Haplogroup J1 | |
---|---|
Possible time of origin | 10,000-36,000 years before present[1] |
Possible place of origin | Western Asia |
Ancestor | J |
Descendants | J1a, J1b, J1c |
Defining mutations | M267 |
Highest frequencies | Semitic populations generally, and also the Caucasus (especially Dagestan) |
In human genetics, Y DNA haplogroup J1, also known as J-M267, is a sub-haplogroup of Haplogroup J, along with its sibling clade Haplogroup J2. It is one of the most commonly shared ancient paternal lineages found amongst men in many parts of North Africa, the Middle East, the Caucasus, and the Horn of Africa. It is also found in Europe and in significant frequencies it is found as far east as the Indian subcontinent and Central Asia.
This type of Y DNA is defined by the presence of the SNP mutation referred to as M267, since its discovery was announced in Cinnioğlu et al. (2004).
Subclades and proposed origins
editJ1 has several recognized sub-clades, some of which were recognized before J1 itself was recognized, for example J-M368.[2] But most of these are not common.[1]
The P58 marker which defines subgroup J1c3 is common, and was first announced in Karafet et al. (2008), but had been announced earlier under the name "Page08" in 2006 in Repping, Van Daalen & Brown (2006). This haplogroup dominates haplogroup J, and is notable as containing the Jewish "Cohen modal haplotype", as well as both the so-called "Galilee modal haplotype" and "P&I Arab modal haplotype" associated with Palestinians and Israeli Arabs by Nebel et al. (2000) .[3]
More generally, the J-P58 group has been shown to be closely associated with a large cluster of J1 which had been recognized before the discovery of P58, and which has been associated with Middle Eastern Semitic migrations including Arabs and Jews. This cluster was identified by STR markers - specifically YCAII as 22-22, and DYS388 having unusual repeat values of 15 or higher, instead of 13.[4] Apart from the Jewish "Cohen" haplotype, Semino et al. (2000) associated this DNA profile with the Arab expansion in the seventh century AD, and noted that it was most frequent amoungst J1 men in the Middle East and North Africa, but less frequent in Ethiopia and Europe. Other authors have claimed that the evidence points to older migrations being possibly more important for explaining the distribution of modern J1.
Similarly, Tofanelli et al. (2009) refer to an "Arabic" and a "Eurasian" type of J1. The Arabic type includes Arabic speakers from Maghreb, Sudan, Iraq and Qatar, and it is a relatively homogeneous group. This is the group with YCAII=22-22 and high DYS388 values. The more diverse "Eurasian" group includes Europeans, Kurds, Iranians and Ethiopians (Ethiopia being outside of Eurasia), and is much more diverse. The authors also say that "Omanis show a mix of Eurasian pool-like and typical Arabic haplotypes as expected, considering the role of corridor played at different times by the Gulf of Oman in the dispersal of Asian and East African genes."
A similar geographical pattern is observed by Chiaroni et al. (2010) looking at J-P58. The authors report that "in Ethiopia, all Cushitic Oromo and ∼29% of Semitic Amharic J1 chromosomes are J1*" (meaning that they are J1 but not in J-P58). And furthermore, both the Caucasus region and the nearby eastern regions of Turkey have also been described by researchers as showing the highest frequency of J1 men who are not in the J-P58 sub-clade.[5][4]
The correspondence between P58 and high DYS388 values, and YCAII=22-22 is not perfect. The J-M368 sub-clade of J-P58 has DYS388=13 and YCAII=19-22, like other types of J1 outside the "Arabic" type of J1, and it is therefore believed to be a relatively old offshoot of J-P58, that did not take part in the most recent waves of J1 expansion in the Middle East.[4] These DYS388=13 haplotypes are most common in the Caucasus and Anatolia, but also found in Ethiopia.[1]
In summary, the above mentioned studies agree upon J1 and J-P58 being male lineages which have expanded more than once within the Middle East and neighbouring regions, with J-P58, and especially part of J-P58, having been carried in the more recent wave of expansion that has become dominant amongst Arabic speakers. Concerning both J1 generally and J-P58 specifically within it, the above surveys all agree that oldest diversity appears to be found today in the Middle East, but not in the most central Arabic areas where the YCAII=22-22 type has become most common, but rather to the north and south of that range. Most recently, concerning J-P58, Chiaroni et al. (2010) propose that the most likely point of original expansion is the north of the fertile crescent in the Taurus and Zagros mountains, during the Neolithic. They propose that the later waves of J-P58 expansion may have been associated with pastoralism and semitic languages. With their method of age estimation, they agree with Tofanelli et al. (2009) in expressing doubt that the Islamic expansions of the 7th century are old enough to explain the most obvious patterns of J1 frequencies within the Middle East.
On the other hand both these authors and others note that in the far south of the J1 range, and the J-P58 range, in Oman, Yemen and Ethiopia, there are also signs of J1 and J-P58 being much older than in the Arabian desert. Chiaroni et al. (2010) attribute this to "either sampling variability and/or demographic complexity associated with multiple founders and multiple migrations".
Tree
editThis phylogenetic tree of haplogroup subclades is based on the ISOGG (2011) tree, which is in turn based upon the YCC 2008 tree[6] and subsequent published research.
- J1 (L255, L321, M267) Typical of populations of the Arabian peninsula, Dagestan, Mesopotamia, the Levant and Semitic-speaking populations of North Africa and Northeast Africa, with a moderate distribution throughout Western Asia'
- J1* -
- J1a (M62) Found at a low frequency in Britain
- J1b (M365.1)
- J1c (L136)
Distribution
edit300px|rightt|thumb|J1 distribution
J1 (or J-M267) is a typical Y chromosome of populations of the Arabian peninsula, Dagestan, Mesopotamia, the Levant and Semitic-speaking populations of North Africa and Northeast Africa, and further distribution throughout Western Asia and other neighbouring areas.
The frequency of Haplogroup J1 is particularly high in Semitic-speaking populations (often much higher in those populations than in neighbouring ones). For comparison with non-semitic Middle Eastern populations only about 10% of men in Iran and Turkey are J1, compared to much high frequencies elsewhere.[7][8]
In the Caucasus and Eastern Turkey, J1 is common in several areas, and particularly common in Daghestan. It is also distinctive. While Middle Eastern J1 tends to be dominated by one sub-clade of J1, called J-P58 or J1c3. Both the Caucusus region and the nearby eastern regions of Turkey have been described by researchers as showing the highest frequencies of J1 which is not in the J-P58 sub-clade.[5][4]
The distribution of J1 outside of the Middle East and Caucasus may be associated with Semites who traded, migrated or conquered in various periods, to such areas as Sicily, southern Italy, Tunisia, Spain, Khartoum, Ethiopia, and Pakistan.[citation needed]
Arabian Plate
editHaplogroup J1, defined by the M267 marker is the most common Y haplogroup in Yemen(80%), Qatar (58%).[9] and Saudi Arabia (64%=68/106).[10]
J1 is generally frequent amongst Negev Bedouins (62%=21/31)[11]. It is also very common among other Arabs such as those of the Levant, i.e. Palestinian (38.4%).[12]
Caucasus
editHaplogroup J1 is the most frequent Y haplogroup in Dagestan among Kubachi (99%), Kaitak (85%), Avars (58%), Dargins (69%), Lezgins (44%) and Chechens (21%).[5]
North Africa and Horn Of Africa
editIn North Africa, J1 is found at the highest rates among the Sudanese of Khartoum (74%) [4]. The Haplogroup's frequency rates among the Sudanese Arabs is (45%),Nubians (41%), Copts is (39%), Beja is (36%), and present with lower frequency in the region of Darfur: Masalit (6%), Fur (6%).[13] J1 is also found with high frequency among Algerians (15-35%) and Tunisians (15-34%),where as in Morocco it is (20%).Haplogroup J1 may be found in as many as 20% of Egyptian males,[14] with the frequency of this haplogroup tending to be comparatively high in the south of the country.[15] J1 is also Observed among East Africans Amhara 36% , Arsi Ethiopians 26%, Oromo Ethiopians 2.56%.[16]
Europe
editIn general J1 has a very low frequency in Europe. However, higher frequencies has been reported in the central Adriatic regions of Italy Gargano (17.2%),[17] Pescara (15%),[17] in the Mediterranean Paola (11.1%),[17], Crete (8.3%),[18] Malta (7.8%), South Portugal (7%)[19], Cyprus (6.2%),[20] Greece (5.3%).[18] and Sicily(3.8%).[21]
Summary
editThe following gives a summary of most of the studies which specifically tested for M267, showing where its distribution is greater than 1%. {{citation}}
: Empty citation (help)
Country or region | Sampling | N | J1 | Source |
Albania | 56 | 3.6 | Semino et al. (2004) | |
Algeria | 20 | 35 | Semino et al. (2004) | |
Algeria | Oran | 102 | 22.5 | Robino et al. (2008) |
Caucasus | 1525 | 23.2 | Balanovsky et al. (2011) | |
Caucasus | Avars | 115 | 58.3 | Balanovsky et al. (2011) |
Caucasus | Dargins | 101 | 69.3 | Balanovsky et al. (2011) |
Caucasus | Kubachi | 65 | 98.5 | Balanovsky et al. (2011) |
Caucasus | Kaitak | 33 | 84.8 | Balanovsky et al. (2011) |
Caucasus | Lezghins | 81 | 44.4 | Balanovsky et al. (2011) |
Caucasus | Chechens | 330 | 20.9 | Balanovsky et al. (2011) |
Caucasus | Circassians | 142 | 4.9 | Balanovsky et al. (2011) |
Caucasus | Ingush | 143 | 2.8 | Balanovsky et al. (2011) |
Caucasus | Ossets | 357 | 2.2 | Balanovsky et al. (2011) |
Central Asia | 184 | 9.7 | Semino et al. (2004) | |
Cyprus | 164 | 12.9 | El-Sibai et al. (2009)[22] | |
Egypt | 147 | 19.7 | Flores et al. (2005) | |
Egypt | Western Desert (el-Hayez) | 35 | 31.4 | Kujanová et al. (2009) |
Ethiopia | Amhara | 48 | 33.3 | Semino et al. (2004) |
Ethiopia | Oromo | 78 | 2.6 | Semino et al. (2004) |
Europe | Ashkenazim Jewish | 442 | 19 | Behar et al. (2004) |
Europe | Ashkenazim Jewish | 82 | 14.6 | Semino et al. (2004) |
Europe | Sephardim Jewish | 42 | 11.9 | Semino et al. (2004) |
Georgia | 45 | 6.6 | Semino et al. (2004) | |
Greece | 92 | 2.2 | Semino et al. (2004) | |
Greece | 442 | 2.5 | Flores et al. (2005) | |
Greece | Nea Nikomedeia | 57 | 10.5 | King et al. (2008) |
Greece | Sesklo/Dimini | 57 | 3.5 | King et al. (2008) |
Greece | Lerna/Franchthi | 57 | 1.8 | King et al. (2008) |
Greece | Crete | 193 | 8.3 | King et al. (2008) |
Greece | Macedonia | 56 | 1.8 | Semino et al. (2004) |
Iberia | 655 | 2.1 | Fregel et al. (2009) | |
Iran | 318 | 13.8 | Flores et al. (2005) | |
Iraq | 156 | 28.2 | Semino et al. (2004) | |
Iraq | 203 | 31 | Flores et al. (2005) | |
Israel | Palestinians | 143 | 38.4 | Semino et al. (2004) |
Israel | Bedouin | 32 | 62.5 | Semino et al. (2004) |
Italy | Calabria | 57 | 1.8 | Semino et al. (2004) |
Italy | Apulia | 86 | 2.3 | Semino et al. (2004) |
Italy | Sicily | 42 | 7.1 | Semino et al. (2004) |
Italy | Sicily | 236 | 3.8 | Gaetano al. (2008) |
Italy | Sardinia | 144 | 2.8 | Semino et al. (2004) |
Jordania | 101 | 40.6 | Flores et al. (2005) | |
Jordania | 273 | 31.9 | El-Sibai et al. (2009) | |
Kuwait | 42 | 33.3 | El-Sibai et al. (2009) | |
Lebanon | 40 | 12.5 | Semino et al. (2004) | |
Lebanon | 914 | 20.1 | Zalloua et al. (2008) | |
Lebanon | 104 | 16.3 | Flores et al. (2005) | |
Malta | 90 | 7.8 | El-Sibai et al. (2009) | |
Morocco | Arab | 49 | 10.2 | Semino et al. (2004) |
Morocco | Arab | 44 | 13.6 | Semino et al. (2004) |
Morocco | Berber | 64 | 6.3 | Semino et al. (2004) |
Morocco | Berber | 103 | 7.8 | Semino et al. (2004) |
Morocco | Residents in Italy | 51 | 19.6 | Onofri et al. (2008) |
Morocco | Bosh et al 2001 | 221 | 5 | Fregel et al. (2009) |
North Africa | Saharawish | 29 | 17.2 | Semino et al. (2004) |
North Africa | Saharawish | 89 | 20.3 | Fregel et al. (2009) |
North Africa | Algeria, Tunisia | 202 | 29.2 | Fregel et al. (2009) |
Oman | 121 | 37.2 | Flores et al. (2005) | |
Pakistan | Hunza | 38 | 2.6 | Semino et al. (2004) |
Pakistan-India | 88 | 7.9 | Semino et al. (2004) | |
Pakistan | 718 | 4.9 | Flores et al. (2005) | |
Portugal | North, Center, South | 303 | 4.3 | Goncalves et al. (2005) |
Portugal | North | 101 | 1 | Goncalves et al. (2005) |
Portugal | Center | 102 | 4.9 | Goncalves et al. (2005) |
Portugal | South | 100 | 7 | Goncalves et al. (2005) |
Portugal | Tras-os-Montes (Jews) | 57 | 12.3 | Nogueiro et al. (2010) |
Qatar | 72 | 58.3 | Cadenas et al. 2008 | |
Saudi Arabia | 106 | 64.2 | Alshamaly et al. 2009 | |
Saudi Arabia | 157 | 40.1 | Abu-Amero et al. 2009 | |
Somali | 201 | 2.5 | Flores et al. (2005) | |
Spain | Andalusia | 93 | 1.1 | Semino et al. (2004) |
Spain | Canary Islands | 652 | 3.53 | Fregel et al. (2009) |
Sudan | Khartoum | 35 | 74 | Chiaroni el al. (2010)[4] |
Syria | 111 | 32.4 | Flores et al. (2005) | |
Syria | 554 | 33.8 | El-Sibai et al. (2009) | |
Tunisia | 73 | 30.1 | Semino et al. (2004) | |
Tunisia | 52 | 34.6 | Onofri et al. (2008) | |
Turkey | Muslim Kurd | 95 | 11.6 | Semino et al. (2004) |
Turkey | Muslim Kurd | 251 | 11.2 | Flores et al. (2005) |
Turkey | Istanbul | 73 | 6.9 | Semino et al. (2004) |
Turkey | Konya | 129 | 3.9 | Semino et al. (2004) |
Turkey | 523 | 9.2 | Flores et al. (2005) | |
UAE | 164 | 35 | Abu-Amero et al. 2009 | |
Yemen | 104 | 80.8 | Alshamaly et al. 2009 | |
Yemen | 62 | 72.7 | Cadenas et al. 2008 |
References
edit- ^ a b c Tofanelli et al. (2009)
- ^ Y Chromosome Consortium "YCC" (2002)
- ^ Hammer et al. (2009)
- ^ a b c d e f Chiaroni et al. (2011)
- ^ a b c Balanovsky et al. (2011)
- ^ Karafet et al. (2008)
- ^ Cinnioğlu et al. (2004)
- ^ Regueiro et al. (2006)
- ^ Cadenas AM, Zhivotovsky LA, Cavalli-Sforza LL, Underhill PA, Herrera RJ (2008). "Y-chromosome diversity characterizes the Gulf of Oman". Eur. J. Hum. Genet. 16 (3): 374–86. doi:10.1038/sj.ejhg.5201934. PMID 17928816.
{{cite journal}}
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ignored (help)CS1 maint: multiple names: authors list (link) Yemen 45/62 = 72.6% J1-M267 Qatar 42/72 = 58.3% J1-M267 - ^ Alshamaly et al. 2009
- ^ Nebel et al. 2001
- ^ Semino et al. (2004)
- ^ Hisham Y. Hassan et al., "Y-Chromosome Variation Among Sudanese: Restricted Gene Flow, Concordance With Language, Geography, and History," American Journal of Physical Anthropology (2008). J-12f2(xJ2-M172) in 46/102 Sudanese Arabs of the Gaalien, Meseria, and Arakien tribes, 16/39 Nubians, 13/33 Copts, 15/42 Beja, 2/32 Masalit, and 2/32 Fur.
- ^ Luis JR, Rowold DJ, Regueiro M; et al. (2004). "The Levant versus the Horn of Africa: evidence for bidirectional corridors of human migrations". Am. J. Hum. Genet. 74 (3): 532–44. doi:10.1086/382286. PMC 1182266. PMID 14973781.
{{cite journal}}
: Explicit use of et al. in:|author=
(help); Unknown parameter|month=
ignored (help)CS1 maint: multiple names: authors list (link) - ^ Barbara Arredi, Estella S. Poloni, Silvia Paracchini, Tatiana Zerjal, Dahmani M. Fathallah, Mohamed Makrelouf, Vincenzo L. Pascali, Andrea Novelletto and Chris Tyler-Smith, "A Predominantly Neolithic Origin for Y-Chromosomal DNA Variation in North Africa," American Journal of Human Genetics, Volume 75, Issue 2, August 2004, Pages 338-345. Haplogroup J-12f2(xJ2-M172) in 4/44 = 9.1% of a sample of Egyptians from the vicinity of Mansoura in northern Egypt, and 6/29 = 40.7% of a sample of Egyptians from the vicinity of Luxor in southern Egypt.
- ^ http://www.nature.com/ejhg/journal/v18/n3/abs/ejhg2009166a.html
- ^ a b c http://www.familytreedna.com/pdf/italy.pdf
- ^ a b King RJ, Ozcan SS, Carter T; et al. (2008). "Differential Y-chromosome Anatolian influences on the Greek and Cretan Neolithic". Ann. Hum. Genet. 72 (Pt 2): 205–14. doi:10.1111/j.1469-1809.2007.00414.x. PMID 18269686.
{{cite journal}}
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ignored (help)CS1 maint: multiple names: authors list (link),http://www.atlascom.gr/HELLENIC_DNA_PAPER.PDF,
Crete 16 out of 193
Greece 9 out of 171 - ^ 7/100,"Y-chromosome Lineages from Portugal, Madeira and A¸cores Record Elements of Sephardim and Berber Ancestry," Annals of Human Genetics (2005) 69,443–454, Goncalves et al. (2005) http://onlinelibrary.wiley.com/doi/10.1111/j.1529-8817.2005.00161.x/pdf
- ^ Structure in the Mediterranean Basin: A Y Chromosome Perspective, Capelli et al. 2005
- ^ http://volgagermanbrit.us/documents/ejhg2008120a.pdf
- ^ Geographical Structure of the Y-chromosomal Genetic Landscape of the Levant: A coastal-inland contrast,Annals of Human Genetics (2009) 73,568–581,El-Sibai et al.(2009) (reported results from several studies : Di Giacomo et al. 2003, Al-Zahery et al. 2003, Flores et al.2004, Cinnioglu et al. 2004, Capelli et al. 2005, Goncalves et al. 2005, Zalloua et al. 2008, Cadenas et al. 2008),http://www.lbem.icb.ufmg.br/pdf/elsibai09ahg-levantegenografico.pdf
General listing
edit- Arredi, B; Poloni, E; Paracchini, S; Zerjal, T; Fathallah, D; Makrelouf, M; Pascali, V; Novelletto, A; Tylersmith, C (2004), "A Predominantly Neolithic Origin for Y-Chromosomal DNA Variation in North Africa", American Journal of Human Genetics, 75 (2): 338–345, doi:10.1086/423147, PMC 1216069, PMID 15202071
{{citation}}
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specified (help) - Balanovsky, Oleg; Dibirova, Khadizhat; Dybo, Anna; Mudrak, Oleg; Frolova, Svetlana; Pocheshkhova, Elvira; Haber, Marc; Platt, Daniel; Schurr, Theodore; Haak, Wolfgang; Kuznetsova, Marina; Radzhabov, Magomed; Balaganskaya, Olga; Zakharova, Tatiana; Soria Hernanz, David F.; Zalloua, Pierre; Koshel, Sergey; Ruhlen, Merritt; Renfrew, Colin; Wells, R. Spencer; Tyler-Smith, Chris; Balanovska, Elena (2011), Mol Biol Evol, doi:10.1093/molbev/msr126 Evolution of Genes and Languages in the Caucasus Region http://mbe.oxfordjournals.org/content/early/2011/05/13/molbev.msr126.abstract Evolution of Genes and Languages in the Caucasus Region
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- Behar; Thomas, Mark G.; Skorecki, Karl; Hammer, Michael F.; Bulygina, Ekaterina; Rosengarten, Dror; Jones, Abigail L.; Held, Karen; Moses, Vivian; et al. (2003), "Multiple Origins of Ashkenazi Levites: Y Chromosome Evidence for Both Near Eastern and European Ancestries", Am. J. Hum. Genet., vol. 73, no. 4, pp. 768–779, doi:10.1086/378506, PMC 1180600, PMID 13680527
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- Cann, HM; de Toma, C; Cazes, L; Legrand (2002), "A human genome diversity cell panel", Science, 296: 261–262
- Capelli; Onofri, Valerio; Brisighelli, Francesca; Boschi, Ilaria; Scarnicci, Francesca; Masullo, Mara; Ferri, Gianmarco; Tofanelli, Sergio; Tagliabracci, Adriano (2009), "Moors and Saracens in Europe: estimating the medieval North African male legacy in southern Europe", European Journal of Human Genetics, 17 (6): 848–852, doi:10.1038/ejhg.2008.258, PMC 2947089, PMID 19156170
- Cinnioğlu, Cengiz; King, Roy; Kivisild, Toomas; Kalfoglu, Ersi; Atasoy, Sevil; Cavalleri, Gianpiero L.; Lillie, Anita S.; Roseman, Charles C.; Lin, Alice A. (2004), "Excavating Y-chromosome haplotype strata in Anatolia" (PDF), Hum Genet, 114 (2): 127, doi:10.1007/s00439-003-1031-4, PMID 14586639
- Chiaroni, Jacques; King, Roy J.; Myres, Natalie M.; Henn, Brenna M.; Mitchell, Michael J.; Boetsch, Gilles; Sheikha, Issa; Lin, Alice A.; Nik-Ahd, Mahnoosh; Ahmad, Jabeen; Lattanzi, Francesca; Herrera, Rene J.; Ibrahim, Muntaser E.; Brody, Aaron; Semino, Ornella; Kivisild, Toomas; Underhill, Peter A. (2010). "The emergence of Y-chromosome haplogroup J1e among Arabic-speaking populations". European Journal of Human Genetics. 18: 348–353. doi:10.1038.
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value (help) - Di Gaetano; Cerutti; Crobu; Robino (2009), "Differential Greek and northern African migrations to Sicily are supported by genetic evidence from the Y chromosome", European Journal of Human Genetics, 17 (1): 91–99, doi:10.1038/ejhg.2008.120, PMC 2985948, PMID 18685561
- El-Sibai, Mirvat; Platt, Daniel E.; Haber, Marc; Xue, Yali; Youhanna, Sonia C.; Wells, R. Spencer; Izaabel, Hassan; Sanyoura, May F.; Harmanani, Haidar; Ashrafian Bonab, Maziar; Behbehani, Jaafar; Hashwa, Fuad; Tyler-Smith, Chris; Zalloua, Pierre A. (2009), Geographical Structure of the Y-chromosomal Genetic Landscape of the Levant: A coastal-inland contrast journal=Annals of Human Genetics, doi:10.1111/j.1469-1809.2009.00538.x
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- ISOGG (2011), Y-DNA Haplogroup J and its Subclades - 2011, International Society of Genetic Genealogists "ISOGG"
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: CS1 maint: unflagged free DOI (link) - Tofanelli, Sergio; Ferri, Gianmarco; Bulayeva, Kazima; Caciagli, Laura; Onofri, Valerio; Taglioli, Luca; Bulayev, Oleg; Boschi, Ilaria; Alù, Milena; Berti, Andrea; Rapone, Cesare; Beduschi, Giovanni; Luiselli, Donata; M Cadenas, Alicia; Dafaallah Awadelkarim, Khalid; Mariani-Costantini, Renato; Eldin Elwali, Nasr; Verginelli, Fabio; Pilli, Elena; J Herrera, Rene; Gusmão, Leonor; Paoli, Giorgio; Capelli, Cristian (2009), "J1-M267 Y lineage marks climate-driven pre-historical human displacements", European Journal of Human Genetics, 17: 1520–1524, doi:10.1038/ejhg.2009.58
- Y Chromosome Consortium "YCC" (2002), "A Nomenclature System for the Tree of Human Y-Chromosomal Binary Haplogroups", Genome Research, vol. 12, no. 2, pp. 339–348, doi:10.1101/gr.217602, PMC 155271, PMID 11827954
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- Zalloua; Platt, Daniel E.; El Sibai, Mirvat; Khalife, Jade; Makhoul, Nadine; Haber, Marc; Xue, Yali; Izaabel, Hassan; Bosch, Elena (2008), "Identifying Genetic Traces of Historical Expansions: Phoenician Footprints in the Mediterranean" (PDF), The American Journal of Human Genetics, 83 (5): 633–642, doi:10.1016/j.ajhg.2008.10.012, PMC 2668035, PMID 18976729