Haplogroup E-Z827
Haplogroup E-Z827 | |
---|---|
Possible time of origin | approx 24,100 years BP [1] |
Possible place of origin | Northern Africa |
Ancestor | E-M215/M35 |
Descendants | E-L19, E-Z830 |
Defining mutations | Z827 |
In human genetics, E-Z827, also known as E1b1b1b,[2] is the name of a major Y chromosome haplogroup. It is defined as the lineage which combines the haplogroups E-Z830 and E-V257, and defines their common ancestry. The former is predominantly found in the Horn of Africa and the Middle East, while the latter is most frequently observed in North Western Africa; it is also found at lower frequencies in Europe, and in isolated parts of Southeast Africa.
Subclades of E-Z827 and Distribution
Family Tree
The following phylogeny is based on the YCC 2008 tree and subsequent published research as summarized by ISOGG.[3][4][5]
- E-Z827 (Z827) - E1b1b1b[6]
- E-V257/L19 (L19, V257) - E1b1b1b1[6]
- E-PF2431
- E-M81 (M81)[7]
- E-M81*
- E-PF2546
- E-PF2546*
- E-CTS12227
- E-MZ11
- E-MZ12
- E-MZ11
- E-A929
- E-Z5009
- E-Z5009*
- E-Z5010
- E-Z5013
- E-Z5013*
- E-A1152
- E-A2227
- E-A428
- E-MZ16
- E-PF6794
- E-PF6794*
- E-PF6789
- E-MZ21
- E-MZ23
- E-MZ80
- E-A930
- E-Z2198/E-MZ46
- E-A601
- E-L351
- E-Z5009
- E-Z830 (Z830) - E1b1b1b2[6]
- E-M123 (M123)
- E-M34 (M34)
- E-M84 (M84)
- E-M136 (M136)
- E-M290 (M290)
- E-V23 (V23)
- E-L791 (L791,L792)
- E-M84 (M84)
- E-M34 (M34)
- E-V1515
- E-V1515*
- E-V1486
- E-V1486*
- E-V2881
- E-V2881*
- E-V1792
- E-V92
- E-M293 (M293)
- E-M293*
- E-P72 (P72)
- E-V3065*
- E-V1700
- E-V42 (V42)
- E-V1785
- E-V1785*
- E-V6 (V6)
- E-M123 (M123)
- E-V257/L19 (L19, V257) - E1b1b1b1[6]
E-V257/L19 (E1b1b1b1)
E-V257/L19 showed a parallel with its sibling clade E-V68 in the way that both clades show signs of having migrated from North Africa to southern Europe across the Mediterranean sea. 6 "E-V257/L19*" individuals were found in published samples who were E-V257/L19, but not E-M81. a Marrakesh Berber, a Corsican, a Sardinian, A Borana from Kenya, a southern Spaniard and a Cantabrian.
Within E-M35, there are striking parallels between two haplogroups, E-V68 and E-V257. Both contain a lineage which has been frequently observed in North Africa (E-M78 and E-M81, respectively) and a group of undifferentiated chromosomes that are mostly found in southern Europe. An expansion of E-M35 carriers, possibly from the Middle East as proposed by other authors, and split into two branches separated by the geographic barrier of the Mediterranean Sea, would explain this geographic pattern. However, the absence of E-V68* and E-V257* in the Middle East makes a maritime spread between northern Africa and southern Europe a more plausible hypothesis.
A project dedicated to researching and understanding the origins of E-V257/L19* is underway at FamilyTreeDNA.com. The name of the project is E1b1b1b*-A
E-V257's dominant sub-clade E-M81 is thought to have originated in the area of North Africa 14,200 years ago.[8]
E-M81
E-M81 is the most common subclade of E-L19/V257 and found in the Maghreb, dominated by its sub-clade E-M183. This haplogroup reaches a mean frequency of 42% in North Africa, decreasing in frequency from 100% in some isolated Berber populations to approximately 10% to the east of this range in Egypt.[9][10][11] Because of its prevalence among these groups and also others such as Mozabite, Middle Atlas, Kabyle and other Berber groups, it is sometimes referred to as a genetic "Berber marker". Pereira et al. (2010) report high levels among Tuareg in two Saharan populations - 77.8% near Gorom-Gorom, in Burkina Faso, and 81.8% from Gossi in Mali. There was a much lower frequency of 11.1% in the vicinity of Tanut in the Republic of Niger.
E-M81 is also quite common among North African Arabic-speaking groups. It is generally found at frequencies around 45% in coastal cities of the Maghreb (Oran, Tunis, Tizi Ouzou, Algiers).[12]
In this key area from Egypt to the Atlantic Ocean, Arredi et al. (2004) report a pattern of decreasing STR haplotype variation (implying decreasing lineage age in those areas) from East to West, accompanied by a substantial increasing frequency. At the eastern extreme of this core range, Kujanova et al. (2009) found M81 in 28.6% (10 out of 35 men) in El-Hayez in the Western desert in Egypt
Arredi et al. (2004) believe the pattern of distribution and variance to be consistent with the hypothesis of a post Paleolithic "demic diffusion" from the East. The ancestral lineage of E-M81 in their hypothesis could have been linked with the spread of Neolithic food-producing technologies from the Fertile Crescent via the Nile, although pastoralism rather than agriculture. E-M81 and possibly proto-Afroasiatic language may have been carried either all the way from Asia, or they may represent a "local contribution to the North African Neolithic transition". According to Shomarka Keita, a Near Eastern origin of proto-Afroasiatic speakers carrying E-M81, or its ancestral lineage, is inconsistent with the linguistic evidence, which seems to indicate an African origin of Proto-Afro-Asiatic speakers. Keita argues that there is no autochthonous presence of E-M81 in the Near East, indicating that M81 most likely emerged from its parent clade M35 either in the Maghreb, or possibly as far southeast as the Horn of Africa.[13]
Europe
In Europe, E-M81 has a widespread distribution at very low frequencies but is common mostly in the Iberian Peninsula, where unlike in the rest of Europe[Note 1] it is more common than E-M78, with an average frequency around 5%. Its frequencies are higher in the western half of the peninsula with frequencies reaching 8% in Extremadura and South Portugal, 4% in one study and 9% in another in Galicia, 10% in Western Andalusia and Northwest Castile and 9% to 17% in Cantabria.[14][15][16][17][18] The highest frequencies of this clade found so far in Europe were observed in the Pasiegos from Cantabria, ranging from 18% (8/45)[18] to 41% (23/56).[19] An average frequency of 8.28% (54/652) has also been reported in the Spanish Canary Islands with frequencies over 10% in the three largest islands of Tenerife (10.68%), Gran Canaria (11.54%) and Fuerteventura (13.33%).[20]
E-M81 is also found in France, 2.70% (15/555) overall with frequencies surpassing 5% in Auvergne (5/89) and Île-de-France (5/91),[21][22] in Sicily (approximately 2% overall, but up to 7% in Piazza Armerina),[23] and in slightly lower frequencies in continental Italy (especially near Lucera)[17] due to historic colonization during the Islamic, Roman, and Carthaginian empires or ancient migrations in the Metals Ages through maritime means. E-M81 was also found in 2013 at 5.8% in a large sample of 1 204 Sardinians.[24]
Latin America
As a result of Spanish and Portuguese colonization of the Americas, this sub-clade is found throughout Latin America, for example 6.1% in Cuba,[25] 5.4% in Brazil (Rio de Janeiro), [Note 2] and among Hispanic men from California and Hawaii 2.4%.[26]
Others
In smaller numbers, E-M81 men can be found in areas in contact with the Maghreb, both around the Sahara, in places like Sudan, and around the Mediterranean in places like Lebanon, Turkey, and amongst Sephardic Jews.
Distribution
The following gives a summary of most of the studies which specifically tested for E-M81, showing where its distribution is greater than 1% in Europe, North Africa, the Middle East and Latin America.
Country/Region | Sampling | n | %E-M81 | Source |
---|---|---|---|---|
Algeria | Mozabite Berbers | 67 | 86.6 | Dugoujon et al. (2009) |
Algeria | Mozabite Berbers | 20 | 80 | Cruciani et al. (2004) |
Algeria | Oran | 102 | 45.1 | Robino et al. (2008) |
Algeria | Algiers | 35 | 42.9 | Arredi et al. (2004) |
Algeria | Kabyles from Tizi Ouzou | 19 | 47.4 | Arredi et al. (2004) |
Brazil | Rio de Janeiro | 112 | 5.4 | Silva et al. (2006) |
Burkina Faso | Tuaregs | 38 | 77.8 | Pereira et al. (2010) |
Canary Islands | Fuerteventura | 75 | 13.3 | Fregel et al. (2009) |
Canary Islands | Gran Canaria | 78 | 11.5 | Fregel et al. (2009) |
Canary Islands | Tenerife | 178 | 10.7 | Fregel et al. (2009) |
Canary Islands | Lanzarote | 97 | 6.2 | Fregel et al. (2009) |
Canary Islands | La Palma | 85 | 5.9 | Fregel et al. (2009) |
Canary Islands | Gomera | 92 | 4.4 | Fregel et al. (2009) |
Canary Islands | Hierro | 47 | 2.1 | Fregel et al. (2009) |
Cuba | 132 | 6.1 | Mendizabal et al. (2008) | |
Cyprus | Turkish Cypriots | 46 | 8.7 | Cruciani et al. (2004) |
Egypt | Northern Egyptians | 21 | 4.8 | Cruciani et al. (2004) |
Egypt | Western Desert | 35 | 28.6 | Kujanová et al. (2009) |
Egypt | 147 | 8.2 | Flores et al. (2005) | |
France | 85 | 3.5 | Cruciani et al. (2004) | |
France | Auvergne | 89 | 5.6 | Ramos-Luisa et al. (2009) |
France | Île-de-France | 91 | 5.5 | Ramos-Luisa et al. (2009) |
France | Nord-Pas-de-Calais | 68 | 4.4 | Ramos-Luisa et al. (2009) |
France | Provence-Alpes-Côte d'Azur | 45 | 2.2 | Ramos-Luisa et al. (2009) |
France | Midi-Pyrénées | 67 | 1.5 | Ramos-Luisa et al. (2009) |
Iberia | Spain, Portugal | 655 | 5.2 | Fregel et al. (2009) |
Iberia | Spain, Portugal | 1140 | 4.3 | Adams et al. (2008) |
Israel | Bedouins | 28 | 3.6 | Cruciani et al. (2004) |
Italy | Central Italians | 89 | 2.2 | Cruciani et al. (2004) |
Italy | Northern Italians | 67 | 1.5 | Cruciani et al. (2004) |
Italy | North-West Apulia | 46 | 4.3 | Capelli et al. (2009) |
Italy | East Campania | 84 | 2.4 | Capelli et al. (2009) |
Italy | Veneto | 55 | 1.8 | Capelli et al. (2009) |
Italy | North-East Latium | 55 | 1.8 | Capelli et al. (2009) |
Italy | Lucera | 60 | 1.7 | Capelli et al. (2009) |
Italy | Sicily | 236 | 2.1 | Gaetano et al. (2008) |
Italy | Sardinia | 1204 | 5.8 | Francalacci et al. (2013)[24] |
Jordania | 101 | 4 | Flores et al. (2005) | |
Lebanon | 104 | 1.9 | Flores et al. (2005) | |
Lebanon | 914 | 1.2 | Zalloua et al. (2008) | |
Libya | Tuaregs | 47 | 48.9 | Ottoni et al. (2011) |
Libya | Arabs | 215 | 35.9 | Fadhlaoui-Zid et al. (2013) |
Mali | Tuaregs (Gozi) | 21 | 81.8 | Pereira et al. (2010) |
Morocco | Marrakesh Berbers | 29 | 72.4 | Cruciani et al. (2004) |
Morocco | Southern Moroccan Berbers | 187 | 98.5 | Ahmed Reguig et al. (2014)[27] |
Morocco | Moyen Atlas Berbers | 69 | 71 | Cruciani et al. (2004) |
Morocco | Moroccan Arabs | 54 | 31.5 | Cruciani et al. (2004) |
Morocco | Marrakesh (Amizmiz Valley) | 33 | 84.8 | Alvarez et al. (2009) |
Morocco | Northern Moroccans (Beni Snassen) | 67 | 79.1 | Dugoujon et al. (2005) |
Morocco | Northern Moroccans (Rhiraya) | 54 | 79.6 | Dugoujon et al. (2005)[28] |
Morocco | Immigrants resident in Italy | 51 | 54.9 | Onofri et al. (2008) |
Morocco | Arabs and Berbers | 221 | 65 | Fregel et al. (2009), from Bosh et al. 2001 |
Niger | Tuaregs | 22 | 9.1 | Cruciani et al. (2004) |
Niger | Tuaregs | 31 | 11.1 | Pereira et al. (2010) |
North Africa | Sahara | 89 | 59.6 | Fregel et al. (2009) |
North Africa | Algeria, Tunisia | 202 | 39.1 | Fregel et al. (2009) |
Portugal | North | 109 | 5.5 | Flores et al. (2004) |
Portugal | South | 49 | 12.2 | Cruciani et al. (2004) |
Portugal | North | 50 | 4 | Cruciani et al. (2004) |
Portugal | South | 78 | 7.7 | Adams et al. (2008) |
Portugal | North | 60 | 3.3 | Adams et al. (2008) |
Portugal | 303 | 5.6 | Goncalves et al. (2005) | |
Portugal | North | 101 | 6 | Goncalves et al. (2005) |
Portugal | Center | 102 | 4.9 | Goncalves et al. (2005) |
Portugal | South | 100 | 6 | Goncalves et al. (2005) |
Portugal | Madeira | 129 | 5.4 | Goncalves et al. (2005) |
Portugal | Açores | 121 | 5 | Goncalves et al. (2005) |
Portugal | 657 | 5.6 | Beleza et al. (2006) | |
Portugal | Entre Douro e Minho | 228 | 6.6 | Beleza et al. (2006) |
Portugal | Tras os Montes | 64 | 3.1 | Beleza et al. (2006) |
Portugal | Beira Litoral | 116 | 5.2 | Beleza et al. (2006) |
Portugal | Beira Interior | 58 | 5.3 | Beleza et al. (2006) |
Portugal | Estremadura | 43 | 4.6 | Beleza et al. (2006) |
Portugal | Lisboa e Setubal | 62 | 6.5 | Beleza et al. (2006) |
Portugal | Alentejo | 65 | 7.7 | Beleza et al. (2006) |
Portugal | Coruche | 64 | 9.4 | Pereira et al. (2010) |
Portugal | Pias | 46 | 4.3 | Pereira et al. (2010) |
Portugal | Alcacer do Sal | 21 | 4.8 | Pereira et al. (2010) |
Portugal | Tras-os-Montes (Jews) | 57 | 5.3 | Nogueiro et al. (2010) |
Portugal | Tras-os-Montes (Non Jews) | 30 | 10 | Nogueiro et al. (2010) |
Somalia | 201 | 1.5 | Flores et al. (2005) | |
Spain | Pasiegos from Cantabria | 19 | 36.8 | Scozzari et al. (2001) |
Spain | Pasiegos from Cantabria | 56 | 41.1 | Cruciani et al. (2004) |
Spain | Pasiegos from Cantabria | 45 | 17.8 | Maca-Meyer et al. (2003) |
Spain | Spanish Basques | 55 | 3.6 | Cruciani et al. (2004) |
Spain | Asturians | 90 | 2.2 | Cruciani et al. (2004) |
Spain | Southern Spaniards | 62 | 1.6 | Cruciani et al. (2004) |
Spain | Castile, NorthWest | 100 | 10 | Adams et al. (2008) |
Spain | Andalucia, West | 73 | 9.6 | Adams et al. (2008) |
Spain | Galicia | 19 | 10.5 | Flores et al. (2004) |
Spain | Galicia | 292 | 4.1 | Brion et al. (2004) |
Spain | Galicia | 88 | 9.1 | Adams et al. (2008) |
Spain | Galicia | 44 | 9.1 | Santos et al. (2013) |
Spain | Galicia | 164 | 9.1 | Regueiro et al. (2015) |
Spain | Extremadura | 52 | 7.7 | Adams et al. (2008) |
Spain | Valencia | 73 | 4.1 | Adams et al. (2008) |
Spain | Castile, NorthEast | 31 | 3.2 | Adams et al. (2008) |
Spain | Aragon | 34 | 2.9 | Adams et al. (2008) |
Spain | Minorca | 37 | 2.7 | Adams et al. (2008) |
Spain | Andalucia, East | 95 | 2.1 | Adams et al. (2008) |
Spain | Majorca | 62 | 1.6 | Adams et al. (2008) |
Spain | Castile, La Mancha | 63 | 1.6 | Adams et al. (2008) |
Spain | Catalonia | 80 | 1.3 | Adams et al. (2008) |
Spain | Catalonia | 111 | 3.6 | Santos et al. (2013) |
Spain | Cantabria | 161 | 13 | Capelli et al. (2009) |
Spain | Malaga | 26 | 11.5 | Flores et al. (2004) |
Spain | Cantabria | 70 | 8.6 | Flores et al. (2004) |
Spain | Cordoba | 27 | 7.4 | Flores et al. (2004) |
Spain | Valencia | 31 | 6.5 | Flores et al. (2004) |
Spain | Valencia | 59 | 5.1 | Santos et al. (2013) |
Spain | Almeria | 36 | 5.6 | Santos et al. (2013) |
Spain | Leon | 60 | 5 | Flores et al. (2004) |
Spain | Castile | 21 | 4.8 | Flores et al. (2004) |
Spain | Seville | 155 | 4.5 | Flores et al. (2004) |
Spain | Huelva | 22 | 4.5 | Flores et al. (2004) |
Spain | Basques | 45 | 2.2 | Flores et al. (2004) |
Spain | Huelva | 167 | 3 | Ambrosio et al. (2010) |
Spain | Granada | 250 | 3.6 | Ambrosio et al. (2010) |
Spain | Pedroches Valley | 68 | 1.5 | Alvarez et al. (2009) |
Spain | Andalusians | 94 | 2.1 | Alvarez et al. (2009) |
Spain | Zamora | 235 | 5.5 | Alvarez et al. (2014) |
Tunisia | Tunis | 148 | 37.9 | Arredi et al. (2004) |
Tunisia | Immigrants resident in Italy | 52 | 32.7 | Onofri et al. (2008) |
Tunisia | Berbers from Bou Omrane | 40 | 87.5 | Ennafaa et al. (2011) |
Tunisia | Berbers from Bou Saad | 40 | 92.5 | Ennafaa et al. (2011) |
Tunisia | Jerbian Arabs | 46 | 60.9 | Ennafaa et al. (2011) |
Tunisia | Jerbian Berbers | 47 | 76.6 | Ennafaa et al. (2011) |
Tunisia | Berbers from Chenini–Douiret | 27 | 100 | Fadhlaoui-Zid et al. (2011) |
Tunisia | Berbers from Sened | 35 | 65.7 | Fadhlaoui-Zid et al. (2011) |
Tunisia | Berbers from Jradou | 32 | 100 | Fadhlaoui-Zid et al. (2011) |
Tunisia | Andalusian Zaghouan | 32 | 40.6 | Fadhlaoui-Zid et al. (2011) |
Tunisia | Cosmopolitan Tunis | 33 | 54.4 | Fadhlaoui-Zid et al. (2011) |
Turkey | Istanbul Turkish | 35 | 5.7 | Cruciani et al. (2004) |
Turkey | Sephardi Turkish | 19 | 5.3 | Cruciani et al. (2004) |
Turkey | Southwestern Turkish | 40 | 2.5 | Cruciani et al. (2004) |
Turkey | Northeastern Turkish | 41 | 2.4 | Cruciani et al. (2004) |
E-Z830 (E1b1b1b2)
A recently confirmed sub-clade of E-Z827, Z830, includes the confirmed sub-clades of E-M123, E-M293, and E-V42, and is a sibling clade to E-L19. Currently, the E-M35 phylogeny project recognizes four distinct clusters of Z830* carriers, two of which are exclusively Jewish in origin. The remaining two are significantly smaller, and include scattered individuals in Germany, Spain, Latin America, Egypt, and Ethiopia.[29][30][31][32]
E-M123
E-M123 is mostly known for its major subclade E-M34, which dominates this clade.[Note 3]
E-V1515
A new clade (E-V1515) was defined by Trombetta et al. 2015, which originated about 12 kya (95% CI 8.6-16.4) in eastern Africa where it is currently mainly distributed. This clade includes all the sub-Saharan haplogroups (E-V42, E-M293, E-V92, E-V6) reported as E-M35 basal clades in a previous phylogeny.[33]
E-M293
E-M293 is a subclade of E-V1515. It was identified by ISOGG as the second clade within E-Z830. It was discovered before E-Z830, being announced in Henn 2008 , which associated it with the spread of pastoralism from Eastern Africa into Southern Africa. So far high levels have been found in specific ethnic groups in Tanzania and Southern Africa. Highest were the Datog (43%), Khwe (Kxoe) (31%), Burunge (28%), and Sandawe (24%). Henn (2008) in their study also found two Bantu-speaking Kenyan males with the M293 mutation.[34] Other E-M215 subclades are rare in Southern Africa. The authors state...
Without information about M293 in the Maasai, Hema, and other populations in Kenya, Sudan, and Ethiopia, we cannot pinpoint the precise geographic source of M293 with greater confidence. However, the available evidence points to present-day Tanzania as an early and important geographic locus of M293 evolution.
They also say that "M293 is only found in sub-Saharan Africa, indicating a separate phylogenetic history for M35.1 * (former) samples further north". E-P72 appears in Karafet (2008). Trombetta et al. 2011 announced that this is a subclade of E-M293.
E-V42
Trombetta et al. 2011 announced the discovery of E-V42 in two Ethiopian Jews. It was suggested that it may be restricted to the region around Ethiopia. However, further testing by commercial DNA testing companies confirmed positive results for this subclade in Arabia as well.[35]
E-V6
The E-V6 subclade of E-V1515 is defined by V6. Cruciani et al. (2004) identified a significant presence of these lineages in Ethiopia, and also some in the neighboring Somali population. Among the Ethiopian and Somali samples, the highest were 14.7% among the Ethiopian Amhara, and 16.7% among the Ethiopian Wolayta.
E-V92
Trombetta et al. 2011 announced the discovery of E-V92 in two Ethiopian Amhara. Like E-V6 and E-V42 it possibly only exists in the area of Ethiopia.
Phylogenetics
Phylogenetic History
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 |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
E-P29 | 21 | III | 3A | 13 | Eu3 | H2 | B | E* | E | E | E | E | E | E | E | E | E | E |
E-M33 | 21 | III | 3A | 13 | Eu3 | H2 | B | E1* | E1 | E1a | E1a | E1 | E1 | E1a | E1a | E1a | E1a | E1a |
E-M44 | 21 | III | 3A | 13 | Eu3 | H2 | B | E1a | E1a | E1a1 | E1a1 | E1a | E1a | E1a1 | E1a1 | E1a1 | E1a1 | E1a1 |
E-M75 | 21 | III | 3A | 13 | Eu3 | H2 | B | E2a | E2 | E2 | E2 | E2 | E2 | E2 | E2 | E2 | E2 | E2 |
E-M54 | 21 | III | 3A | 13 | Eu3 | H2 | B | E2b | E2b | E2b | E2b1 | - | - | - | - | - | - | - |
E-P2 | 25 | III | 4 | 14 | Eu3 | H2 | B | E3* | E3 | E1b | E1b1 | E3 | E3 | E1b1 | E1b1 | E1b1 | E1b1 | E1b1 |
E-M2 | 8 | III | 5 | 15 | Eu2 | H2 | B | E3a* | E3a | E1b1 | E1b1a | E3a | E3a | E1b1a | E1b1a | E1b1a | E1b1a1 | E1b1a1 |
E-M58 | 8 | III | 5 | 15 | Eu2 | H2 | B | E3a1 | E3a1 | E1b1a1 | E1b1a1 | E3a1 | E3a1 | E1b1a1 | E1b1a1 | E1b1a1 | E1b1a1a1a | E1b1a1a1a |
E-M116.2 | 8 | III | 5 | 15 | Eu2 | H2 | B | E3a2 | E3a2 | E1b1a2 | E1b1a2 | E3a2 | E3a2 | E1b1a2 | E1b1a2 | E1ba12 | removed | removed |
E-M149 | 8 | III | 5 | 15 | Eu2 | H2 | B | E3a3 | E3a3 | E1b1a3 | E1b1a3 | E3a3 | E3a3 | E1b1a3 | E1b1a3 | E1b1a3 | E1b1a1a1c | E1b1a1a1c |
E-M154 | 8 | III | 5 | 15 | Eu2 | H2 | B | E3a4 | E3a4 | E1b1a4 | E1b1a4 | E3a4 | E3a4 | E1b1a4 | E1b1a4 | E1b1a4 | E1b1a1a1g1c | E1b1a1a1g1c |
E-M155 | 8 | III | 5 | 15 | Eu2 | H2 | B | E3a5 | E3a5 | E1b1a5 | E1b1a5 | E3a5 | E3a5 | E1b1a5 | E1b1a5 | E1b1a5 | E1b1a1a1d | E1b1a1a1d |
E-M10 | 8 | III | 5 | 15 | Eu2 | H2 | B | E3a6 | E3a6 | E1b1a6 | E1b1a6 | E3a6 | E3a6 | E1b1a6 | E1b1a6 | E1b1a6 | E1b1a1a1e | E1b1a1a1e |
E-M35 | 25 | III | 4 | 14 | Eu4 | H2 | B | E3b* | E3b | E1b1b1 | E1b1b1 | E3b1 | E3b1 | E1b1b1 | E1b1b1 | E1b1b1 | removed | removed |
E-M78 | 25 | III | 4 | 14 | Eu4 | H2 | B | E3b1* | E3b1 | E1b1b1a | E1b1b1a1 | E3b1a | E3b1a | E1b1b1a | E1b1b1a | E1b1b1a | E1b1b1a1 | E1b1b1a1 |
E-M148 | 25 | III | 4 | 14 | Eu4 | H2 | B | E3b1a | E3b1a | E1b1b1a3a | E1b1b1a1c1 | E3b1a3a | E3b1a3a | E1b1b1a3a | E1b1b1a3a | E1b1b1a3a | E1b1b1a1c1 | E1b1b1a1c1 |
E-M81 | 25 | III | 4 | 14 | Eu4 | H2 | B | E3b2* | E3b2 | E1b1b1b | E1b1b1b1 | E3b1b | E3b1b | E1b1b1b | E1b1b1b | E1b1b1b | E1b1b1b1 | E1b1b1b1a |
E-M107 | 25 | III | 4 | 14 | Eu4 | H2 | B | E3b2a | E3b2a | E1b1b1b1 | E1b1b1b1a | E3b1b1 | E3b1b1 | E1b1b1b1 | E1b1b1b1 | E1b1b1b1 | E1b1b1b1a | E1b1b1b1a1 |
E-M165 | 25 | III | 4 | 14 | Eu4 | H2 | B | E3b2b | E3b2b | E1b1b1b2 | E1b1b1b1b1 | E3b1b2 | E3b1b2 | E1b1b1b2a | E1b1b1b2a | E1b1b1b2a | E1b1b1b2a | E1b1b1b1a2a |
E-M123 | 25 | III | 4 | 14 | Eu4 | H2 | B | E3b3* | E3b3 | E1b1b1c | E1b1b1c | E3b1c | E3b1c | E1b1b1c | E1b1b1c | E1b1b1c | E1b1b1c | E1b1b1b2a |
E-M34 | 25 | III | 4 | 14 | Eu4 | H2 | B | E3b3a* | E3b3a | E1b1b1c1 | E1b1b1c1 | E3b1c1 | E3b1c1 | E1b1b1c1 | E1b1b1c1 | E1b1b1c1 | E1b1b1c1 | E1b1b1b2a1 |
E-M136 | 25 | III | 4 | 14 | Eu4 | H2 | B | E3ba1 | E3b3a1 | E1b1b1c1a | E1b1b1c1a1 | E3b1c1a | E3b1c1a | E1b1b1c1a1 | E1b1b1c1a1 | E1b1b1c1a1 | E1b1b1c1a1 | E1b1b1b2a1a1 |
Original Research Publications
The following research teams per their publications were represented in the creation of the YCC Tree.
See also
Genetics
Y-DNA E Subclades
Y-DNA Backbone Tree
Notes
- ^ Adams et al. (2008) , shows an average frequency of 4.3% (49/1140) in the Iberian Peninsula with frequencies reaching 4% and 9% in two separate surveys of Galicia, 10% in Western Andalusia and Northwest Castile. However this study also includes 153 individuals from Majorca, Minorca and Ibiza islands as well as 24 individuals from Gascony which are not in the Iberian Peninsula. Without these 177 individuals, real average for Iberian Peninsula is 4.9% (47/963) , see table.
- ^ (6 out of 112), "The presence of chromosomes of North African origin (E3b1b-M81; Cruciani et al., 2004) can also be explained by a Portuguese-mediated influx, since this haplogroup reaches a frequency of 5.6% in Portugal (Beleza et al., (2006) ), quite similar to the frequency found in Rio de Janeiro (5.4%) among European contributors." Silva et al. (2006)
- ^ As of 11 November 2008 for example, the E-M35 phylogeny project had records of four E-M123* tests, compared to 93 test results with E-M34.
- ^ https://www.yfull.com/tree/E-Z827/
- ^ ISOGG (2015), Y-DNA Haplogroup E and its Subclades - 2015
- ^ ISOGG (2008)
- ^ Karafet et al. (2008)
- ^ Y Chromosome Consortium "YCC" (2002)
- ^ a b c ISOGG 2015
- ^ "E-M81 YTree". www.yfull.com. Retrieved 2016-07-09.
- ^ "E-M81 YTree". www.yfull.com. Retrieved 2016-07-10.
- ^ Arredi et al. (2004)
- ^ Alvarez et al. (2009)
- ^ Bosch et al. (2006)
- ^ Robino (2008), Arredi (2004)
- ^ Keita (2008), "Geography, selected Afro-Asiatic families, and Y chromosome lineage variation", In Hot Pursuit of Language
{{citation}}
: External link in
(help); Unknown parameter|chapterurl=
|chapterurl=
ignored (|chapter-url=
suggested) (help) - ^ Adams et al. (2008)
- ^ Flores et al. (2005)
- ^ Beleza et al. (2006)
- ^ a b Capelli et al. (2009)
- ^ a b Maca-Meyer, N.; Sánchez-Velasco, P.; Flores, C.; Larruga, J. M.; González, A. M.; Oterino, A.; Leyva-Cobián, F. (2003), "Y Chromosome and Mitochondrial DNA Characterization of Pasiegos, a Human Isolate from Cantabria (Spain)", Annals of Human Genetics, 67 (Pt 4): 329–339, doi:10.1046/j.1469-1809.2003.00045.x, PMID 12914567.
- ^ Cruciani et al. (2004)
- ^ Fregel et al. (2009), Demographic history of Canary Islands male gene-pool: replacement of native lineages by European, see table
- ^ Ramos-Luisa et al. (2009)
- ^ exluding recent immigration as only men with French surname were analysed
- ^ Di Gaetano et al. (2009)
- ^ a b Francalacci et al. (2013), Low-Pass DNA Sequencing of 1200 Sardinians Reconstructs European Y-Chromosome Phylogeny
- ^ (8 out of 132), Mendizabal et al. (2008)
- ^ (7 out of 295), Paracchini et al. (2003)
- ^ Ahmed, Reguig,; Nourdin, Harich,; Abdelhamid, Barakat,; Hassan, Rouba, (2014-01-01). "Phylogeography of E1b1b1b-M81 Haplogroup and Analysis of its Subclades in Morocco". 86 (2).
{{cite journal}}
: Cite journal requires|journal=
(help)CS1 maint: extra punctuation (link) CS1 maint: multiple names: authors list (link) - ^ "The Berbers Linguistic and enetic diversity" (PDF). ddl.ish-lyon.cnrs.fr.
- ^ "E-M35 Project Data". haplozone.net.
- ^ "E-M35 Project Data". haplozone.net.
- ^ "E-M35 Project Data". haplozone.net.
- ^ "E-M35 Project Data". haplozone.net.
- ^ Trombetta et al. 2015, Phylogeographic refinement and large scale genotyping of human Y chromosome haplogroup E provide new insights into the dispersal of early pastoralists in the African continent
- ^ Henn et al. (2008)
- ^ "E-M35 Project Data". haplozone.net.
References
- Ambrosio, B; Dugoujon, JM; Hernández, C; De La Fuente, D; González-Martín, A; Fortes-Lima, CA; Novelletto, A; Rodríguez, JN; Calderón, R; et al. (2010), "The Andalusian population from Huelva reveals a high diversification of Y-DNA paternal lineages from haplogroup E: Identifying human male movements within the Mediterranean space", Annals of Human Biology, 37 (1): 86–107, doi:10.3109/03014460903229155, PMID 19939195
- Battaglia, Vicenza; Fornarino, Simona; Al-Zahery, Nadia; Olivieri, Anna; Pala, Maria; Myres, Natalie M; King, Roy J; Rootsi, Siiri; et al. (2008), "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 2947100, PMID 19107149
- Bird, Steven (2007), "Haplogroup E3b1a2 as a Possible Indicator of Settlement in Roman Britain by Soldiers of Balkan Origin", Journal of Genetic Genealogy, 3 (2)
- Bosch, E; Calafell, F; González-Neira, A; Flaiz, C; Mateu, E; Scheil, HG; Huckenbeck, W; Efremovska, L; et al. (2006), "Paternal and maternal lineages in the Balkans show a homogeneous landscape over linguistic barriers, except for the isolated Aromuns", Annals of Human Genetics, 70 (Pt 4): 459–487, doi:10.1111/j.1469-1809.2005.00251.x, PMID 16759179
- Cadenas; Zhivotovsky, Lev A; Cavalli-Sforza, Luca L; Underhill, Peter A; Herrera, Rene J (2007), "Y-chromosome diversity characterizes the Gulf of Oman", European Journal of Human Genetics, 16 (3): 1–13, doi:10.1038/sj.ejhg.5201934, PMID 17928816
- Capelli, C; Redhead, N; Abernethy, JK; Gratrix, F; Wilson, JF; Moen, T; Hervig, T; Richards, M; et al. (2003), "A Y Chromosome Census of the British Isles", Current Biology, 13 (11): 979–84, doi:10.1016/S0960-9822(03)00373-7, PMID 12781138 also at [1]
- Caratti; Gino, S.; Torre, C.; Robino, C. (2009), "Subtyping of Y-chromosomal haplogroup E-M78 (E1b1b1a) by SNP assay and its forensic application", International Journal of Legal Medicine, 123 (4): 357, doi:10.1007/s00414-009-0350-y, PMID 19430804
- Cruciani, F; Santolamazza, P; Shen, P; MacAulay, V; Moral, P; Olckers, A; Modiano, D; Holmes, S; et al. (2002), "A Back Migration from Asia to Sub-Saharan Africa Is Supported by High-Resolution Analysis of Human Y-Chromosome Haplotypes" (PDF), American Journal of Human Genetics, 70 (5): 1197–1214, doi:10.1086/340257, PMC 447595, PMID 11910562
- Cruciani, F; La Fratta, R; Santolamazza, P; Sellitto, D; Pascone, R; Moral, P; Watson, E; Guida, V; et al. (May 2004), "Phylogeographic Analysis of Haplogroup E3b (E-M215) Y Chromosomes Reveals Multiple Migratory Events Within and Out Of Africa" (PDF), American Journal of Human Genetics, 74 (5): 1014–1022, doi:10.1086/386294, PMC 1181964, PMID 15042509
- Cruciani, Fulvio; La Fratta, Roberta; Torroni, Antonio; Underhill, Peter A.; Scozzari, Rosaria (2006), "Molecular Dissection of the Y Chromosome Haplogroup E-M78 (E3b1a): A Posteriori Evaluation of a Microsatellite-Network-Based Approach Through Six New Biallelic Markers" (PDF), Human Mutation, 27 (8): 831, doi:10.1002/humu.9445, PMID 16835895
- Cruciani, F; La Fratta, R; Trombetta, B; Santolamazza, P; Sellitto, D; Colomb, EB; Dugoujon, JM; Crivellaro, F; et al. (2007), "Tracing Past Human Male Movements in Northern/Eastern Africa and Western Eurasia: New Clues from Y-Chromosomal Haplogroups E-M78 and J-M12", Molecular Biology and Evolution, 24 (6): 1300–1311, doi:10.1093/molbev/msm049, PMID 17351267 Also see Supplementary Data.
- Di Gaetano (2008), "Differential Greek and northern African migrations to Sicily are supported by genetic evidence from the Y chromosome", European Journal of Human Genetics, 17, doi:10.1038/ejhg.2008.120, PMC 2985948, PMID 18685561
- Di Giacomo; Luca, F.; Anagnou, N.; Ciavarella, G.; Corbo, R.M.; Cresta, M.; Cucci, F.; Di Stasi, L.; Agostiano, V.; Giparaki, M.; Loutradis, A.; Mammi’, C.; Michalodimitrakis, E.N.; Papola, F.; Pedicini, G.; Plata, E.; Terrenato, L.; Tofanelli, S.; Malaspina, P.; Novelletto, A. (2003), "Clinal patterns of human Y chromosomal diversity in continental Italy and Greece are dominated by drift and founder effects" (PDF), Molecular Phylogenetics and Evolution, 28 (3): 387–395, doi:10.1016/S1055-7903(03)00016-2, PMID 12927125
- Firasat; 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): 121–126, doi:10.1038/sj.ejhg.5201726, PMC 2588664, PMID 17047675
- Hassan, HY; Underhill, PA; Cavalli-Sforza, LL; Ibrahim, ME (2008), "Y-Chromosome Variation Among Sudanese: Restricted Gene Flow, Concordance With Language, Geography, and History" (PDF), American Journal of Physical Anthropology, 137 (3): 316, doi:10.1002/ajpa.20876, PMID 18618658
- ISOGG (2008), Y-DNA Haplogroup E and its Subclades - 2008, International Society of Genetic Genealogists "ISOGG"
- Karafet, TM; Mendez, FL; Meilerman, MB; Underhill, PA; Zegura, SL; Hammer, MF (May 2008), Abstract "New Binary Polymorphisms Reshape and Increase Resolution of the Human Y-Chromosomal Haplogroup Tree", Genome Research, 18 (5): 830, doi:10.1101/gr.7172008, PMC 2336805, PMID 18385274
{{citation}}
: Check|url=
value (help). Published online April 2, 2008. See also Supplementary Material. - King, RJ; Ozcan, SS; Carter, T; Kalfoğlu, E; Atasoy, S; Triantaphyllidis, C; Kouvatsi, A; Lin, AA; et al. (2008), "Differential Y-chromosome Anatolian Influences on the Greek and Cretan Neolithic" (PDF), Annals of Human Genetics, 72 (Pt 2): 205–214, doi:10.1111/j.1469-1809.2007.00414.x, PMID 18269686
- King and Underhill (2002), "Congruent distribution of Neolithic painted pottery and ceramic figurines with Y-chromosome lineages", Antiquity, 76: 707–14, doi:10.1017/s0003598x00091158
- Kujanova; Pereira, Luísa; Fernandes, Verónica; Pereira, Joana B.; čErný, Viktor (2009), "Near Eastern Neolithic Genetic Input in a Small Oasis of the Egyptian Western Desert", American Journal of Physical Anthropology, 140 (2): 336, doi:10.1002/ajpa.21078, PMID 19425100
- Lancaster, Andrew (2009), "Y Haplogroups, Archaeological Cultures and Language Families: a Review of the Multidisciplinary Comparisons using the case of E-M35" (PDF), Journal of Genetic Genealogy, 5 (1)
- Onofri; Alessandrini, F; Turchi, C; Pesaresi, M; Buscemi, L; Tagliabracci, A (2006), "Development of multiplex PCRs for evolutionary and forensic applications of 37 human Y chromosome SNPs" (PDF), Forensic Science International, 157 (1): 23:35, doi:10.1016/j.forsciint.2005.03.014, PMID 15896936
- Pelotti, S; Ceccardi, S; Lugaresi, F; Trane, R; Falconi, M; Bini, C; Willuweit, S; Roewer, L (2008), "Microgeographic genetic variation of Y chromosome in a population sample of Ravenna's area in the Emilia-Romagna region (North of Italy)", Forensic Science International: Genetics Supplement Series, 1 (1): 242, doi:10.1016/j.fsigss.2007.10.025
- Peričic; Lauc, LB; Klarić, IM; Rootsi, S; Janićijevic, B; Rudan, I; Terzić, R; Colak, I; et al. (2005), "High-resolution phylogenetic analysis of southeastern Europe traces major episodes of paternal gene flow among Slavic populations", Mol. Biol. Evol., vol. 22, no. 10, pp. 1964–75, doi:10.1093/molbev/msi185, PMID 15944443.
- Pontikos D. "Phylogeographic refinement of haplogroup E" http://dienekes.blogspot.ru/2015/07/phylogeographic-refinement-of.html
- Rosa; Ornelas, Carolina; Jobling, Mark A; Brehm, António; Villems, Richard (2007), "Y-chromosomal diversity in the population of Guinea-Bissau: a multiethnic perspective" (PDF), BMC Evolutionary Biology, 7 (1): 124, doi:10.1186/1471-2148-7-124, PMC 1976131, PMID 17662131
{{citation}}
: CS1 maint: unflagged free DOI (link) - Rosser, Z; Zerjal, T; Hurles, M; Adojaan, M; Alavantic, D; Amorim, A; Amos, W; Armenteros, M; et al. (2000), "Y-Chromosomal Diversity in Europe Is Clinal and Influenced Primarily by Geography, Rather than by Language", American Journal of Human Genetics, 67 (6): 1526–1543., doi:10.1086/316890, PMC 1287948, PMID 11078479
- Sanchez, JJ; Hallenberg, C; Børsting, C; Hernandez, A; Morling, N (2005), "High frequencies of Y chromosome lineages characterized by E3b1, DYS19-11, DYS392-12 in Somali males", European Journal of Human Genetics, 13 (7): 856–866, doi:10.1038/sj.ejhg.5201390, PMID 15756297. Published online 9 March 2005
- Scozzari; Cruciani, F; Pangrazio, A; Santolamazza, P; Vona, G; Moral, P; Latini, V; Varesi, L; et al. (2001), "Human Y-Chromosome Variation in the Western Mediterranean Area: Implications for the Peopling of the Region" (PDF), Human Immunology, 62 (9): 871–884, doi:10.1016/S0198-8859(01)00286-5, PMID 11543889.
- Semino; Passarino, G; Oefner, PJ; Lin, AA; Arbuzova, S; Beckman, LE; De Benedictis, G; Francalacci, P; et al. (2000), "The Genetic Legacy of Paleolithic Homo sapiens sapiens in Extant Europeans: A Y Chromosome Perspective" (PDF), Science, vol. 290, no. 5494, pp. 1155–59, doi:10.1126/science.290.5494.1155, PMID 11073453.
- Semino; Santachiarabenerecetti, A; Falaschi, F; Cavallisforza, L; Underhill, P (2002), "Ethiopians and Khoisan share the deepest clades of the human Y-chromosome phylogeny" (PDF), Am J Hum Genet, vol. 70, no. 1, pp. 265–268, doi:10.1086/338306, PMC 384897, PMID 11719903
- Semino; Magri, C; Benuzzi, G; Lin, A; Alzahery, N; Battaglia, V; MacCioni, L; Triantaphyllidis, C; et al. (2004), "Origin, Diffusion, and Differentiation of Y-Chromosome Haplogroups E and J: Inferences on the Neolithization of Europe and Later Migratory Events in the Mediterranean Area", American Journal of Human Genetics, vol. 74, no. 5, pp. 1023–1034, doi:10.1086/386295, PMC 1181965, PMID 15069642
- Shen; Lavi, Tal; Kivisild, Toomas; Chou, Vivian; Sengun, Deniz; Gefel, Dov; Shpirer, Issac; Woolf, Eilon; et al. (2004), "Reconstruction of Patrilineages and Matrilineages of Samaritans and Other Israeli Populations From Y-Chromosome and Mitochondrial DNA Sequence Variation" (PDF), Human Mutation, vol. 24, no. 3, pp. 248–60, doi:10.1002/humu.20077, PMID 15300852
- Thomas; Stumpf, M. P.H; Harke, H. (2006), "Evidence for an apartheid-like social structure in early Anglo-Saxon England" (PDF), Proceedings of the Royal Society B, 273 (273): 2651–2657, doi:10.1098/rspb.2006.3627, PMC 1635457, PMID 17002951
- Tillmar, Hisham Y.; Underhill, Peter A.; Cavalli-Sforza, Luca L.; Ibrahim, Muntaser E. (2009), "Population data of 12 Y-STR loci from a Somali population" (PDF), Forensic Science International: Genetics Supplement Series, 137 (3): 316, doi:10.1002/ajpa.20876, PMID 18618658
- Trombetta, Vincent; Cruciani, Beniamino; Sellitto, Fulvio; Scozzari, Daniele; Scozzari, Rosaria (2011), MacAulay, Vincent (ed.), "A New Topology of the Human Y Chromosome Haplogroup E1b1 (E-P2) Revealed through the Use of Newly Characterized Binary Polymorphisms", PLoS ONE, 6 (1): e16073, doi:10.1371/journal.pone.0016073, PMC 3017091, PMID 21253605
{{citation}}
: CS1 maint: unflagged free DOI (link) - Trombetta B. "Phylogeographic Refinement and Large Scale Genotyping of Human Y Chromosome Haplogroup E Provide New Insights into the Dispersal of Early Pastoralists in the African Continent" http://gbe.oxfordjournals.org/content/7/7/1940.long
- Underhill; Shen, Peidong; Lin, Alice A.; Jin, Li; Passarino, Giuseppe; Yang, Wei H.; Kauffman, Erin; Bonné-Tamir, Batsheva; et al. (2000), "Y chromosome sequence variation and the history of human populations", Nat Genet, vol. 26, no. 3, pp. 358–361, doi:10.1038/81685, PMID 11062480
- Underhill; Passarino, G.; Lin, A. A.; Shen, P.; Mirazon Lahr, M.; Foley, R. A.; Oefner, P. J.; Cavalli-Sforza, L. L. (2001), "The phylogeography of Y chromosome binary haplotypes and the origins of modern human populations" (PDF), Ann Hum Genet, vol. 65, no. 1, pp. 43–62, doi:10.1046/j.1469-1809.2001.6510043.x, PMID 11415522
- Underhill (2002), Bellwood and Renfrew (ed.), Inference of Neolithic Population Histories using Y-chromosome Haplotypes, Cambridge: McDonald Institute for Archaeological Research, ISBN 1-902937-20-1
{{citation}}
: Unknown parameter|booktitle=
ignored (help) - Underhill; Kivisild, T (2007), "Use of Y Chromosome and Mitochondrial DNA Population Structure in Tracing Human Migrations", Annu. Rev. Genet., 41 (1): 539–64, doi:10.1146/annurev.genet.41.110306.130407, PMID 18076332
- Weale; Zhivotovsky, LA; King, R; Mehdi, SQ; Edmonds, CA; Chow, CE; Lin, AA; Mitra, M; Sil, SK (2002), "Y Chromosome Evidence for Anglo-Saxon Mass Migration" (PDF), Mol. Biol. Evol., vol. 19, no. 7, pp. 1008–1021, doi:10.1093/oxfordjournals.molbev.a004160, PMID 12082121.
- Weale (September 1, 2003), "Rare Deep-Rooting Y Chromosome Lineages in Humans: Lessons for Phylogeography", Genetics, vol. 165, no. 1, pp. 229–234
- 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
- Zalloua; Platt, D; Elsibai, M; Khalife, J; Makhoul, N; Haber, M; Xue, Y; Izaabel, H; et al. (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