Haplogroup G-M201

Haplogroup G
Possible time of origin 9,500-30,000 years BP[1]
Possible place of origin Caucasus or Southwest Asia
Ancestor Haplogroup GHIJK
Descendants G1, G2 and their subclades
Defining mutations L116, L154, L204, L240, L269, L402, L520, L521, L522, L523, L605, L769, L770, L836, L837, M201, P257/U6, Page94/U17, U2, U3, U7, U12, U20, U21, U23, U33

Haplogroup G (M201) is a human Y-chromosome haplogroup. It is one of two branches of Haplogroup GHIJK, the other being Haplogroup HIJK.

At the level of national populations, G-M210 is most commonly found in Georgia; it is found at even higher levels among many other regional and minority populations in The Caucasus. G-M210 is also widely distributed at low frequencies, among ethnic groups of Europe, South Asia, Central Asia, and North Africa.

The most commonly occurring subclades of Haplogroup G (with their corresponding SNPs in parentheses) are: G1 (M285), G2a (P15), G2a1 (L293), G2a3a (M406), G2a3b1a (L140), G2a3b1a2 (L497), G2a3b1a1a (L13/S131/U13), G2a3b1a3 (Z1903) and G2b (M377).

Origins

Various estimated dates and locations have been proposed for the origin of Haplogroup G.

In 2012, a paper by Siiri Rootsi et al. suggested that: "We estimate that the geographic origin of hg G plausibly locates somewhere nearby eastern Anatolia, Armenia or western Iran." [2]

Previously the National Geographic Society places its origins in the Middle East 30,000 years ago and presumes that people carrying the haplogroup took part in the spread of the Neolithic.[3]

Two scholarly papers have also suggested an origin in the Middle East, while differing on the date. Semino et al. (2000) suggested 17,000 years ago.[4] Cinnioglu et al. (2004) suggested the mutation took place only 9,500 years ago.[5]

The only known example of the basal paragroup G(xG1,G2) (G-M210*) is a blood sample said to have come from a man executed on a guillotine during the French Revolution.[6] (See below for further details.)

Structure

G* (M201)

(Subclades here conform to the Y-DNA SNP definitions used by ISOGG In 2012, several categories found only in one man in research studies were removed from the ISOGG tree causing some renaming.)

Geographic distribution

Prehistoric presence

Haplogroup G2a (G-P15) has been identified in neolithic human remains in Europe dating between 5000-3000 BC.

Furthermore, the majority of all the male skeletons from the European Neolithic period have so far yielded Y-DNA belonging to this haplogroup. The oldest skeletons confirmed by ancient DNA testing as carrying haplogroup G2a were five found in the Avellaner cave burial site, near Les Planes d'Hostoles, in Catalonia, Spain and were dated by radiocarbon dating to about 5000 BCE.[7]

A skeleton found at the Neolithic cemetery known as Derenburg Meerenstieg II, in Saxony-Anhalt Germany, apparently belonged to G2a3 (G-S126) or a subclade. It was found with burial artifacts belonging to the Linearbandkeramische Kultur ("Linear Band Ceramic Culture"; LBK). This skeleton could not be dated by radiocarbon dating, but other skeletons there were dated to between 5,100 and 6,100 years old. The most detailed SNP mutation identified was S126 (L30), which defines G2a3.[8]

G2a was found also in 20 out of 22 samples of ancient Y-DNA from Treilles, the type-site of a Late Neolithic group of farmers in the South of France, dated to about 5000 years ago.[9] The fourth site also from the same period is the Ötztal of the Italian Alps where the mummified remains of Ötzi the Iceman were discovered. The Iceman belongs to haplogroup G2a2b [10] (earlier called G2a4).

Haplogroup G2a2b is a rare group today in Europe. The authors of the Spanish study indicated that the Avellaner men had rare marker values in testing of their short tandem repeat (STR) markers.

Historic presence

Two men found in a high-status 7th Century burial at Ergolding in Bavaria, Germany,[11] were found to belong to haplogroup G2a (P15+). The burials have been linked to the Merovingian dynasty.

The only known example of the basal paragroup G(xG1,G2) (G-M210*) was blood found on a handkerchief, sealed in a gourd in France during the 18th Century. The handkerchief reputed to have come from a man who underwent public execution by guillotine during the French Revolution.[6] The blood is said to have belonged to King Louis XVI of France (1754 – 1793), although its provenance is far from certain. Public executions were notoriously chaotic at the time and some of those executed were neither members of the royal family, or even the aristocracy.

Modern presence

In Russia, Ukraine and Central Asia, members of various ethnic minorities and/or residents in particular localities possess G-M201 at its highest levels in the world – even though the average rate at the national level is about 1% or less. The Madjars and Argyn klans of Kazakhstan were found to possess the highest levels of G-M201 among any modern ethnic group. Amongst the Madjars, G1 was found at a rate of 87%. A separate study on the Argyns found that 71% of males belong to G1. In the Russian North Caucasus the Kabardinian and Ossetian populations are also notable for high rates of G-M201. Digora, North Ossetia has the highest known concentration of G in a single city, as 74% of the tested men were G.[12] Haplogroup G is found as far east as northern China in small percentages where G can reach more substantial percentages in minority groups such as the Uyghurs.

In Turkey, the South Caucasus and Iran, haplogroup G reaches the highest percentage of national populations. Among Turkish males 11% of the population is G.[5] In Iran, Haplogroup G reaches 13 to 15% of the population in various parts of the country. While it is found in percentages higher than 10% among the Bakhtiari, Gilaki and Mazandarani, it is closer to 5% among the Iranian Arabs and in some large cities.[13] Among the samples in the YHRD database from the southern Caucasus countries, 29% of the samples from Abazinia, 31% from Georgia, 18% from Azerbaijan and 11% from Armenia appear to be G samples.

In Europe west of the Black Sea, Haplogroup G is found at about 5% of the population on average throughout most of the continent.[14] The concentration of G falls below this average in Scandinavia, the westernmost former Soviet republics and Poland, as well as in Iceland and the British Isles. There are seeming pockets of unusual concentrations within Europe. In Wales, a distinctive G2a3b1 type (DYS388=13 and DYS594=11) dominates there and pushes the G percentage of the population higher than in England.

In the Tirol (Tyrol) of western Austria, the percentage of G-M210 can reach 40% or more; perhaps the most famous example is the ancient remains of the so-called "Iceman", Ötzi. In the northern and highland areas of the island of Sardinia off western Italy, G percentages reach 11% of the population in one study[15] and reached 21% in the town of Tempio in another study. In the Greek island of Crete, approximately 7%[16] to 11%[17] of males belong to haplogroup G. In north-eastern Croatia, in the town of Osijek, G was found in 14% of the males.[18] The city is on the banks of the river Drava, which notably begins in the Tirol/Tyrol region of the Alps, another haplogroup G focus area in Europe. Farther north, 8% of ethnic Hungarian males and 5.1% of ethnic Bohemian (Czech) males have been found to belong to Haplogroup G.

In South Asia, some ethnic minorities possess haplogroup G at concentrations of approximately 18%[19] to 20%[20] of Kalash, approximately 16% of Brahui,[20] and approximately 11.5% of sampled Pashtun,[19] but in only about 3% of the general Pakistani population.[19] The many groups in India and Bangladesh have not been well studied. About 6% of the samples from Sri Lanka and Malaysia were reported as haplogroup G, but none were found in the other coastal lands of the Indian Ocean or Pacific Ocean in Asia.[21]

In the Middle East, haplogroup G accounts for about 3% of the population in almost all areas.[22][23] Among the Druze mostly residents of Israel 10% were found to be haplogroup G.[24]

Around 10% of Jewish males are Haplogroup G.

In Africa, haplogroup G is rarely found in sub-Saharan Africa or south of the horn of Africa among native populations. In Egypt, studies have provided information that pegs the G percentage there to be between 2% and 9%.[25][26][27] 3% of North African Berbers were found to be haplogroup G.[28] 2% of Arab Moroccans and 8% of Berber Moroccans were likewise found to be G.[29]

In the Americas, the percentage of haplogroup G corresponds to the numbers of persons from Old World countries who emigrated. It is not found among Native Americans except where intermarriage with non-native persons has occurred. It has been found in Mexican mestizos.

Major subclades and their distribution

G1 (M285+ or M342+) and its subclades

Main article: Haplogroup G1 (Y-DNA)

Almost all haplogroup G1 persons have the value of 12 at short tandem repeat (STR) marker DYS392 and all will have the M285 or M342 SNP mutation which characterizes this group. This value of 12 is uncommon in other G categories other than G1.

subclades of G1a, G1a1, G1b exist.

The highest reported concentration of G1 and its subclades in a single country is in Iran, with next most frequent concentrations in neighboring countries to the west.

There are distinctive Ashkenazi Jewish and Kazakh subclades based on STR marker value combinations.

G2 (P287+)

Men who belong to this group but are negative for all G2 subclades represent a small number of haplogroup G men. P287 was identified at the University of Arizona and became widely known in late 2007. Its identification caused considerable renaming of G categories.

G2a (P15+)

Haplogroup G men who belong to this group, but are negative for all G2a subclades, are uncommon in Europe but may represent a sizeable group in so far poorly tested areas east of Turkey. P15 was identified at the University of Arizona and became widely known by 2002. Its chromosome location listed as 21653414. G2a was found in medieval remains in a 7th- century CE high-status tomb in Ergolding, Bavaria, Germany, but G2a subclades were not tested.[11]

There are multiple SNPs which so far have the same coverage as P15. They are—with accompanying Y-chromosome locations—U5 (rs2178500), L149 (8486380) and L31 (also called S149) (rs35617575..12538148). Should any man with the P15 mutation test negative (ancestral) for any of these or vice versa, that finding would be the basis of a new G2a category.

G2a1 (L293+) and its subclades

Haplogroup G2a1 and its subclades represent the majority of haplogroup G samples in some parts of the Caucasus Mountains area. They are found only in tiny numbers elsewhere. So far all G2a1 persons have a value of 10 at STR marker DYS392. G2a1a persons also typically have higher values for DYS385b, such as 16, 17 or 18, than seen in most G persons.

The North Ossetians in the mid northern Caucasus area of Russia belong overwhelmingly to the G2a1 subclade based on available samples. The South Ossetians and Svans generally south of North Ossetia have significant number of G2a1 persons, but population percentages have not yet been provided.

The presence of the SNP P18 mutation characterizes G2a1a's only subclade, G2a1a. The reliability of both P16 and P18 in identifying everyone in each of these categories has been questioned and individual components of the SNP have to be examined.

Ashkenazi Jewish G2a1a men with northeastern European ancestry form a distinct cluster based on STR marker values. Men from the Caucasus and men from eastern Europe also form distinctive STR clusters.

G2a2 (L223+) and its subclades

This group is charactized by having the L223 mutation. L223 is found on the Y chromosome at rs13304806 and 6405148 with a mutation from C to G. L223 was first identified in samples at 23andMe in 2009 but proved problematic as an individual test, the first successful results being reported at Family Tree DNA in late 2011 under its assigned L223 label. It was then learned that three subclades as follows belong under L223:

The G2a2a subclade (M286) is tiny. Samples indicating British Isles, Turkish and Lebanese ancestry have been identified. The British samples have inconsistent double values for STR marker DYS19 in many cases. M286 was first identified at Stanford University at chromosome position 21151187, and is a mutation from G to A.

G2a2b (G-L91) was identified in 2009. Its members include "Öetzi", the so-called Iceman, who died at least 5,000 years BP in the European Alps. G2a2b would seem to encompass a significant proportion of men belonging to G. L91 is found so far in scattered parts of Europe and North Africa and in Armenia. Included within G2a2b are some men with double values for STR marker DYS19, but there are also G2a2 men with this finding who are not G2a2b. The double 19 value situation is not seen in the G2a1 and G2a3 subclades. The L91 mutation is found at 21327383 and rs35474563 on the Y-chromosome. The forward primer is GTATTGAACTTACAATTCACGTCCC, and the reverse is CTCTCCAAATCGGGTTTCCT. The mutation involves a change from C to T.[16] L223 is found on the Y chromosome at rs13304806.

The L293 SNP that characterizes the G2a2c subclade was identified in June 2010 at Family Tree DNA. It encompasses a small group of Hispanic men who also so far all have the odd value of 13,21 at the YCA marker. The mutation is found on the Y chromosome at 10595022 and is a change from G to C.

G2a3 (L30+, S126+, U8+)

Men who belong to this group but are negative for all its subclades represent a small number today. This haplogroup was found in a Neolithic skeleton from around 5000 BC, in the cemetery of Derenburg Meerenstieg II, Germany, which forms part of the Linear Pottery culture, known in German as Linearbandkeramik (LBK),[8] but was not tested for G2a3 subclades.

G2a3a (M406+) and its subclades

G2a3a and its several subclades seem most commonly found in Turkey and the coastal areas of the eastern Mediterranean where it can constitute up to 50% of haplogroup G samples. G2a3a is more common in southern Europe than northern Europe. In Europe—except in Italy – G2a3a constitutes less than 20% of G samples. G2a3a so far has seldom surfaced in northern Africa or southern Asia, but represents a small percentage of the G population in the Caucasus Mountains region and in Iran.

A relatively high percentage of G2a3a persons have a value of 21 at STR marker DYS390. The DYS391 marker has mostly a value of 10, but sometimes 11, in G2a3a persons, and DYS392 is almost always 11. If a sample meets the criteria indicated for these three markers, it is likely the sample is G2a3a.

G2a3a has two known subclades. Both are relatively common among G2a3a persons.

G2a3b (L141+)

The SNP that defines this group was identified only in mid-2009 at Family Tree DNA. Almost all L141+ men belong to L141 subclades. Samples from persons with British Isles, Sicilian and Turkish ancestry have been identified. L141+ persons who do not belong to any L141 subclade so far have the value of 11 at STR marker DYS490 — a finding rare in other G categories. The L141 mutation is found on the Y chromosome at 2948607. The L141 mutation involves an insertion.[30]

G2a3b1 (P303+ or S135+) and its subclades

Main article: Haplogroup G-P303

The G2a3b1 definable subclades are heavily concentrated throughout Europe west of the Black Sea and Russia where G2a3b1 is often in the majority among G persons. Small percentages of G2a3b1 are found primarily in the area encompassed by Turkey, the Caucasus countries, Iran and the Middle East where the G2a3b1 SNP may have originated. G2a3b1 is also found in India. The great majority of P303+ men belong to one of its subclades.

The largest G2a3b1 subclade based on available samples is one in which almost all persons have the value of 13 at STR marker DYS388. The L497 SNP (G2a3b1a2) encompasses these men, but most men L497 men belong to its subclade Z725. There are additional subclades of DYS388=13 men characterized by the presence of specific SNPs or uncommon STR marker oddities. Members of this group have been found in Europe and the Middle East.[31]

The next largest G2a3b1 subclade is characterized by the presence of the U1 mutation (G2a3b1a1) But a high percentage of U1+ men belong to its two subclades, L13/S13 (G2a3b1a1a)and Z1266 (G2a3b1a1b). The L13 subclade is most common in north central Europe, and Z1266 is most common in the western Caucasus Mountains.

The final major subclade is characterized by presence of the Z1903 SNP and so far by the value of 9 at marker DYS568. A high percentage of Z1903+ men belong to its subclade, Z724. The Z724 subclade contains a further large subclade consisting overwhelmingly of Ashkenazi Jews.

The highest percentage of G2a3b1 persons in a discrete population so far described is on the island of Ibiza off the eastern Spanish coast. This group has been linked with the Crypto-Jewish population which fled to the island during the time of the Spanish Inquisition, of which a significant portion are identifiable as DYS388=13 persons.[32]

G2a3b2 (L177+)

This G2a3b2 group is certainly smaller in numbers of men included than G2a3b1, but only a small amount of testing has occurred for the L177 mutations. So far the men positive for this have listed Irish, English, Dutch, Lebanese and Turkish (Armenian surname) ancestry. Several L177 subclades based on shared STR marker oddities exist.

The number of STR marker values separating men in this group suggest G2a3b2 is a relatively old group despite the small number of men involved.[33] The mutations involved are complicated and difficult to interpret. The L177.1 component is found at Y chromosome position 23397163; L177.2 at 25030912; L177.3 at 25750264.[34] This SNP was first identified at Family Tree DNA in 2009.

G2b (M377+) and its subclade

Until August, 2012, this subclade was designated G2c. A clade of closely related Ashkenazi Jews represent virtually all G2b persons, with just three other G2b haplotypes having been reported so far: one Turk from Kars in northeast Turkey near Armenia, one Pashtun, and one Burusho in Pakistan.

The extreme rarity of G2b in northern Pakistan could indicate that G2b in this area originates outside the region and was brought there in the historic period, perhaps from further west (Pakistan was part of both the Achaemenid Persian Empire, conquered by Alexander the Great, and then formed a part of the Greco-Bactrian Kingdom). These two reported Pakistani G2c haplotypes are quite divergent from the Ashkenazi Jewish clade, and therefore do not at all indicate a recent common origin. The Turkish G2b is somewhat closer, but not identical. It remains to be seen if testing will reveal G2b haplotypes in other populations — this is some indication that G2b occurs at low levels in the Near East. All G2b men tested so far have a rare null value for the DYS425 marker, (a missing "T" allele of the DYS371 palindromic STR), the result of a RecLOH event, a finding not yet seen among most other G haplotypes. Among Jews in Israel drawn from many areas of the world, G2b constituted 3.7% in one study.[35]

Haplogroup G2b has been found at a frequency of 60% out of a sample of five Pashtuns in the Wardak region of Afghanistan. This is likely due to a local founder effect.[36]

Defining SNPs

The International Society of Genetic Genealogy (ISOGG) maintains the most up-to-date consensus version of haplogroup categories.[37] These classifications are based on shared SNP mutations. The discovery of new SNPs can result in assignment of new names to haplogroup categories. There were only a few G categories until 2008 when major revisions to categories were made. Even more G SNPs were identified in 2009 to 2012 leading to more changes. Until 2008, new G SNPs were reported from labs at the University of Arizona (P designations), Stanford University (M designations) or the University of Central Florida (U designations). Beginning in 2008, additional G SNPs were identified at Family Tree DNA (L designations) and Ethnoancestry (S designations). These latter labs also made use of raw data results reported by individuals tested for about 2,000 SNPs at 23andMe to provide new L or S-designated SNP tests. In 2009-10, Family Tree DNA's Walk through the Y Project, sequencing certain Y-chromosome segments, provided a number of new G SNPs with the L designation. In 2012, SNPs with the Z designation as first identified by citizen researchers from 1000 Genomes Project data began to appear.

Because SNPs provide the most reliable method of categorization, each is allowed to represent an official G category. Categories have alternating letters and numbers. But unusual values or unusual value combinations found at short tandem repeat markers (STRs) can also provide the basis of additional taxonomisation. The identification of a new SNP can necessitate renaming of one or more categories.

The M201 SNP mutation that characterizes haplogroup G was identified at Stanford University and was first reported in 2001.[38] The technical specifications of M201 are given as: refSNPid is rs2032636.....Y chromosome location of 13536923....forward primer is tatgcatttgttgagtatatgtc.....reverse primer is gttctgaatgaaagttcaaacg.....the mutation involves a change from G to T.

A number of SNPs have been identified with seemingly the same coverage in the population as M201. Because M201 was identified first, it is the standard SNP test used when testing for G persons. In order to determine if one of these alternative SNPs represents a subclade of M201, the alternative SNPs must be tested in G persons who are negative for the known subclades of G. There are only a tiny number of persons in such a category, and only a tiny number of persons have been tested for G equivalent SNPs other than M201.

The following SNPs are so far identified as M201 equivents: L116, L154, L269, L294, L240, P257, L402, L520, L521, L522, L523, L605, Page 94, U2, U3, U6, U7, U12, U17, U20, U21, U23 and U33. P257 was first reported in 2008.[39] L240 was identified in 2009.[40] The "U" SNPs were identified in 2006 but not published until 2009.[41]

In addition, there are multiple other SNPs thought to have the same coverage as M201. These are found at: rs9786910, rs9786537, rs2713254, rs35567891 and rs34621155 on the Y chromosome. No labs have yet assigned them shorthand names.

Prominent members of G-M201

The man known as "Ötzi" – whose remains, dating from the 4th millennium BCE, were found on the modern border between Austria and Italy – is a member of G-L91 (G2a2b).

Recently, the English Plantagenet king, Richard III, has been forensically confirmed and verified as a bearer of a sub-variant of G-M201.[42]

Joseph Stalin, from a genetic test on his grandson (his son Vasily's son; Alexander Burdonsky), shows his Y-DNA haplogroup to be G2a1a:[43]

DYS 393 390 19 391 385A 385B 426 388 439 389I 392 389II 458 459A 459B 455 454 447 437 448 449 464A 464B 464C 464D
Alleles 14 23 15 9 15 16 11 12 11 11 10 28 17 9 9 11 11 25 16 21 28 13 13 14 14

Other people purportedly members of Haplogroup G include: King Louis XVI of France,[6] German-American pioneer and soldier Phillip Hamman, politician Linn Banks, physicist John G. Cramer, actor James Franciscus, lawyer and administrator Newton Minow, and male members of the Swedish Bure kinship (G2a3b1a2-Y12970).

See also

References

  1. ↑ Learn about Y-DNA Haplogroup G Genebase.com
  2. ↑ S. Rootsi et al. 2012, "Distinguishing the co-ancestries of haplogroup G Y-chromosomes in the populations of Europe and the Caucasus", European Journal of Human Genetics, v. 20, no. 11 (Dec.), pp. 1275-82.
  3. ↑ Atlas of the Human Journey: Haplogroup G (M201)
  4. ↑ 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 (November 2000). "The genetic legacy of Paleolithic Homo sapiens sapiens in extant Europeans: a Y chromosome perspective". Science. 290 (5494): 1155–9. Bibcode:2000Sci...290.1155S. doi:10.1126/science.290.5494.1155. PMID 11073453.
  5. 1 2 Cinnioğlu C, et al. (January 2004). "Excavating Y-chromosome haplotype strata in Anatolia". Human Genetics. 114 (2): 127–48. doi:10.1007/s00439-003-1031-4. PMID 14586639.
  6. 1 2 3 Maarten H. D. Larmuseau et al., 2014, "Genetic genealogy reveals true Y haplogroup of House of Bourbon contradicting recent identification of the presumed remains of two French Kings", European Journal of Human Genetics, vol. 22, no. 5 (May), pp. 681–687.
  7. ↑ Lacan M, Keyser C, Ricaut FX, Brucato N, Tarrús J, Bosch A, Guilaine J, Crubézy E, Ludes B (November 2011). "Ancient DNA suggests the leading role played by men in the Neolithic disseminationtolia". Proc Natl Acad Sci USA. 108 (45): 18255–9. Bibcode:2011PNAS..10818255L. doi:10.1073/pnas.1113061108. PMC 3215063Freely accessible. PMID 22042855.
  8. 1 2 Haak W, Balanovsky O, Sanchez JJ, Koshel S, Zaporozhchenko V, Adler CJ, Der Sarkissian CS, Brandt G, Schwarz C, Nicklisch N, Dresely V, Fritsch B, Balanovska E, Villems R, Meller H, Alt KW, Cooper A (2010). Penny D, ed. "Ancient DNA from European Early Neolithic Farmers Reveals Their Near Eastern Affinities". PLoS Biology. 8 (11): e1000536. doi:10.1371/journal.pbio.1000536. PMC 2976717Freely accessible. PMID 21085689.
  9. ↑ Marie Lacan, Christine Keyser, François-Xavier Ricaut, Nicolas Brucato, Francis Duranthon, Jean Guilaine, Eric Crubézy, and Bertrand Ludes, Ancient DNA reveals male diffusion through the Neolithic Mediterranean route, Proceedings of the National Academy of Sciences of the USA, online May 31, 2011 before print.
  10. ↑ Interview Dr. Eduard Egarter-Vigl, Head of Conservation and Assistant to research projects of the Archaeological Museum in Bozen. From the Docu-Movie: "Ötzi, ein Archäologiekrimi" [Ötzi, a Archaeology Crime] by Christine Sprachmann. TV-Broadcast by 3sat 10 August 2011 and br-alpha 13 September 2011
  11. 1 2 Vanek D, Saskova L, Koch H (2009). "Kinship and Y-Chromosome Analysis of 7th Century Human Remains: Novel DNA Extraction and Typing Procedure for Ancient Material". Croatian Medical Journal. 50 (3): 286–95. doi:10.3325/cmj.2009.50.286. PMC 2702742Freely accessible. PMID 19480023.
  12. ↑ Nasidze I, Sarkisian T, Kerimov A, Stoneking M (2003). "Testing Hypotheses of Language Replacement in the Caucasus: Evidence from the Y Chromosome" (PDF). Human Genetics. 112 (3): 255–61. doi:10.1007/s00439-002-0874-4 (inactive 2015-01-13). PMID 12596050.
  13. ↑ Nasidze I, et al. (2008). "Close Genetic Relationship Between Semitic-speaking and Indo-European-speaking Groups in Iran". Annals of Human Genetics. 72 (Pt 2): 241–52. doi:10.1111/j.1469-1809.2007.00413.x. PMID 18205892.
  14. ↑ "G-Z7941 - Haplogroup G Project". google.com.
  15. ↑ Zei G, et al. (2003). "From surnames to the history of Y chromosomes: the Sardinian population as a paradigm". European Journal of Human Genetics. 11 (10): 802–07. doi:10.1038/sj.ejhg.5201040. PMID 14512971.
  16. ↑ Martinez L, et al. (April 2007). "Paleolithic Y-haplogroup heritage predominates in a Cretan highland plateau". European Journal of Human Genetics. 15 (4): 485–93. doi:10.1038/sj.ejhg.5201769. PMID 17264870.
  17. ↑ King RJ, et al. (March 2008). "Differential Y-chromosome Anatolian influences on the Greek and Cretan Neolithic". Annals of Humam Genetics. 72 (Pt 2): 205–14. doi:10.1111/j.1469-1809.2007.00414.x. PMID 18269686.
  18. ↑ Battaglia V, et al. (2009). "Y-chromosomal evidence of the cultural diffusion of agriculture in southeast Europe". European Journal of Human Genetics. 17 (6): 820–30. doi:10.1038/ejhg.2008.249. PMC 2947100Freely accessible. PMID 19107149.
  19. 1 2 3 Firasat S, Khaliq S, Mohyuddin A, Papaioannou M, Tyler-Smith C, Underhill PA, Ayub Q (2006). "Y Chromosomal Evidence for a Limited Greek Contribution to the Pathan Population of Pakistan". European Journal of Human Genetics. 15 (1): 121–26. doi:10.1038/sj.ejhg.5201726. PMC 2588664Freely accessible. PMID 17047675.
  20. 1 2 Sengupta S, 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". American Journal of Human Genetics. 78 (2): 202–21. doi:10.1086/499411. PMC 1380230Freely accessible. PMID 16400607.
  21. ↑ Hammer MF, et al. (2006). "Dual Origins of the Japanese: Common Ground for Hunter-Gatherer and Farmer Y-Chromosomes". Journal of Human Genetics. 51 (1): 47–58. doi:10.1007/s10038-005-0322-0. PMID 16328082.
  22. ↑ Flores C, et al. (2005). "Isolates in a corridor of migrations: a high-resolution analysis of Y-chromosome variation in Jordan" (PDF). Journal of Human Genetics. 50 (9): 435–41. doi:10.1007/s10038-005-0274-4. PMID 16142507.
  23. ↑ Cadenas AM, et al. (2008). "Chromosome Diversity Characterizes the Gulf of Oman". European Journal of Human Genetics. 16 (3): 374–86. doi:10.1038/sj.ejhg.5201934. PMID 17928816.
  24. ↑ Shlush LI, et al. (2008). Gemmell NJ, ed. "The Druze: A Population Genetic Refugium of the Near East". PLoS ONE. 3 (5): e2105. Bibcode:2008PLoSO...3.2105S. doi:10.1371/journal.pone.0002105. PMC 2324201Freely accessible. PMID 18461126.
  25. ↑ Elizabeth T Wood, Daryn A Stover, Christopher Ehret et al., "Contrasting patterns of Y chromosome and mtDNA variation in Africa: evidence for sex-biased demographic processes," European Journal of Human Genetics (2005) 13, 867–876. (cf. Appendix A: Y Chromosome Haplotype Frequencies)
  26. ↑ Luis JR, et al. (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.
  27. ↑ El-Sibai M, Platt DE, Haber M, Xue Y, Youhanna SC, Wells RS, Izaabel H, Sanyoura MF, Harmanani H, Bonab MA, Behbehani J, Hashwa F, Tyler-Smith C, Zalloua PA (2009). "Geographical Structure of the Y-Chromosomal Genetic Landscape of the Levant: A Coastal-Inland Contrast". Annals of Human Genetics. 73 (Pt6): 561–81. doi:10.1111/j.1469-1809.2009.00538.x. PMC 3312577Freely accessible. PMID 19686289.
  28. ↑ Alonso S, Flores C, Cabrera V, Alonso A, Martín P, Albarrán C, Izagirre N, de la Rúa C, García O (2005). "The place of the Basques in the European Y-chromosome diversity landscape" (PDF). European Journal of Human Genetics. 13 (12): 1293–1302. doi:10.1038/sj.ejhg.5201482. PMID 16094307.
  29. ↑ Cruciani F, et al. (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): 1197–1214. doi:10.1086/340257. PMC 447595Freely accessible. PMID 11910562.
  30. ↑ http://ymap.ftdna.com/cgi-bin/gb2/gbrowse_details/hs_chrY?ref=ChrY;start=2948607;end=2948607;name=L141;class=Sequence;feature_id=41210;db_id=ymap%3Adatabase
  31. ↑ Rootsi S, Myres NM, Lin AA, et al. (16 May 2012). "Distinguishing the co-ancestries of haplogroup G Y-chromosomes in the populations of Europe and the Caucasus". European Journal of Human Genetics. 20: 1275–1282. doi:10.1038/ejhg.2012.86. PMID 22588667. Retrieved 4 January 2015.
  32. ↑ Adams SM, Bosch E, Balaresque PL, Ballereau SJ, Lee AC, Arroyo E, López-Parra AM, Aler M, Grifo MS, Brion M, Carracedo A, Lavinha J, Martínez-Jarreta B, Quintana-Murci L, Picornell A, Ramon M, Skorecki K, Behar DM, Calafell F, Jobling MA (December 2008). "The genetic legacy of religious diversity and intolerance: paternal lineages of Christians, Jews, and Muslims in the Iberian Peninsula". American Journal of Human Genetics. 83: 725–36. doi:10.1016/j.ajhg.2008.11.007. PMC 2668061Freely accessible. PMID 19061982.
  33. ↑ "..Project Rosters - Haplogroup G Project". google.com.
  34. ↑ http://ytree.ftdna.com/index.php?name=Draft&parent=20173662
  35. ↑ Hammer MF, et al. (2009). "Extended Y chromosome haplotypes resolve multiple and unique lineages of the Jewish priesthood". Human Genetics. 126 (5): 707–17. doi:10.1007/s00439-009-0727-5. PMC 2771134Freely accessible. PMID 19669163.
  36. ↑ Habe, M; et al. (March 28, 2012). "Afghanistan's Ethnic Groups Share a Y-Chromosomal Heritage Structured by Historical Events". PLoS One. 7 (3): e34288. doi:10.1371/journal.pone.0034288. PMC 3314501Freely accessible.
  37. ↑ Copyright 2015 ISOGG. "ISOGG 2015 Y-DNA Haplogroup Tree". isogg.org.
  38. ↑ Underhill PA, Passarino G, Lin AA, Shen P, Mirazón Lahr M, Foley RA, Oefner PJ, Cavalli-Sforza LL (2001). "The phylogeography of Y chromosome binary haplotypes and the origins of modern human populations". Annals of Human Genetics. 65 (Pt1): 43–62. doi:10.1046/j.1469-1809.2001.6510043.x. PMID 11415522.
  39. ↑ Karafet TM, Mendez FL, Meilerman MB, Underhill PA, Zegura SL, Hammer MF (2008). "New binary polymorphisms reshape and increase resolution of the human Y chromosomal haplogroup tree". Genome Research. 18 (5): 830–38. doi:10.1101/gr.7172008. PMC 2336805Freely accessible. PMID 18385274.
  40. ↑ http://ymap.ftdna.com/cgi-bin/gbrowse_details/hs_chrY?name=L240;class=Sequence;ref=ChrY;start=3191153;end=3191153;feature_id=40369
  41. ↑ Sims LM, et al. (2009). Batzer MA, ed. "Improved Resolution Haplogroup G Phylogeny in the Y Chromosome, Revealed by a Set of Newly Characterized SNPs". PLoS ONE. 4 (6): 1–5. Bibcode:2009PLoSO...4.5792S. doi:10.1371/journal.pone.0005792. PMC 2686153Freely accessible. PMID 19495413.
  42. ↑ King TE, Fortes GG, Balaresque P, Thomas MG, Balding D, Maisano Delser P, Neumann R, Parson W, Knapp M, Walsh S, Tonasso L, Holt J, Kayser M, Appleby J, Forster P, Ekserdjian D, Hofreiter M, Schürer K (2014). "Identification of the remains of King Richard III". Nat Commun. 2 (5): 5631. doi:10.1038/ncomms6631. PMC 4268703Freely accessible. PMID 25463651.
  43. ↑
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.
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