Haplogroup L-M20 Explained

L-M20
Map:Distribution Haplogroup L Y-DNA.svg
Origin-Date:30,000[1] - 43,000 years BP[2]
Ancestor:LT
Mutations:M11, M20, M61, M185, L656, L863, L878, L879
Members:Syria Raqqa, Balochistan, Northern Afghanistan, Karnataka, Tarkhan, Jats, Kalash, Nuristanis, Burusho, Pashtuns, Lazs, Afshar village, 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, haplogroup L currently has the alternative phylogenetic name of K1a, and is a sibling of haplogroup T (a.k.a. K1b).

The presence of L-M20 has been observed at varying levels throughout South Asia, peaking in populations native to Balochistan (28%), Northern Afghanistan (25%),[3] and Southern India (19%). The clade also occurs in Tajikistan and Anatolia, as well as at lower frequencies in Iran. It has also been present for millennia at very low levels in the Caucasus, Europe and Central Asia. The subclade L2 (L-L595) has been found in Europe and Western Asia, but is extremely rare.

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:

Origins

L-M20 is a descendant of Haplogroup LT,[4] which is a descendant of haplogroup K-M9.[5] [6] According to Dr. Spencer Wells, L-M20 originated in the Eurasian K-M9 clan that migrated eastwards from the Middle East, and later southwards from the Pamir Knot into present-day Pakistan and India.[7] [8] These people arrived in India approximately 30,000 years ago. Hence, it is hypothesized that the first bearer of M20 marker was born either in India or the Middle East. Other studies have proposed either a West Asian or South Asian origin for L-M20 and associated its expansion in the Indus valley to Neolithic farmers.[9] [10] [11] [12] [13] Genetic studies suggest that L-M20 may be one of the haplogroups of the original creators of the Indus Valley Civilisation. McElreavy and Quintana-Murci, writing on the Indus Valley Civilisation, state that

Sengupta et al. (2006) observed three subbranches of haplogroup L: L1-M76 (L1a1), L2-M317 (L1b) and L3-M357 (L1a2), with distinctive geographic affiliations. Almost all Indian members of haplogroup L are L1 derived, with L3-M357 occurring only sporadically (0.4%). Conversely in Pakistan, L3-M357 subclade account for 86% of L-M20 chromosomes and reaches an intermediate frequency of 6.8%, overall. L1-M76 occurs at a frequency of 7.5% in India and 5.1% in Pakistan, exhibiting peak variance distribution in the Maharashtra region in coastal western India.

Geographical distribution

In India, L-M20 has a higher frequency among Dravidian castes, but is somewhat rarer in Indo-Aryan castes. In Pakistan, it has highest frequency in Balochistan.

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, 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.

South Asia

India

It has higher frequency among Dravidian castes (ca. 17-19%) but is somewhat rarer in Indo-Aryan castes (ca. 5-6%). The presence of haplogroup L-M20 is quite rare among tribal groups (ca. 5,6-7%) (and). However, the Korova tribe of Uttara Kannada in which L-M11 occurs at 68% is an exception.

L-M20 was found at 38% in the Bharwad caste and 21% in Charan caste from Junagarh district in Gujarat. It has also been reported at 17% in the Kare Vokkal tribe from Uttara Kannada in Karnataka. It is also found at low frequencies 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.[14] [15] It also occurs at 18.6% among the Konkanastha Brahmins of the Konkan region[16] and at 15% among the Maratha's of Maharashtra.[17] L-M20 is also found at 32.35% in the Vokkaligas and at 17.82% in the Lingayats of Karnataka.[18]

And available data[19] shows that among Tamils, L-M20 is found at 48% among Kallar, 20.56% among Tamil yadavas, 28.57% among Vanniyars, 28.81% among Nadar, and 26% among the Saurashtra people, 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 frequencies of 8-16% among Indian Jews.[20] L-M20 has an overall frequency of 12% in Punjab. 2% of Siddis have also been reported with L-M11. Haplogroup L-M20 is currently present in the Indian population at an overall frequency of ca. 7-15%.[21]

Pakistan

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 and 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%). 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%) and Pashtuns (approx. 7%),

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.[22] L-M20 is found at 17.78% among the Parsis.[9] L3a is found at 23% among the Nuristanis in both Pakistan and Afghanistan.

L-PK3 is found in approximately 23% of Kalash in northwest Pakistan .

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 . 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,[23] 8% in Lebanese Druze,[24] and it was not found in a sample of 59 Syrian Druze. Haplogroup L-M20 has been found in 2.0% (1/50) to 5.25% (48/914) of Lebanese .

PopulationsDistributionSource
Turkey57% 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)),
Iran54.9% (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)		,,
Syria51.0% (33/65) of Syrians in Raqqa, 31.0% of Eastern Syrians
Laz41.7% (15/36) L1b-M317
Saudi Arabians15.6% (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)
Kurds3.2% of Kurds in Southeast Turkey
Iraq3.1% (2/64) L-M22
Armenians1.63% (12/734) to 4.3% (2/47) and
Omanis1% L-M11
Qataris2.8% (2/72 L-M76)
UAE Arabs3.0% (4/164 L-M76 and 1/164 L-M357)

Central Asia

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.[25] 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.[25] 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%).[25] An earlier study involving a lesser number of samples had reported that L1c comprises 12.24% of the Afghan Pashtun male lineages.[26] [27] L1c is also found at 7.69% among the Balochs of Afghanistan.[26] However L1a-M76 occurs in a much more higher frequency among the Balochs (20[27] to 61.54%),[27] and is found at lower levels in Kyrgyz, Tajik, Uzbek and Turkmen populations.[27]

PopulationsDistributionSource
Tajiks22.5% (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 Badakhshan Province
Uzbeks20% (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), and
Uyghurs16.7% (1/6) L1c-M357 in Kyrgyzstan
Pamiris16% (7/44) of Shugnanis, 12% 3/25 of Ishkashimis, 0/30 Bartangis
Hazaras12.5% (1/8) L1a in Balkh Province, 1.9% (2/69) L1a in Bamiyan Province
Yagnobis9.7% (3/31)
Bukharan Arabs9.5% (4/42)
Pashtuns9.4% (5/53) L1a and L1b in Kunduz Province, 2.9% (1/34) L1c in Baghlan Province
Dungans8.3% (1/12) L1c in Kyrgyzstan
Uyghurs (Lopliks)7.8% (5/64) L-M357 in Qarchugha Village, Lopnur County, Xinjiang
Karakalpaks4.5% (2/44)
Uyghurs4.4% (3/68) and [28]
Turkmens4.1% (3/74) L1a in Jawzjan Province
Chelkans4.0% (1/25) and
Kyrgyzes2.7% (1/37) L1c in Northwest Kyrgyzstan and 2.5% (1/40) L1a in Central Kyrgyzstan
Kazan Tatars2.6% (1/38)
Hui1.9% (1/54)
Bashkirs0.64% (3/471)

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), Han Chinese (1/57 = 1.8%), Dolgans from Sakha and Taymyr (1/67 = 1.5% L-M20) and Koreans (3/506 = 0.6% L-M20).

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 (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.

PopulationsDistributionSource
Fascia, Italy19.2% of Fascians L-M20
Nonstal, Italy10% of Nonesi L-M20
Samnium, Italy10% of Aquilanis L-M20
Vicenza, Italy10% of Venetians L-M20
South Tyrol, Italy8.9% 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 and .
Georgians20% (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, and
Daghestan, Russia10% of Chechens in Daghestan, 9.5% (4/42) of Avars, 8.3% (2/24) of Tats, 3.7% (1/27) of Chamalins,
Arkhangelsk Oblast, Russia5.9% of Russians L1c-M357
EstoniaL2-L595 and L1-M22 are found in 5.3%, 3.5%, 1.4% and 0.8% of Estonians and
Balkarians, Russia5.3% (2/38) L-M317
Portugal5.0% of Coimbra
Bulgaria3.9% of Bulgarians
FlandersL1a*: 3.17% of Mechelen 2.4% of Turnhout and 1.3% of Kempen. L1b*: 0.74% of West Flanders and East Flanders and
Antsiferovo, Novgorod2.3% of Russians
East Tyrol, AustriaL-M20 is found in 1.9% of Tyroleans in Region B (Isel, Lower Drau, Defereggen, Virgen, and Kals valley)
Gipuzkoa, Basque CountryL1b is found in 1.7% of Gipuzkoans
North Tyrol, AustriaL-M20 is found in 0.8% of Tyroleans in Reutte

Southern Africa and the Swahili Coast

Researchers in 2013 studying the origins of the Lemba people - who are of paternal South Arabian ancestry - found that 13.8% of Lemba males carried the Y-DNA L-M20, specifically the subclade L-M349 making it the 4th most common lineage amongst them.[29] A Lemba sample from South Africa submitted to Familytreedna in 2023 was found to carry a yet unnamed L-M349 subclade of L-FT408126 which was closest to 2 samples from Iraq and Iran.[30]

Researchers also found traces of traces of L-M20 on the Swahili coast in Kenya amounting to 4.2% of the total population.

Subclade distribution

L1 (M295)

L-M295 is found from Western Europe to South Asia.[31]

The L1 subclade is also found at low frequencies on the Comoros Islands.[32]

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 Europe, but slightly higher Middle East Arab populations. There is a very minor presence among Siddi's (2%),[33] as well.

L1a2 (M357)

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

A Chinese study published in 2018 found L-M357/L1307 in 7.8% (5/64) of a sample of Loplik Uyghurs from Qarchugha Village, Lopnur County, Xinjiang.[34]

L-PK3L-PK3, which is downstream of L-M357,[35] is found frequently among Kalash.

L1b (M317)

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

In Europe, L-M317 has been found in Northeast Italians (3/67 = 4.5%) and Greeks (1/92 = 1.1%).

In Caucasia, L-M317 has been found in Mountain Jews (2/10 = 20%), Avars (4/42 = 9.5%, 3%), Balkarians (2/38 = 5.3%),[36] Abkhaz (8/162 = 4.9%,[37] 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-M317 has been found in Makranis (2/20 = 10%) in Pakistan, Iranians (3/186 = 1.6%), Pashtuns in Afghanistan(1/87 = 1.1%), and Uzbeks in Afghanistan (1/127 = 0.79%).[27]

L1b1 (M349)

L-M349 is found in some Crimean Karaites who are Levites.[38] Some of L-M349's branches are found in West Asia, including L-Y31183 in Lebanon, L-Y31184 in Armenia, and L-Y130640 in Iraq, Iran, Yemen and South Africa. Others are found in Europe, such as L-PAGE116 in Italy, L-FT304386 in Slovenia, and L-FGC36841 in Moldova.[39] 13.8% of Lemba males carry L-M349 under the clade L-Y130640.[29] This percentage is most likely due to a founder effect in their population making them the only group on the African continent with any substantial proportion of L-M20.

L2 (L595)

L2-L595 is extremely rare, and has been identified by private testing in individuals from Europe and Western Asia.

Two confirmed L2-L595 individuals from Iran were reported in a 2020 study supplementary.[40] Possible but unconfirmed cases of L2 include 4% (1/25) L-M11(xM76, M27, M317, M357) in a sample of Iranians in Kordestan and 2% (2/100) L-M20(xM27, M317, M357) in a sample of Shapsugs,[41] among other rare reported cases of L which don't fall into the common branches.

L2 in modern populations!Region!Population!n/Sample size!Percentage!Source
West AsiaAzerbaijan2/2041[42]
Central EuropeGermany1/86410.0000115[43]
Southern EuropeGreece1/7530.1[44]
West AsiaIran2/8000.25
Southern EuropeItaly3/9130.3[45]

Ancient DNA

Chalcolithic South Caucasus

Areni-1 Cave
PropertyAreni-IAreni-IIAreni-III
IDAR1/44 I1634AR1/46 I1632ARE12 I1407
Y DNAL1aL1a1-M27L1a
PopulationChalcolithic (Horizon III)Chalcolithic (Horizon III)Chalcolithic (Horizon II)
Language
CultureLate ChalcolithicLate ChalcolithicLate 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[52]
NotesWorld's earliest evidence of footwear and wine making

Nomenclature

See 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 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 2006ISOGG 2007ISOGG 2008ISOGG 2009ISOGG 2010ISOGG 2011ISOGG 2012
L-M2028VIII1U27Eu17H5FL*LLL-------
L-M2728VIII1U27Eu17H5FL1L1L1L1-------
The Y-Chromosome Consortium treeThis is the official scientific tree produced by the Y-Chromosome Consortium (YCC). The last major update was in 2008. Subsequent updates have been quarterly and biannual. The current version is a revision of the 2010 update.[53]
Original research publicationsThe following research teams per their publications were represented in the creation of the YCC Tree.

Sources

Sources for conversion tables

÷

External links

Notes and References

  1. https://web.archive.org/web/20121023014422/http://www.genebase.com/learning/article/13 Learn about Y-chromosome Haplogroup L
  2. https://www.yfull.com/tree/L/ Yfull Tree L
  3. Lacau H, Gayden T, Regueiro M, Chennakrishnaiah S, Bukhari A, Underhill PA, Garcia-Bertrand RL, Herrera RJ . Afghanistan from a Y-chromosome perspective . En . European Journal of Human Genetics . 20 . 10 . 1063–1070 . October 2012 . 22510847 . 3449065 . 10.1038/ejhg.2012.59 .
  4. http://www.isogg.org/tree/ISOGG_YDNATreeTrunk.html International Society of Genetic Genealogy, 2015, Y-DNA Haplogroup Tree 2015
  5. http://www.isogg.org/tree/ISOGG_HapgrpK.html International Society of Genetic Genealogy, 2015 Y-DNA Haplogroup K and its Subclades – 2015 (5 April 2015).
  6. Chiaroni J, Underhill PA, Cavalli-Sforza LL . Y chromosome diversity, human expansion, drift, and cultural evolution . Proceedings of the National Academy of Sciences of the United States of America . 106 . 48 . 20174–20179 . December 2009 . 19920170 . 2787129 . 10.1073/pnas.0910803106 . free . 2009PNAS..10620174C . 25593348 .
  7. Book: Wells S . Deep Ancestry: The Landmark DNA Quest to Decipher Our Distant Past . 20 November 2007 . National Geographic Books . 978-1-4262-0211-7 . 161–162 . This part of the M9 Eurasian clan migrated south once they reached the rugged and mountainous Pamir Knot region. The man who gave rise to marker M20 was possibly born in India or the Middle East. His ancestors arrived in India around 30,000 years ago and represent the earliest significant settlement of India..
  8. Book: Wells S . The Journey of Man: A Genetic Odyssey . 28 March 2017 . Princeton University Press . 978-0-691-17601-7 . 111–113 .
  9. Qamar R, Ayub Q, Mohyuddin A, Helgason A, Mazhar K, Mansoor A, Zerjal T, Tyler-Smith C, Mehdi SQ . Y-chromosomal DNA variation in Pakistan . American Journal of Human Genetics . 70 . 5 . 1107–1124 . May 2002 . 11898125 . 447589 . 10.1086/339929 .
  10. Zhao Z, Khan F, Borkar M, Herrera R, Agrawal S . Presence of three different paternal lineages among North Indians: a study of 560 Y chromosomes . Annals of Human Biology . 36 . 1 . 46–59 . 2009 . 19058044 . 2755252 . 10.1080/03014460802558522 .
  11. Thanseem I, Thangaraj K, Chaubey G, Singh VK, Bhaskar LV, Reddy BM, Reddy AG, Singh L . Genetic affinities among the lower castes and tribal groups of India: inference from Y chromosome and mitochondrial DNA . BMC Genetics . 7 . 1 . 42 . August 2006 . 16893451 . 1569435 . 10.1186/1471-2156-7-42 . free .
  12. Cordaux R, Aunger R, Bentley G, Nasidze I, Sirajuddin SM, Stoneking M . Independent origins of Indian caste and tribal paternal lineages . Current Biology . 14 . 3 . 231–235 . February 2004 . 14761656 . 10.1016/j.cub.2004.01.024 . 5721248 . free . 2004CBio...14..231C .
  13. Thangaraj K, Naidu BP, Crivellaro F, Tamang R, Upadhyay S, Sharma VK, Reddy AG, Walimbe SR, Chaubey G, Kivisild T, Singh L . The influence of natural barriers in shaping the genetic structure of Maharashtra populations . PLOS One . 5 . 12 . e15283 . December 2010 . 21187967 . 3004917 . 10.1371/journal.pone.0015283 . free . 2010PLoSO...515283T .
  14. Mahal DG, Matsoukas IG . Y-STR Haplogroup Diversity in the Jat Population Reveals Several Different Ancient Origins . Frontiers in Genetics . 8 . 121 . 20 September 2017 . 28979290 . 5611447 . 10.3389/fgene.2017.00121 . free .
  15. Sharma S, Rai E, Sharma P, Jena M, Singh S, Darvishi K, Bhat AK, Bhanwer AJ, Tiwari PK, Bamezai RN . 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 . January 2009 . 19158816 . 10.1038/jhg.2008.2 . free .
  16. Kivisild T, Rootsi S, Metspalu M, Mastana S, Kaldma K, Parik J, Metspalu E, Adojaan M, Tolk HV, Stepanov V, Gölge M, Usanga E, Papiha SS, Cinnioğlu C, King R, Cavalli-Sforza L, Underhill PA, Villems R . The genetic heritage of the earliest settlers persists both in Indian tribal and caste populations . American Journal of Human Genetics . 72 . 2 . 313–332 . February 2003 . 12536373 . 379225 . 10.1086/346068 .
  17. Sengupta S, Zhivotovsky LA, King R, Mehdi SQ, Edmonds CA, Chow CE, Lin AA, Mitra M, Sil SK, Ramesh A, Usha Rani MV, Thakur CM, Cavalli-Sforza LL, Majumder PP, Underhill PA . 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 . American Journal of Human Genetics . 78 . 2 . 202–221 . February 2006 . 16400607 . 1380230 . 10.1086/499411 .
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