|Possible time of origin||25,000–30,000 years BP|
|Possible place of origin||South Asia|
|Defining mutations||M11, M20, M61, M185, L656, L863, L878, L879[web 1]|
|Highest frequencies||South Asians, Burusho, Kalash, Pashtuns, Tamil Kallars, Afshar village, Raqqa, Balochistan, northern Afghanistan, Fascia, Veneto, southern Tyrol|
Haplogroup L-M20 is a human Y-DNA haplogroup, which is defined by SNPs M11, M20, M61 and M185. As a secondary descendant of haplogroup K and a primary branch of haplogroup LT (a.k.a. K1), the basal haplogroup L* currently has the alternative phylogenetic name of K1a, and is a sibling of haplogroup T* (a.k.a. K1b).
L-M20 is most commonly found in populations native to South Asia. The clade also occurs in Tajikistan and Anatolia, as well as at lower frequencies in Iran. It has also been present at low frequencies for millennia at very low levels in the Caucasus, Europe and Central Asia. The subclade L2 (L-L595), while it is extremely rare, is seldom found outside Western Europe.
- 1 Phylogenetic tree
- 2 Origins
- 3 Geographical distribution
- 4 Subclade distribution
- 5 Ancient DNA
- 6 Nomenclature
- 7 See also
- 8 Footnotes
- 9 References
- 10 Sources
- 11 External links
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.[web 1] The International Society of Genetic Genealogy (ISOGG) also provides an amateur tree.
This section needs expansion. You can help by adding to it. (January 2013)
This is Thomas Krahn at the Genomic Research Center's Draft tree Proposed Tree for haplogroup L-M20:[web 1]
- L-M20 M11, M20, M61, M185, L656, L863, L878, L879
- L-M22 (L1) M22, M295, PAGES00121
- L-M317 (L1b) M317, L655
- L-M349 (L1b1) M349
- L-M274 M274
- L-L1310 L1310
- L-L1304 L1304
- L-M27 (L1a1) M27, M76, P329.1, L1318, L1319, L1320, L1321
- L-M357 (l1a2) M357
- L-PK3 PK3
- L-L1305 L1305, L1306, L1307
- L-M317 (L1b) M317, L655
- L-L595 (L2) L595
- L-L864 L864, L865, L866, L867, L868, L869, L870, L877
- L-M22 (L1) M22, M295, PAGES00121
L-M20 is a descendant of Haplogroup LT, which is a descendant of haplogroup K-M9. According to Dr. Spencer Wells, L-M20 originated in the rugged and mountainous Pamir Knot region in Tajikistan and migrated into Pakistan and India ca. 30,000 years ago. However, most other studies have proposed a West Asian origin for L-M20 and associated its expansion in the Indus valley to neolithic farmers. McElreavy and Quintana-Murci, writing on the Indus Valley Civilisation, state that
One Y-chromosome haplogroup (L-M20) has a high mean frequency of 14% in Pakistan and so differs from all other haplogroups in its frequency distribution. L-M20 is also observed, although at lower frequencies, in neighbouring countries, such as India, Tajikistan, Uzbekistan and Russia. Both the frequency distribution and estimated expansion time (~7,000 YBP) of this lineage suggest that its spread in the Indus Valley may be associated with the expansion of local farming groups during the Neolithic period.
Sengupta et al. (2006) observed three subbranches of haplogroup L: L1-M76 (L1a1), L2-M317 (L1b) and L3-M357 (L1a2). All three are found mostly in South Asia. According to Sengupta et al. (2006), the L-M20 subclade L-M76 (L1a1) "underwent early diversification in South India and subsequently expanded toward peripheral regions." They note that the frequency and microsatellite variance of L1 are highest in south India, the southwest and the west coast, proposing "a pattern of spread emanating from southern India." They further note that haplogroup L-M76 "is clearly predominant in Dravidian speakers," concluding that "our data provide overwhelming support for an Indian origin of Dravidian speakers."
Sengupta et al. (2006) further note that L3-M357 (L1a2) "occurs with an intermediate frequency in Pakistan (6.8%), [while] it is very rare in India (0.4%). Conversely, L1-M76 occurs at a frequency of 7.5% in India and 5.1% in Pakistan," which may be an indication that L-M20 originated in the Pakistan region.
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).
Preliminary evidence gleaned from non-scientific sources, such as individuals who have had their Y-chromosomes tested by commercial labs,[web 2] suggests that most European examples of Haplogroup L-M20 might belong to the subclade L2-M317, which is, among South Asian populations, generally the rarest of the subclades of Haplogroup L.[web 2]
It has higher frequency among Dravidian castes (ca. 17-19%) but is somewhat rarer in Indo-Aryan castes (ca. 5-6%). It reaches up to 68% in some tribes and castes of Karnataka, 38% in some castes in Gujarat, 48% in some castes in Tamil Nadu and an overall frequency of 12% in Punjab. 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. L-M20 is the single largest male lineage (36.8%) among the Jat people of Northern India and is found at 16.33% among the Gujar's of Jammu and Kashmir. It also occurs at 18.6% among the Konkanastha Brahmins of the Konkan region and at 15% among the Maratha's of Maharashtra. L-M20 is also found at 32.35% in the Vokkaligas and at 17.82% in the Lingayats of Karnataka. 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. L-M11 is found in frequncies of 8-16% among Indian Jews. 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]
The greatest concentration of Haplogroup L-M20 is along the Indus River in Pakistan where the Indus Valley Civilization flourished during 3300–1300 BC with its mature period between 2600–1900 BCE. L-M357's highest frequency and diversity is found in the Balochistan province at 28% with a moderate distribution among the general Pakistani population at 11.6% (Firasat 2007)). It is also found in Afghanistan ethnic 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. L-M20 is found at 17.78% among the Parsis. L3a is found at 23% among the Nuristanis in both Pakistan and 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. 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. 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%). An earlier study involving a lesser number of samples had reported that L1c comprises 12.24% of the Afghan Pashtun male lineages. L1c-M357 occurs significantly in the Burusho and Kalash(15% and 25%), as well. L1c is also found at 7.69% among the Balochs of Afghanistan. However L1a-M76 occurs in a much more higher frequency among the Balochs (20 to 61.54%), and is found at lower levels in Kyrgyz, Tajik, Uzbek and Turkmen populations.
Middle East and Anatolia
L-M20 was found in 51% of Syrians from Raqqa, a northern Syrian city whose previous inhabitants were wiped out by Mongol genocides and repopulated in recent times by local Bedouin populations and Chechen war refugees from Russia (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).
|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|
|Iran||54.9% (42/71) L in Priest Zoroastrian Parsis
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 2006, Cristofaro 2013, Malyarchuk 2013 and Lopez 2017|
|Syria||51.0% (33/65) of Syrians in Raqqa, 31.0% of Eastern Syrians||El-Sibai 2009|
|Laz||41.7% (15/36) L1b-M317||O. Balanovsky 2017|
|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|
|Kurds||3.2% of Kurds in Southeast Turkey||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|
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).
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 (Italy), and Andalusia (Spain). 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.
|Fascia, Italy||19.2% of Fascians L-M20||Valentina Coia 2013|
|Nonstal. Italy||10% of Nonesi L-M20||F. di Giacomo 2003|
|Samnium, Italy||10% of Aquilanis L-M20||Alessio Boattini 2013|
|Vicenza, Italy||10% of Venetians L-M20||Alessio Boattini 2013|
|South Tyrol, Italy||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.|
|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|
|Daghestan, Russia||10% of Chechens in Daghestan, 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|
|Arkhangelsk Oblast, Russia||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|
|Balkarians, Russia||5.3% (2/38) L-M317||Battaglia 2008|
|Portugal||5.0% of Coimbra||Beleza 2006|
|Bulgaria||3.9% of Bulgarians||Karafet 2016|
|Flanders, Belgium||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, Austria||L-M20 is found in 1.9% of Tyroleans in Region B (Isel, Lower Drau, Defereggen, Virgen, and Kals valley)||H.Niederstätter 2012|
|Gipuzkoa, Spain||L1b is found in 1.7% of Gipuzkoans||Young 2011|
|North Tyrol, Austria||L-M20 is found in 0.8% of Tyroleans in Reutte||D.Erhart 2012|
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 Europe, but slightly higher Middle East Arab populations (Karafet 2016). There is a very minor presence among Siddi's (2%), as well.
L-M357 is found frequently among Burushos, Kalashas, and Pashtuns, with a moderate distribution among other populations in Pakistan, Georgia, Chechens, Ingushes, northern Iran, India, the UAE, and Saudi Arabia.
In Caucasia, L-M317 has been found in Mountain Jews (2/10 = 20%), Avars (4/42 = 9.5%, 3%), Balkarians (2/38 = 5.3%), Abkhaz (8/162 = 4.9%, 2/58 = 3.4%), Chamalals (1/27 = 3.7%), Abazins (2/88 = 2.3%), Adyghes (3/154 = 1.9%), Chechens (3/165 = 1.8%), Armenians (1/57 = 1.8%), Lezgins (1/81 = 1.2%), and Ossetes (1/132 = 0.76% North Ossetians, 2/230 = 0.9% Iron).
L-M349 is principally found in Europe.
Three individuals who lived in the Chalcolithic era (c. 5700–6250 years BP), found at the Areni-1 ("Bird's Eye") cave in the South Caucasus mountains (present-day Vayots Dzor Province, Armenia), was also identified as belonging to haplogroup L1a. The individual's genome also indicated that he had red hair and blue eyes.
Elite Hun grave
|Language||unknown; possibly Hunnic|
|Date (YBP)||1540–1500 ybp|
|Burial / Location||Hungary|
|Members / Sample Size||1/1|
|Eye color (HIrisPlex System)|
|Hair color (HIrisPlex System)|
|ABO Blood Group|
|Diet (d13C%0 / d15N%0)|
|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|
|Age at Death|
|Source||Laboratory of population genetics of Kazakhstan|
Chalcolithic South Caucasus
|ID||AR1/44 I1634||AR1/46 I1632||ARE12 I1407|
|Population||Chalcolithic (Horizon III)||Chalcolithic (Horizon III)||Chalcolithic (Horizon II)|
|Culture||Late Chalcolithic||Late Chalcolithic||Late Chalcolithic|
|Date (YBP)||6161 ± 89||6086 ± 72||6025 ± 325|
|Burial / Location||Burial 2 / Areni-1 Cave||Burial 3 / Areni-1 Cave||Trench 2A, Unit 7, Square S33/T33, Locus 9, Spit 23 / Areni-1 Cave|
|Members / Sample Size||1/3||1/3||1/3|
|Eye color (HIrisPlex System)||Likely Blue|
|Hair color (HIrisPlex System)||Likely Red|
|Skin pigmentation||Likely light|
|ABO Blood Group||Likely O or B|
|Diet (d13C%0 / d15N%0)|
|Lactase Persistence||Likely 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|
|Age at Death||11 ± 2.5||15 ± 2.5|
|Notes||World’s earliest evidence of footwear and wine making|
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|
- 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.
This section needs expansion. You can help by adding to it. (January 2013)
- Original research publications
The following research teams per their publications were represented in the creation of the YCC Tree.
|Phylogenetic tree of human Y-chromosome DNA haplogroups [χ 1][χ 2]|
|A00||A0-T [χ 3]|
|A0||A1 [χ 4]|
|I||J||LT [χ 5]||K2 [χ 6]|
|L||T||K2a [χ 7]||K2b [χ 8]||K2c||K2d||K2e [χ 9]|
|K-M2313 [χ 10]||K2b1 [χ 11]||P [χ 12]|
|NO||S [χ 13]||M [χ 14]||P1||P2|
- International Society of Genetic Genealogy, 2015, Y-DNA Haplogroup Tree 2015 (30 May 2015).
- Chiaroni, J.; Underhill, P. A.; Cavalli-Sforza, L. L. (December 2009). "Y chromosome diversity, human expansion, drift, and cultural evolution". Proc. Natl. Acad. Sci. U.S.A. 106 (48): 20174–49. Bibcode:2009PNAS..10620174C. doi:10.1073/pnas.0910803106. JSTOR 25593348. PMC . PMID 19920170.
- International Society of Genetic Genealogy, 2015 Y-DNA Haplogroup K and its Subclades – 2015 (5 April 2015).
- Wells, Spencer (2007). Deep ancestry : inside the Genographic project. Washington, D.C.: National Geographic. ISBN 1426201184.
- Mahal, David G.; Matsoukas, Ianis G. (20 September 2017). "Y-STR Haplogroup Diversity in the Jat Population Reveals Several Different Ancient Origins". Frontiers in Genetics. 8. doi:10.3389/fgene.2017.00121. ISSN 1664-8021. PMC . PMID 28979290.
- Spencer Wells (2003), The Journey of Man. A Genetic Odyssey. New Delhi: Penguin Books India, p. 167
- Qamar, Raheel; Ayub, Qasim; Mohyuddin, Aisha; Helgason, Agnar; Mazhar, Kehkashan; Mansoor, Atika; Zerjal, Tatiana; Tyler-Smith, Chris; Mehdi, S. Qasim (2002). "Y-Chromosomal DNA Variation in Pakistan". American Journal of Human Genetics. 70 (5): 1107–1124. ISSN 0002-9297. PMC .
- Zhao, Zhongming; Khan, Faisal; Borkar, Minal; Herrera, Rene; Agrawal, Suraksha (2009). "Presence of three different paternal lineages among North Indians: A study of 560 Y chromosomes". Annals of human biology. 36 (1): 46–59. doi:10.1080/03014460802558522. ISSN 0301-4460. PMC .
- Thanseem, Ismail; Thangaraj, Kumarasamy; Chaubey, Gyaneshwer; Singh, Vijay Kumar; Bhaskar, Lakkakula VKS; Reddy, B Mohan; Reddy, Alla G; Singh, Lalji (7 August 2006). "Genetic affinities among the lower castes and tribal groups of India: inference from Y chromosome and mitochondrial DNA". BMC Genetics. 7: 42. doi:10.1186/1471-2156-7-42. ISSN 1471-2156. PMC .
- Cordaux, Richard; Aunger, Robert; Bentley, Gillian; Nasidze, Ivane; Sirajuddin, S. M.; Stoneking, Mark (3 February 2004). "Independent origins of Indian caste and tribal paternal lineages". Current Biology. 14 (3): 231–235. doi:10.1016/j.cub.2004.01.024. ISSN 0960-9822. PMID 14761656.
- McElreavey, K.; Quintana-Murci, L. (2005). "A population genetics perspective of the Indus Valley through uniparentally-inherited markers". Annals of Human Biology. 32 (2): 154–162. doi:10.1080/03014460500076223. ISSN 0301-4460. PMID 16096211.
- Thangaraj, Kumarasamy; Naidu, B. Prathap; Crivellaro, Federica; Tamang, Rakesh; Upadhyay, Shashank; Sharma, Varun Kumar; Reddy, Alla G.; Walimbe, S. R.; Chaubey, Gyaneshwer; Kivisild, Toomas; Singh, Lalji (20 December 2010). "The Influence of Natural Barriers in Shaping the Genetic Structure of Maharashtra Populations". PLoS ONE. 5 (12). Bibcode:2010PLoSO...515283T. doi:10.1371/journal.pone.0015283. ISSN 1932-6203. PMC .
- K. McElreavy and L. Quintana-Murci (2005), A population genetics perspective of the Indus Valley through uniparentally-inherited markers
- Sengupta 2006.
- Sengupta 2006, p. 219.
- Sengupta 2006, p. 218.
- Y HAPLOGROUP L
- Qamar 2002.
- Shah 2011.
- Kivisild, T; Rootsi, S; Metspalu, M; et al. (February 2003). "The Genetic Heritage of the Earliest Settlers Persists Both in Indian Tribal and Caste Populations". Am. J. Hum. Genet. 72 (2): 313–32. doi:10.1086/346068. PMC . PMID 12536373.
- Sharma, S; Rai, E; Sharma, P; et al. (January 2009). "The Indian origin of paternal haplogroup R1a1* substantiates the autochthonous origin of Brahmins and the caste system". Journal of Human Genetics. 54 (1): 47–55. doi:10.1038/jhg.2008.2. PMID 19158816.
- Sengupta, S; Zhivotovsky, LA; King, R; et al. (February 2006). "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". Am. J. Hum. Genet. 78 (2): 202–21. doi:10.1086/499411. PMC . PMID 16400607.
- "Analysis of Y-chromosome Diversity in Lingayat and Vokkaliga Populations of Southern India".
- Chaubey, Gyaneshwer (2016). "Genetic affinities of the Jewish populations of India". Scientific Reports. 6: 19166. Bibcode:2016NatSR...619166C. doi:10.1038/srep19166.
- Qamar, R; Ayub, Q; Mohyuddin, A; et al. (May 2002). "Y-Chromosomal DNA Variation in Pakistan". Am. J. Hum. Genet. 70 (5): 1107–24. doi:10.1086/339929. PMC . PMID 11898125.
- Firasat, S; Khaliq, S; Mohyuddin, A; et al. (January 2007). "Y-chromosomal evidence for a limited Greek contribution to the Pathan population of Pakistan". Eur. J. Hum. Genet. 15 (1): 121–26. doi:10.1038/sj.ejhg.5201726. PMC . PMID 17047675.
- Lacau, H; Gayden, T; Regueiro, M; Chennakrishnaiah, S; Bukhari, A; Underhill, PA; Garcia-Bertrand, RL; Herrera, RJ (Oct 2012). "Afghanistan from a Y-chromosome perspective". European Journal of Human Genetics. 20 (10): 1063–70. doi:10.1038/ejhg.2012.59. PMC . PMID 22510847.
- Haber, M; Platt, DE; Ashrafian Bonab, M; et al. (2012). "Afghanistan's Ethnic Groups Share a Y-Chromosomal Heritage Structured by Historical Events". PLoS ONE. 7 (3): e34288. Bibcode:2012PLoSO...734288H. doi:10.1371/journal.pone.0034288. PMC . PMID 22470552.
- Di Cristofaro, J; Pennarun, E; Mazières, S; Myres, NM; Lin, AA; Temori, SA; Metspalu, M; Metspalu, E; Witzel, M; King, RJ; Underhill, PA; Villems, R; Chiaroni, J (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. PMC . PMID 24204668.
- Fedorova 2013.
- Karafet 2010.
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