil-speaking higher ruler caste of Tamil Nadu, among whom 40 (approx. 48%) displayed the M20 mutation that defines Haplogroup L. The presence of haplogroup L-M20 is rare among tribal groups (ca. 5,6-7%) (Cordaux 2004, Sengupta 2006, and Thamseem 2006).

L-M20 was found 68% in the Korova tribe from Karnataka, 38% in the Bharwad caste from Junagarh district in Gujarat, 21% in Charan caste from Junagarh district in Gujarat and 17% in the Kare Vokkal tribe from Uttara Kannada in Karnataka.(Shah 2011) Also found at low frequency in other populations from Junagarh district and Uttara Kannada. It is found at 16.33% among the Gujar's of Jammu and Kashmir.[5] It also occurs at 18.6% among the Konkanastha Brahmins of the Konkan region[6] and at 15% among the Maratha's of Maharashtra.[7] L-M20 is also found at 32.35% in the Vokkaligas and at 17.82% in the Lingayats of Karnataka.[8] L-M20 is also found at 20.7% among the Ambalakarar, 16.7% among the Iyengar and 17.2% among the Iyer castes of Tamil Nadu.[7] L-M11 is found in frequncies of 8-16% among Indian Jews.[9] 2% of Siddis have also been reported with L-M11.(Shah 2011) Haplogroup L-M20 is currently present in the Indian population at an overall frequency of ca. 7-15%.[Footnote 1]

Pakistan

L-M357 highest frequency and diversity is found in Balochistan at 28%.[2] with a moderate distribution among the general Pakistani population at 11.6% (Firasat 2007)). Many of these are found in their Afghanistan counterparts as well, such as with the Pashtuns and Balochis. L-M357 is found frequently among Burusho (approx. 12% (Firasat 2007)) and Pashtuns (approx. 7% (Firasat 2007)),

L1a and L1c-M357 are found at 24% among Balochis, L1a and L1c are found at 8% among the Dravidian-speaking Brahui, L1c is found at 25% among Kalash, L1c is found at 15% among Burusho, L1a-M76 and L1b-M317 are found at 2% among the Makranis, and L1c is found at 3.6% of Sindhis according to Julie di Cristofaro et al. 2013.[10] L3a is found at 23% among the Nuristanis in both Pakistan and Afghanistan.[11]

L-PK3 is found in approximately 23% of Kalash in northwest Pakistan(Firasat 2007).

Afghanistan

A study on the Pashtun male lineages in Afghanistan, found that Haplogroup L-M20, with an overall frequency of 9.5%, is the second most abundant male lineage among them.[12] It exhibits substantial disparity in its distribution on either side of the Hindu Kush range, with 25% of the northern Afghan Pashtuns belonging to this lineage, compared with only 4.8% of males from the south.[13] Specifically, paragroup L3*-M357 accounts for the majority of the L-M20 chromosomes among Afghan Pashtuns in both the north (20.5%) and south (4.1%). [14] An earlier study involving a lesser number of samples had reported that L1c comprises 12.24% of the Afghan Pashtun male lineages.[15] L1c-M357 occurs significantly in the Burusho and Kalash(15% and 25%), as well.[16] L1c is also found at 7.69% among the Balochs of Afghanistan.[17] However L1a-M76 occurs in a much more higher frequency among the Balochs (20[18] to 61.54%),[19] and is found at lower levels in Kyrgyz, Tajik, Uzbek and Turkmen populations.[20]

Caucasus & Middle East

L-M20 was found in 51% of Syrians from Al-Raqqah, a northern Syrian city in which its previous inhabitants have been wiped out by the Mongols and repopulated in recent times by local Bedouin populations and Chechen war refugees (El-Sibai 2009). In a small sample of Israeli Druze haplogroup L-M20 was found in 7 out of 20 (35%). However, studies done on bigger samples showed that L-M20 averages 5% in Israeli Druze,[Footnote 2] 8% in Lebanese Druze,[Footnote 3] and it was not found in a sample of 59 Syrian Druze. Haplogroup L-M20 has been found in 2.0% (1/50) (Wells 2001) to 5.25% (48/914) of Lebanese (Zalloua 2008).

Populations Distribution Source
Syria 51.0% (33/65) of Syrians in Al-Raqqah, 31.0% of Eastern Syrians El-Sibai 2009
Iran 22.2% L1b and L1c in South Iran (2/9)
%8 to 16% L2-L595, L1a, L1b and L1c of Kurds in Kordestan (2-4/25)
9.1% L-M20 (7/77) of Persians in Eastern Iran
3.4% L-M76 (4/117) and 2.6% L-M317 (3/117)
for a total of 6.0% (7/117) haplogroup L-M20 in Southern Iran
3.0% (1/33) L-M357 in Northern Iran
4.2% L1c-M357 of Azeris in East Azeris (1/21)
4.8% L1a and L1b of Persians in Esfahan (2/42)		 Regueiro 2006Cristofaro 2013Malyarchuk 2013
Turkey 57% in Afshar village, 12% (10/83) in Black Sea Region, 6.6% (7/106) of Turks in Turkey, 4.2% (1/523 L-M349 and 21/523 L-M11(xM27, M349)) Cinnioğlu 2004, Gokcumen 2008 and Karafet 2016
Georgians 20% (2/10) of Georgians in Gali, 14.3% (2/14) of Georgians in Chokhatauri, 12.5% (2/16) of Georgians in Martvili, 11.8% (2/17) of Georgians in Abasha, 11.1% (2/18) of Georgians in Baghdati, 10% (1/10) of Georgians in Gardabani, 9.1% (1/11) of Georgians in Adigeni, 6.9% (2/29) of Georgians in Omalo, 5.9% (1/17) of Georgians in Gurjaani, 5.9% (1/17) of Georgians in Lentekhi and 1.5% (1/66) L-M357(xPK3) to 1.6% (1/63) L-M11 Battaglia 2008, Semino 2000 and Tarkhnishvili 2014
Saudi Arabians 15.6% ( 4/32 of L-M76 and 1/32 of L-317 ) 1.91% (2/157=1.27% L-M76 and 1/157=0.64% L-M357) Karafet 2016 and AbuAmero 2009
Daghestan 10% of Chechens, 9.5% (4/42) of Avars, 8.3% (2/24) of Tats, 3.7% (1/27) of Chamalins Yunusbaev 2006, Caciagli 2009 and Karafet 2016
Balkarians 5.3% (2/38) L-M317 Battaglia 2008
Southeastern Turkey 3.2% in Kurds Flores 2005
Iraq 3.1% (2/64) L-M22 Sanchez 2005
Armenians 1.63% (12/734) to 4.3% (2/47) Weale 2001 and Wells 2001
Omanis 1% L-M11 Luis 2004
Qataris 2.8% (2/72 L-M76) Cadenas 2008
UAE Arabs 3.0% (4/164 L-M76 and 1/164 L-M357) Cadenas 2008

Central Asia

Populations Distribution Source
Tajiks 22.5% (9/40), 11.1% (6/54) L1a and L1c in Balkh Province, 9.0% (7/78), 6.3% (1/16) L1c in Samangan Province, 5.4% (2/37) L1c in Badakhstan Province Malyarchuk 2013Wells 2001
Uzbeks 20% (1/5) L1c in Balkh Province, 14.3% (4/28) L1a and L1c in Sar-e-Pol Province, 7.5% (7/94)L1a, L1b and L1c in Jawzjan Province, 3.0% (11/366) to 3.7% (2/54) Wells 2001, Karafet 2001 and Cristofaro 2013
Uyghurs 16.7% (1/6) L1c in Kyrgyzstan Cristofaro 2013
Pamiris 16% (7/44) of Shugnanis, 12% 3/25 of Ishkashimis, 0/30 Bartangis Wells 2001
Hazaras 12.5% (1/8) L1a in Balkh Province, 1.9% (2/69) L1a in Bamiyan Province Cristofaro 2013
Yagnobis 9.7% (3/31) Wells 2001
Bukharan Arabs 9.5% (4/42) Wells 2001
Pashtuns 9.4% (5/53) L1a and L1b in Kunduz Province, 2.9% (1/34) L1c in Baghlan Province Cristofaro 2013
Dungans 8.3% (1/12) L1c in Kyrgyzstan Cristofaro 2013
Karakalpaks 4.5% (2/44) Wells 2001
Uyghurs 4.4% (3/68) Karafet 2001 and Hammer 2005[Footnote 4]
Turkmens 4.1% (3/74) L1a in Jawzjan Province Cristofaro 2013
Chelkans 4.0% (1/25) Dulik 2012 and Dulik 2012
Kyrgyzes 2.7% (1/37) L1c in Northwest Kyrgyzstan and 2.5% (1/40) L1a in Central Kyrgyzstan Cristofaro 2013
Kazan Tatars 2.6% (1/38) Wells 2001
Hui 1.9% (1/54) Karafet 2001
Bashkirs 0.64% (3/471) Lobov 2009

East Asia

Researchers studying samples of Y-DNA from populations of East Asia have rarely tested their samples for any of the mutations that define Haplogroup L. However, mutations for Haplogroup L have been tested and detected in samples of Balinese (13/641 = 2.0% L-M20), Dolgans from Sakha and Taymyr (1/67 = 1.5% L-M20) and Koreans (3/506 = 0.6% L-M20).[21][22][23]

Europe

An article by O. Semino et al. published in the journal Science (Volume 290, 10 November 2000) reported the detection of the M11-G mutation, which is one of the mutations that defines Haplogroup L, in approximately 1% to 3% of samples from Georgia, Greece, Hungary, Calabria, and Andalusia. The sizes of the samples analyzed in this study were generally quite small, so it is possible that the actual frequency of Haplogroup L-M20 among Mediterranean European populations may be slightly lower or higher than that reported by Semino et al., but there seems to be no study to date that has described more precisely the distribution of Haplogroup L-M20 in Southwest Asia and Europe.

Populations Distribution Source
Fascia 19.2% of Fascians L-M20 Valentina Coia 2013
Nonstal 10% of Nonesi L-M20 F. di Giacomo 2003
Samnium 10% of Aquilanis L-M20 Alessio Boattini 2013
Vicenza 10% of Venetians L-M20 Alessio Boattini 2013
South Tyrol 8.9% of Ladin speakers from Val Badia, 8.3% of Val Badia, 2.9% of Puster Valley, 2.2% of German speakers from Val Badia, 2% of German speakers from Upper Vinschgau, 1.9% of German speakers from Lower Vinschgau and 1.7% of Italian speakers from Bolzano Pichler 2006 and Thomas 2007.
Arkhangelsk Oblast 5.9% of Russians L1c-M357 Hongyang Xu 2014
Estonia L2-L595 and L1-M22 are found in 5.3%, 3.5%, 1.4% and 0.8% of Estonians Scozzari 2001 and Lappalainen 2007
Portugal 5.0% of Coimbra Beleza 2006
Bulgaria 3.9% of Bulgarians Karafet 2016
Flanders L1a*: 3.17% of Mechelen 2.4% of Turnhout and 1.3% of Kempen. L1b*: 0.74% of West Flanders and East Flanders Larmuseau 2010 and Larmuseau 2011
East Tyrol L-M20 is found in 1.9% of Tyroleans in Region B (Isel, Lower Drau, Defereggen, Virgen, and Kals valley) H.Niederstätter 2012
Gipuzkoa L1b is found in 1.7% of Gipuzkoans Young 2011
North Tyrol L-M20 is found in 0.8% of Tyroleans in Reutte D.Erhart 2012

Subclade distribution

L1 (M295)

L-M295 is found from Western Europe to South Asia.[Footnote 5]

The L1 subclade is also found at low frequencies on the Comoros Islands.[24]

L1a1 (M27)

L-M27 is found in 14,5% of Indians and 15% of Sri Lankans, with a moderate distribution in other populations of Pakistan, southern Iran, and European but more in Middle East Arab populations.Karafet 2016. There is a very minor presence among Siddi's(2%),[25] as well.

L1a2 (M357)

L-M357 is found frequently among Burushos, Kalashas, Chechens and Pashtuns, with a moderate distribution among other populations in Pakistan, Georgia, northern Iran, India, the UAE, and Saudi Arabia.

L-PK3

L-PK3, which is downstream of L-M357,[26] is found frequently among Kalash.

L1b (M317)

L-M317 is found at low frequency in Central Asia, Southwest Asia, and Central Europe.

L1b1 (M349)

L-M349 is principally found in Europe.

L2 (L595)

L-L595 is found only in Europe from Ireland, Iberian Peninsula and Sardinia to Eastern Europe and their highest frequency is found in Estonia,[Footnote 5] Scozzari 2001, Lappalainen 2008.

Ancient DNA

Areni-1 cave ("Bird's Eye cave"), Vayots-Dzor Province, Armenia, c. 4161 BCE

Areni-1 Cave L1a1 Areni-I Areni-II Areni-III
ID AR1/44 I1634AR1/46 I1632ARE12 I1407
Y DNAL1aL1a1-M27L1a
PopulationChalcolithic (Horizon III)Chalcolithic (Horizon III)Chalcolithic (Horizon II)
Language
CultureEarly Late ChalcolithicEarly Late ChalcolithicMiddle Late Chalcolithic
Date (YBP)6161 ± 896086 ± 726025 ± 325
Burial / LocationBurial 2, / Areni-1 CaveBurial 3, / Areni-1 CaveTrench 2A, Unit 7, Square S33/T33, Locus 9, Spit 23 / Areni-1 Cave
Members / Sample Size1/31/31/3
Percentage33.3%33.3%33.3%
mtDNAH2a1K1a8H*
Isotope Sr
Eye color (HIrisPlex System)Likely Blue
Hair color (HIrisPlex System)Likely Red
Skin pigmentationLikely light
ABO Blood GroupLikely O or B
Diet (d13C%0 / d15N%0)
FADS activity
Lactase PersistenceLikely lactose-intolerant
Oase-1 Shared DNA
Ostuni1 Shared DNA
Neanderthal Vi33.26 Shared DNA
Neanderthal Vi33.25 Shared DNA
Neanderthal Vi33.16 Shared DNA
Ancestral Component (AC)
puntDNAL K12 Ancient
Dodecad [dv3]
Eurogenes [K=36]
Dodecad [Globe13]
Genetic Distance
Parental Consanguinity
Age at Death11 ± 2.515 ± 2.5
Death Position
SNPs
Read Pairs
Sample
Source[27]
NotesWorld’s earliest evidence of footwear and wine making

Elite Hunnic skeleton remains, Hungary, Pannonian Basin

Elite Hunnic L-M20 Elite Hunnic
ID
Y DNAL-M20
PopulationHuns
LanguageHunnic language
Culture
Date (YBP)1540–1500 ybp
Burial / LocationHungary
Members / Sample Size1/1
Percentage
mtDNAD4j12
Isotope Sr
Eye color (HIrisPlex System)
Hair color (HIrisPlex System)
Skin pigmentation
ABO Blood Group
Diet (d13C%0 / d15N%0)
FADS activity
Lactase Persistence
Oase-1 Shared DNA
Ostuni1 Shared DNA
Neanderthal Vi33.26 Shared DNA
Neanderthal Vi33.25 Shared DNA
Neanderthal Vi33.16 Shared DNA
Ancestral Component (AC)
puntDNAL K12 Ancient
Dodecad [dv3]
Eurogenes [K=36]
Dodecad [Globe13]
Genetic Distance
Parental Consanguinity
Age at Death
Death Position
SNPs
Read Pairs
Sample
SourceLaboratory of population genetics of Kazakhstan
Notes

Nomenclature

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
L-M20 28 VIII 1U 27 Eu17 H5 F L* L L L-------
L-M27 28 VIII 1U 27 Eu17 H5 F L1 L1 L1 L1-------
The Y-Chromosome Consortium tree

This is the official scientific tree produced by the Y-Chromosome Consortium (YCC). The last major update was in 2008 (Karafet 2008). Subsequent updates have been quarterly and biannual. The current version is a revision of the 2010 update.[28]

Original research publications

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

See also

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

Footnotes
  1. ↑ see Basu 2003, Cordaux 2004, Sengupta 2006, and Thamseem 2006.
  2. ↑ 12/222 Shlush et al. 2008
  3. ↑ 1/25 Shlush et al. 2008
  4. ↑ In Hammer 2005, see the Supplementary Material.
  5. 1 2 FTDNA lab results, May 2011
Works cited
  1. ↑ http://www.dnaexplain.com/Publications/PDFs/Y-LineDNAHaplogroups.pdf
  2. 1 2 Qamar 2002.
  3. 1 2 3 Sengupta 2006.
  4. 1 2 Shah 2011.
  5. ↑ "The Indian origin of paternal haplogroup R1a1* substantiates the autochthonous origin of Brahmins and the caste system".
  6. ↑ "The Genetic Heritage of the Earliest Settlers Persists Both in Indian Tribal and Caste Populations".
  7. 1 2 "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".
  8. ↑ "Analysis of Y-chromosome Diversity in Lingayat and Vokkaliga Populations of Southern India".
  9. ↑ "Genetic affinities of the Jewish populations of India".
  10. ↑ http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0076748
  11. ↑ "Y-chromosomal evidence for a limited Greek contribution to the Pathan population of Pakistan".
  12. ↑ "Afghanistan from a Y-chromosome perspective".
  13. ↑ "Afghanistan from a Y-chromosome perspective".
  14. ↑ "Afghanistan from a Y-chromosome perspective".
  15. ↑ "Afghanistan's Ethnic Groups Share a Y-Chromosomal Heritage Structured by Historical Events".
  16. ↑ "Afghan Hindu Kush: Where Eurasian Sub-Continent Gene Flows Converge".
  17. ↑ "Afghanistan's Ethnic Groups Share a Y-Chromosomal Heritage Structured by Historical Events".
  18. ↑ "Afghan Hindu Kush: Where Eurasian Sub-Continent Gene Flows Converge".
  19. ↑ "Afghan Hindu Kush: Where Eurasian Sub-Continent Gene Flows Converge".
  20. ↑ "Afghan Hindu Kush: Where Eurasian Sub-Continent Gene Flows Converge".
  21. ↑ Fedorova 2013.
  22. ↑ Karafet 2010.
  23. ↑ Kim 2011.
  24. ↑ Msaidie, Said; et al. (2011). "Genetic diversity on the Comoros Islands shows early seafaring as major determinant of human biocultural evolution in the Western Indian Ocean" (PDF). European Journal of Human Genetics. 19: 89–94. doi:10.1038/ejhg.2010.128.
  25. ↑ "Indian Siddis: African Descendants with Indian Admixture".
  26. ↑ ISOGG 2016.
  27. ↑ Lazaridis, Iosif; et al. (2016). "The genetic structure of the world's first farmers". BioR XIV.
  28. ↑ "Y-DNA Haplotree". Family Tree DNA uses the Y-Chromosome Consortium tree and posts it on their website.

Journals

Websites

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