Haplogroup E-Z827

Haplogroup E-Z827
Possible time of origin approx 24,100 years BP [1]
Possible place of origin Northern Africa[2]
Ancestor E-M215/M35
Descendants E-L19, E-Z830
Defining mutations Z827

E-Z827, also known as E1b1b1b,[3] is a major human Y-chromosome DNA haplogroup. It is the parent lineage to the E-Z830 and E-V257 subclades, and defines their common phylogeny. The former is predominantly found in the Horn of Africa and the Middle East; the latter is most frequently observed in North Africa, with its E-M81 subclade observed among the ancient Guanche natives of the Canary Islands. E-Z827 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.[4][5][6]

  • E-Z827 (Z827) - E1b1b1b[7]
    • E-V257/L19 (L19, V257) - E1b1b1b1[7]
      • E-PF2431 (PF2431)[8]
        • E-PF2438
          • E-Y10561
            • E-FGC18981
              • E-FGC38527
              • E-Y35933
              • E-FGC18960
                • E-Y33020
                • E-FGC18958
          • E-PF2440
            • E-PF2471
              • E-BY9805
      • E-M81 (M81)[9]
        • E-M81*
        • E-PF2546
          • E-PF2546*
          • E-CTS12227
            • E-MZ11
              • E-MZ12
          • 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-Z830 (Z830) - E1b1b1b2[7]
      • E-M123 (M123)
        • E-M34 (M34)
          • E-M84 (M84)
            • E-M136 (M136)
          • E-M290 (M290)
          • E-V23 (V23)
          • E-L791 (L791,L792)
      • 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-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.

—  [10]
Distribution of E-PF2431

PF2431 is the sister branch of M81 which was discovered in Paolo Francalacci (2011). Previously, it was designated L19*/V257*. This mutation has been discovered in North Africa (mainly in Souss in Morocco and West Nile in Egypt), the Sahel (Chad, Niger, Gambia), Western Europe (United Kingdom (Derbyshire), Germany, Switzerland, Spain, Italy) and Near Eastern (Karabakh and Urmia). It would have formed 13800 years ago and is thought to originate from the "green" Sahara. Its TMRCA is estimated at 10600 years by yfull.

E-V257's dominant sub-clade E-M81 is thought to have originated in the area of North Africa 14,000 years ago, but all Yfull members are M183 (because there are only 7 Z827 Moroccans) so have a TMRCA just 2700 years ago.[11]

Distribution of E1b1b1b1a in select areas of Africa, Asia and Europe

E-M81 is the most common subclade of haplogroup E-L19/V257. It is concentrated in the north africa, and is dominated by its E-M183 subclade. E-M183 is believed to have originated in northwestern Africa, and has an estimated age of 2284-2984 ybp.[12]

The E-M183 subhaplogroup reaches a mean frequency of 42% in North Africa. It decreases in frequency from 100% in some isolated Berber populations to approximately 28.6% to the east of this range in Egypt.[13][14][15] Because of its prevalence among these groups and also others such as Mozabite, Middle Atlas, Kabyle and other Berber groups, and coincidence with language dispersal age, it is sometimes referred to as a genetic "Berber marker". High levels among two Tuareg populations inhabiting the Sahara: 77.8% near Gorom-Gorom, in Burkina Faso, and 81.8% from Gossi in Mali are observed.[16] There was a much lower frequency of 11.1% in the vicinity of Tanut in the Republic of Niger.

The E-M81 subclade is also quite common among North African Arabic-speaking groups. It 31.5% among Moroccan Arabs,[17] and is generally found at frequencies around 45% in coastal cities of the Maghreb (Oran, Tunis, Tizi Ouzou, Algiers).[13][18]

In this key area from Egypt to the Atlantic Ocean,[13] report a pattern of decreasing STR haplotype variation (implying decreasing lineage age in those areas) from East to West (but[19] reports West to East for M183), accompanied by a substantial increasing frequency. At the eastern extreme of this core range,[15] M81 is found in 28.6% (10 out of 35 men) in El-Hayez in the Western desert in Egypt

The pattern of distribution and variance to be consistent with the hypothesis of a post Paleolithic "demic diffusion" from the East.[13] 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". 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. 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.[20]

The E-M81 subclade has been found in ancient Guanche (Bimbapes) fossils excavated in Punta Azul, El Hierro, Canary Islands, which are dated to the 10th century (~44%).[21]

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,[22] 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)[22] 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.[22][23][24][25][26] The highest frequencies of this clade found so far in Europe were observed in the Pasiegos from Cantabria, ranging from 18% (8/45)[26] to 41% (23/56).[17] 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%).[27]

E-M81 is also found in other parts of Western Europe, such as Britain – especially Wales and Scotland – and France, where it has an overall incidence of 2.7% (15/555), with frequencies surpassing 5.0% in Auvergne (5/89) and Île-de-France (5/91).[28][29] (There is limited evidence that the data with regard to ethnic French males excludes modern migration into France. For example, E-M81 occurred among men with surnames with French origins in Sicily at a rate of 2.0%, and up to 7.0% in towns such as Piazza Armerina.)[30] It occurs at slightly lower frequencies in continental Italy (especially near Lucera)[25] 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.[31]

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, (8 out of 132),[32] 5.4% in Brazil (Rio de Janeiro) (6 out of 112), "The presence of chromosomes of North African origin (E3b1b-M81;[17] can also be explained by a Portuguese-mediated influx, since this haplogroup reaches a frequency of 5.6% in Portugal,[24] quite similar to the frequency found in Rio de Janeiro (5.4%) among European contributors."[33] and among Hispanic men from California and Hawaii 2.4% (7 out of 295),[34]

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.

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 [35]
Algeria Mozabite Berbers 20 80 [17]
Algeria Oran 102 45.1 [18]
Algeria Algiers 35 42.9 [13]
Algeria Kabyles from Tizi Ouzou 19 47.4 [13]
Algeria Arabs and Berbers 156 44.2 [36]
Brazil Rio de Janeiro 112 5.4 [33]
Burkina Faso Tuaregs 38 77.8 [16]
Canary Islands Fuerteventura 75 13.3 [27]
Canary Islands Gran Canaria 78 11.5 [27]
Canary Islands Tenerife 178 10.7 [27]
Canary Islands Lanzarote 97 6.2 [27]
Canary Islands La Palma 85 5.9 [27]
Canary Islands Gomera 92 4.4 [27]
Canary Islands Hierro 47 2.1 [27]
Cuba 132 6.1 [32]
Cyprus Turkish Cypriots 46 8.7 [17]
Egypt Northern Egyptians 21 4.8 [17]
Egypt Western Desert 35 28.6 [15]
Egypt 147 8.2 [23]
Egypt Arabs 370 11.8 [36]
France 85 3.5 [17]
France Auvergne 89 5.6 [28]
France Île-de-France 91 5.5 [28]
France Nord-Pas-de-Calais 68 4.4 [28]
France Provence-Alpes-CĂŽte d'Azur 45 2.2 [28]
France Midi-Pyrénées 67 1.5 [28]
France BĂ©arnais 56 1.8 [29]
France Bigorre 44 2.3 [29]
Iberia Spain, Portugal 655 5.2 [27]
Iberia Spain, Portugal 1140 4.3 [22]
Israel Bedouins 28 3.6 [17]
Italy Central Italians 89 2.2 [17]
Italy Northern Italians 67 1.5 [17]
Italy North-West Apulia 46 4.3 [25]
Italy East Campania 84 2.4 [25]
Italy Veneto 55 1.8 [25]
Italy North-East Latium 55 1.8 [25]
Italy Lucera 60 1.7 [25]
Italy Sicily 236 2.1 [30]
Italy Sardinia 1204 5.8 [31]
Jordania 101 4 [23]
Lebanon 104 1.9 [23]
Lebanon 914 1.2 [37]
Libya Tuaregs 47 48.9 [38]
Libya Arabs 215 35.9 [39]
Libya Arabs and Berbers 83 45.7 [36]
Mali Tuaregs (Gozi) 21 81.8 [16]
Mauritania Arabs and Berbers 189 55.5 [36]
Morocco Marrakesh Berbers 29 72.4 [17]
Morocco Southern Moroccan Berbers 187 98.5 [40]
Morocco Moyen Atlas Berbers 69 71 [17]
Morocco Moroccan Arabs 54 31.5 [17]
Morocco Marrakesh (Amizmiz Valley) 33 84.8 [14]
Morocco Northern Moroccans (Beni Snassen) 67 79.1 [35]
Morocco Northern Moroccans (Rhiraya) 54 79.6 [35]
Morocco Immigrants resident in Italy 51 54.9 [41]
Morocco Arabs and Berbers 221 65 [27]
Morocco Arabs and Berbers 760 67.3 [36]
Niger Tuaregs 22 9.1 [17]
Niger Tuaregs 31 11.1 [16]
North Africa Sahara 89 59.6 [27]
North Africa Algeria, Tunisia 202 39.1 [27]
Portugal North 109 5.5 [42]
Portugal South 49 12.2 [17]
Portugal North 50 4 [17]
Portugal South 78 7.7 [22]
Portugal North 60 3.3 [22]
Portugal 303 5.6 [43]
Portugal North 101 6 [43]
Portugal Center 102 4.9 [43]
Portugal South 100 6 [43]
Portugal Madeira 129 5.4 [43]
Portugal Açores 121 5 [43]
Portugal 657 5.6 [24]
Portugal Entre Douro e Minho 228 6.6 [24]
Portugal Tras os Montes 64 3.1 [24]
Portugal Beira Litoral 116 5.2 [24]
Portugal Beira Interior 58 5.3 [24]
Portugal Estremadura 43 4.6 [24]
Portugal Lisboa e Setubal 62 6.5 [24]
Portugal Alentejo 65 7.7 [24]
Portugal Coruche 64 9.4 [16]
Portugal Pias 46 4.3 [16]
Portugal Alcacer do Sal 21 4.8 [16]
Portugal Tras-os-Montes (Jews) 57 5.3 [44]
Portugal Tras-os-Montes (Non Jews) 30 10 [44]
Somalia 201 1.5 [23]
Spain Pasiegos from Cantabria 19 36.8 [45]
Spain Pasiegos from Cantabria 56 41.1 [17]
Spain Pasiegos from Cantabria 45 17.8 [26]
Spain Spanish Basques 55 3.6 [17]
Spain Asturians 90 2.2 [17]
Spain Southern Spaniards 62 1.6 [17]
Spain Castile, NorthWest 100 10 [22]
Spain Andalucia, West 73 9.6 [22]
Spain Galicia 19 10.5 [42]
Spain Galicia 292 4.1 [46]
Spain Galicia 88 9.1 [22]
Spain Galicia 44 9.1 [47]
Spain Galicia 164 9.1 [48]
Spain Extremadura 52 7.7 [22]
Spain Valencia 73 4.1 [22]
Spain Castile, NorthEast 31 3.2 [22]
Spain Aragon 34 2.9 [22]
Spain Minorca 37 2.7 [22]
Spain Andalucia, East 95 2.1 [22]
Spain Majorca 62 1.6 [22]
Spain Castile, La Mancha 63 1.6 [22]
Spain Catalonia 80 1.3 [22]
Spain Catalonia 111 3.6 [47]
Spain Cantabria 161 13 [25]
Spain Malaga 26 11.5 [42]
Spain Cantabria 70 8.6 [42]
Spain Cordoba 27 7.4 [42]
Spain Valencia 31 6.5 [42]
Spain Valencia 59 5.1 [47]
Spain Almeria 36 5.6 [47]
Spain Leon 60 5 [42]
Spain Castile 21 4.8 [42]
Spain Seville 155 4.5 [42]
Spain Huelva 22 4.5 [42]
Spain Basques 45 2.2 [42]
Spain Huelva 167 3 [49]
Spain Granada 250 3.6 [49]
Spain Pedroches Valley 68 1.5 [14]
Spain Andalusians 94 2.1 [14]
Spain Zamora 235 5.5 [50]
Tunisia Tunis 148 37.9 [13]
Tunisia Immigrants resident in Italy 52 32.7 [41]
Tunisia Berbers from Bou Omrane 40 87.5 [51]
Tunisia Berbers from Bou Saad 40 92.5 [51]
Tunisia Jerbian Arabs 46 60.9 [51]
Tunisia Jerbian Berbers 47 76.6 [51]
Tunisia Berbers from Chenini–Douiret 27 100 [52]
Tunisia Berbers from Sened 35 65.7 [52]
Tunisia Berbers from Jradou 32 100 [52]
Tunisia Andalusian Zaghouan 32 40.6 [52]
Tunisia Cosmopolitan Tunis 33 54.4 [52]
Tunisia Arabs and Berbers 601 62.7 [36]
Turkey Istanbul Turkish 35 5.7 [17]
Turkey Sephardi Turkish 19 5.3 [17]
Turkey Southwestern Turkish 40 2.5 [17]
Turkey Northeastern Turkish 41 2.4 [17]

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.[53][54][55][56]

E-M123 is mostly known for its major subclade E-M34, which dominates this clade.[57]

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.[2]

We observed the highest frequency and diversity of this haplogroup in the northern part of the Horn of Africa (present day Eritrea and northern Ethiopia), where the majority of the deepest E-V1515 subhaplogroups and paragroups were found. In the southern part of the Horn (southern Ethiopia, Somalia and northern Kenya), haplogroup E-V1515 is almost exclusively represented by the recent (3.5 ka; 95% CI: 1.7–5.9 ka) subhaplogroup E-V1486. Further south, in southern Kenya and southern Africa, a single E-V1486 terminal clade, known as E-M293 (Henn et al. 2008), was found (fig. 3). This phylogeographic pattern is strongly suggestive of human movements from the northern part of the Horn to the Ethiopian/Kenyan borders between 12 ka (the coalescence of E-V1515) and 3.5 ka (the coalescence of E-V1486), and from here toward southern Africa across the equatorial belt in more recent times.[2]

Multiple instances of commercially observed E-V1515 have also been detected in Arabia.[58]

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 and is associated with the spread of pastoralism from Eastern Africa into Southern Africa.[59] 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%). Two Bantu-speaking Kenyan males were found with the M293 mutation.[59] 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.[5] This is a subclade of E-M293.[10]

E-V42 was discovered in two Ethiopian Jews.[10] 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.[60]

The E-V6 subclade of E-V1515 is defined by V6 and has been identified a significant presence of these lineages in Ethiopia, and also some in the neighboring Somali population.[17] Among the Ethiopian and Somali samples, the highest were 14.7% among the Ethiopian Amhara, and 16.7% among the Ethiopian Wolayta.

E-V92 was discovered in two Ethiopian Amhara.[10] Like E-V6 and E-V42 it possibly only exists in the area of Ethiopia.


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) (ÎČ) (Îł) (ÎŽ) (Δ) (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

The following research teams per their publications were represented in the creation of the YCC Tree.

See also


Y-DNA E Subclades

Y-DNA Backbone Tree

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
F1  F2  F3  GHIJK
I   J     LT [χ 5]       K2 [χ 6]
L     T    K2a [χ 7]        K2b [χ 8]     K2c     K2d K2e [χ 9]  
K-M2313 [χ 10]     K2b1 [χ 11] P [χ 12]
NO   S [χ 13]  M [χ 14]    P1     P2


  1. ^ "YFull YTree v6.02". YFull: Y-Chr Sequence Interpretation Service.
  2. ^ a b c Trombetta B, D'Atanasio E, Massaia A, Ippoliti M, Coppa A, Candilio F, et al. (June 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". Genome Biology and Evolution. 7 (7): 1940–50. doi:10.1093/gbe/evv118. PMC 4524485. PMID 26108492.
  3. ^ ISOGG (2015), Y-DNA Haplogroup E and its Subclades - 2015
  4. ^ ISOGG (2008). "Y-DNA Haplogroup E and its Subclades - 2008". International Society of Genetic Genealogists "ISOGG". Cite journal requires |journal= (help)
  5. ^ a b c Karafet TM, Mendez FL, Meilerman MB, Underhill PA, Zegura SL, Hammer MF (May 2008). "New binary polymorphisms reshape and increase resolution of the human Y chromosomal haplogroup tree". Genome Research. 18 (5): 830–8. doi:10.1101/gr.7172008. PMC 2336805. PMID 18385274.
  6. ^ Y Chromosome Consortium "YCC" (February 2002). "A nomenclature system for the tree of human Y-chromosomal binary haplogroups". Genome Research. 12 (2): 339–48. doi:10.1101/gr.217602. PMC 155271. PMID 11827954.
  7. ^ a b c ISOGG 2015
  8. ^ "E-PF2431 YTree".
  9. ^ "E-M81 YTree". www.yfull.com. Retrieved 2016-07-09.
  10. ^ a b c d Trombetta B, Cruciani F, Sellitto D, Scozzari R (January 2011). MacAulay V (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. Bibcode:2011PLoSO...616073T. doi:10.1371/journal.pone.0016073. PMC 3017091. PMID 21253605.
  11. ^ "E-M81 YTree". www.yfull.com. Retrieved 2016-07-10.
  12. ^ SolĂ©-Morata N, GarcĂ­a-FernĂĄndez C, Urasin V, Bekada A, Fadhlaoui-Zid K, Zalloua P, Comas D, Calafell F (November 2017). "Whole Y-chromosome sequences reveal an extremely recent origin of the most common North African paternal lineage E-M183 (M81)". Scientific Reports. 7 (1): 15941. Bibcode:2017NatSR...715941S. doi:10.1038/s41598-017-16271-y. PMC 5698413. PMID 29162904.
  13. ^ a b c d e f g Arredi B, Poloni ES, Paracchini S, Zerjal T, Fathallah DM, Makrelouf M, Pascali VL, Novelletto A, Tyler-Smith C (August 2004). "A predominantly neolithic origin for Y-chromosomal DNA variation in North Africa". American Journal of Human Genetics. 75 (2): 338–45. doi:10.1086/423147. PMC 1216069. PMID 15202071.
  14. ^ a b c d Alvarez L, Santos C, Montiel R, Caeiro B, Baali A, Dugoujona JM, Dugoujon JM, Aluja MP (2009). "Y-chromosome variation in South Iberia: insights into the North African contribution". American Journal of Human Biology. 21 (3): 407–9. doi:10.1002/ajhb.20888. PMID 19213004. S2CID 7041905.
  15. ^ a b c KujanovĂĄ M, Pereira L, Fernandes V, Pereira JB, CernĂœ V (October 2009). "Near eastern neolithic genetic input in a small oasis of the Egyptian Western Desert". American Journal of Physical Anthropology. 140 (2): 336–46. doi:10.1002/ajpa.21078. PMID 19425100.
  16. ^ a b c d e f g Pereira L, CernĂœ V, Cerezo M, Silva NM, HĂĄjek M, VasĂ­kovĂĄ A, KujanovĂĄ M, Brdicka R, Salas A (August 2010). "Linking the sub-Saharan and West Eurasian gene pools: maternal and paternal heritage of the Tuareg nomads from the African Sahel". European Journal of Human Genetics. 18 (8): 915–23. doi:10.1038/ejhg.2010.21. PMC 2987384. PMID 20234393.
  17. ^ a b c d e f g h i j k l m n o p q r s t u v w x y Cruciani F, La Fratta R, Santolamazza P, Sellitto D, Pascone R, Moral P, Watson E, Guida V, Colomb EB, Zaharova B, Lavinha J, Vona G, Aman R, Cali F, Akar N, Richards M, Torroni A, Novelletto A, Scozzari R (May 2004). "Phylogeographic analysis of haplogroup E3b (E-M215) y chromosomes reveals multiple migratory events within and out of Africa". American Journal of Human Genetics. 74 (5): 1014–22. doi:10.1086/386294. PMC 1181964. PMID 15042509.
  18. ^ a b Robino C, Crobu F, Di Gaetano C, Bekada A, Benhamamouch S, Cerutti N, Piazza A, Inturri S, Torre C (May 2008). "Analysis of Y-chromosomal SNP haplogroups and STR haplotypes in an Algerian population sample". International Journal of Legal Medicine. 122 (3): 251–5. doi:10.1007/s00414-007-0203-5. PMID 17909833. S2CID 11556974.
  19. ^ SolĂ©-Morata, Neus; GarcĂ­a-FernĂĄndez, Carla; Urasin, Vadim; Bekada, Asmahan; Fadhlaoui-Zid, Karima; Zalloua, Pierre; Comas, David; Calafell, Francesc (21 November 2017). "Whole Y-chromosome sequences reveal an extremely recent origin of the most common North African paternal lineage E-M183 (M81)". Scientific Reports. 7 (1): 15941. Bibcode:2017NatSR...715941S. doi:10.1038/s41598-017-16271-y. PMC 5698413. PMID 29162904.
  20. ^ Keita SO (2008). "Geography, selected Afro-Asiatic families, and Y chromosome lineage variation". In Bengtson JD (ed.). In hot pursuit of language in prehistory: essays in the four fields of anthropology: in honor of Harold Crane Fleming. Amsterdam: John Benjamins Pub. ISBN 978-90-272-3252-6.
  21. ^ Ordóñez AC, Fregel R, Trujillo-Mederos A, Hervella M, de-la-RĂșa C, Arnay-de-la-Rosa M (2017). "Genetic studies on the prehispanic population buried in Punta Azul cave (El Hierro, Canary Islands)". Journal of Archaeological Science. 78: 20–28. doi:10.1016/j.jas.2016.11.004.
  22. ^ a b c d e f g h i j k l m n o p q r Adams SM, Bosch E, Balaresque PL, Ballereau SJ, Lee AC, Arroyo E, LĂłpez-Parra AM, Aler M, Grifo MS, Brion M, Carracedo A, Lavinha J, MartĂ­nez-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 (6): 725–36. doi:10.1016/j.ajhg.2008.11.007. PMC 2668061. PMID 19061982.
  23. ^ a b c d e Flores C, Maca-Meyer N, Larruga JM, Cabrera VM, Karadsheh N, Gonzalez AM (2005). "Isolates in a corridor of migrations: a high-resolution analysis of Y-chromosome variation in Jordan". Journal of Human Genetics. 50 (9): 435–41. doi:10.1007/s10038-005-0274-4. PMID 16142507.
  24. ^ a b c d e f g h i j Beleza S, GusmĂŁo L, Lopes A, Alves C, Gomes I, Giouzeli M, Calafell F, Carracedo A, Amorim A (March 2006). "Micro-phylogeographic and demographic history of Portuguese male lineages". Annals of Human Genetics. 70 (Pt 2): 181–94. doi:10.1111/j.1529-8817.2005.00221.x. PMID 16626329. S2CID 4652154.
  25. ^ a b c d e f g h Capelli C, Onofri V, Brisighelli F, Boschi I, Scarnicci F, Masullo M, et al. (June 2009). "Moors and Saracens in Europe: estimating the medieval North African male legacy in southern Europe". European Journal of Human Genetics. 17 (6): 848–52. doi:10.1038/ejhg.2008.258. PMC 2947089. PMID 19156170.
  26. ^ a b c Maca-Meyer N, SĂĄnchez-Velasco P, Flores C, Larruga JM, GonzĂĄlez AM, Oterino A, Leyva-CobiĂĄn F (July 2003). "Y chromosome and mitochondrial DNA characterization of Pasiegos, a human isolate from Cantabria (Spain)". Annals of Human Genetics. 67 (Pt 4): 329–39. CiteSeerX doi:10.1046/j.1469-1809.2003.00045.x. PMID 12914567. S2CID 40355653.
  27. ^ a b c d e f g h i j k l Fregel R, Gomes V, GusmĂŁo L, GonzĂĄlez AM, Cabrera VM, Amorim A, Larruga JM (August 2009). "Demographic history of Canary Islands male gene-pool: replacement of native lineages by European". BMC Evolutionary Biology. 9: 181. doi:10.1186/1471-2148-9-181. PMC 2728732. PMID 19650893.
  28. ^ a b c d e f Ramos-Luis E, Blanco-Verea A, Brión M, Van Huffel V, Carracedo A, Sánchez-Diz P (December 2009). "Phylogeography of French male lineages". Forensic Science International: Genetics Supplement Series. 2 (1): 439–441. doi:10.1016/j.fsigss.2009.09.026.
  29. ^ a b c MartĂ­nez-Cruz B, Harmant C, Platt DE, Haak W, Manry J, Ramos-Luis E, Soria-Hernanz DF, Bauduer F, Salaberria J, Oyharçabal B (March 2012). "Evidence of Pre-Roman Tribal Genetic Structure in Basques from Uniparentally Inherited Markers". Molecular Biology and Evolution. 29 (9): 2211–2222. doi:10.1093/molbev/mss091. PMID 22411853.
  30. ^ a b Di Gaetano C, Cerutti N, Crobu F, Robino C, Inturri S, Gino S, et al. (January 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): 91–9. doi:10.1038/ejhg.2008.120. PMC 2985948. PMID 18685561.
  31. ^ a b Francalacci P, Morelli L, Angius A, Berutti R, Reinier F, Atzeni R, et al. (August 2013). "Low-pass DNA sequencing of 1200 Sardinians reconstructs European Y-chromosome phylogeny". Science. 341 (6145): 565–9. Bibcode:2013Sci...341..565F. doi:10.1126/science.1237947. PMC 5500864. PMID 23908240.
  32. ^ a b Mendizabal I, Sandoval K, Berniell-Lee G, Calafell F, Salas A, MartĂ­nez-Fuentes A, Comas D (July 2008). "Genetic origin, admixture, and asymmetry in maternal and paternal human lineages in Cuba". BMC Evolutionary Biology. 8: 213. doi:10.1186/1471-2148-8-213. PMC 2492877. PMID 18644108.
  33. ^ a b Silva DA, Carvalho E, Costa G, Tavares L, Amorim A, GusmĂŁo L (2006). "Y-chromosome genetic variation in Rio de Janeiro population". American Journal of Human Biology. 18 (6): 829–37. doi:10.1002/ajhb.20567. PMID 17039481. S2CID 23778828.
  34. ^ Paracchini S, Pearce CL, Kolonel LN, Altshuler D, Henderson BE, Tyler-Smith C (November 2003). "A Y chromosomal influence on prostate cancer risk: the multi-ethnic cohort study". Journal of Medical Genetics. 40 (11): 815–9. doi:10.1136/jmg.40.11.815. PMC 1735314. PMID 14627670.
  35. ^ a b c Dugoujon JM, Philippson G (2005). "The Berbers: Linguistic and genetic diversity" (PDF). Archived from the original (PDF) on 2012-03-25.
  36. ^ a b c d e f Bekada A, Fregel R, Cabrera VM, Larruga JM, Pestano J, Benhamamouch S, GonzĂĄlez AM (February 2013). "Introducing the Algerian mitochondrial DNA and Y-chromosome profiles into the North African landscape". PLOS ONE. 8 (2): e56775. Bibcode:2013PLoSO...856775B. doi:10.1371/journal.pone.0056775. PMC 3576335. PMID 23431392.
  37. ^ Genographic Consortium, Zalloua PA, Platt DE, El Sibai M, Khalife J, Makhoul N, et al. (November 2008). "Identifying genetic traces of historical expansions: Phoenician footprints in the Mediterranean". American Journal of Human Genetics. 83 (5): 633–42. doi:10.1016/j.ajhg.2008.10.012. PMC 2668035. PMID 18976729.
  38. ^ Ottoni C, Larmuseau MH, Vanderheyden N, MartĂ­nez-Labarga C, Primativo G, Biondi G, Decorte R, Rickards O (May 2011). "Deep into the roots of the Libyan Tuareg: a genetic survey of their paternal heritage". American Journal of Physical Anthropology. 145 (1): 118–24. doi:10.1002/ajpa.21473. PMID 21312181.
  39. ^ Fadhlaoui-Zid K, Haber M, MartĂ­nez-Cruz B, Zalloua P, Benammar Elgaaied A, Comas D (2013). "Genome-wide and paternal diversity reveal a recent origin of human populations in North Africa". PLOS ONE. 8 (11): e80293. Bibcode:2013PLoSO...880293F. doi:10.1371/journal.pone.0080293. PMC 3842387. PMID 24312208.
  40. ^ Reguig A, Harich N, Barakat A, Rouba H (2014). "Phylogeography of E1b1b1b-M81 haplogroup and analysis of its subclades in Morocco". Human Biology. 86 (2): 105–12. doi:10.3378/027.086.0204. PMID 25397701. S2CID 11334895.
  41. ^ a b Onofri V, Alessandrini F, Turchi C, Pesaresi M, Tagliabracci A (August 2008). "Y-chromosome markers distribution in Northern Africa: High-resolution SNP and STR analysis in Tunisia and Morocco populations". Forensic Science International: Genetics Supplement Series. 1 (1): 235–236. doi:10.1016/j.fsigss.2007.10.173.
  42. ^ a b c d e f g h i j k Flores C, Maca-Meyer N, GonzĂĄlez AM, Oefner PJ, Shen P, PĂ©rez JA, Rojas A, Larruga JM, Underhill PA (October 2004). "Reduced genetic structure of the Iberian peninsula revealed by Y-chromosome analysis: implications for population demography". European Journal of Human Genetics. 12 (10): 855–63. doi:10.1038/sj.ejhg.5201225. PMID 15280900.
  43. ^ a b c d e f Gonçalves R, Freitas A, Branco M, Rosa A, Fernandes AT, Zhivotovsky LA, Underhill PA, Kivisild T, Brehm A (July 2005). "Y-chromosome lineages from Portugal, Madeira and Açores record elements of Sephardim and Berber ancestry". Annals of Human Genetics. 69 (Pt 4): 443–54. doi:10.1111/j.1529-8817.2005.00161.x. PMID 15996172. S2CID 3229760.
  44. ^ a b Nogueiro I, Manco L, Gomes V, Amorim A, GusmĂŁo L (March 2010). "Phylogeographic analysis of paternal lineages in NE Portuguese Jewish communities". American Journal of Physical Anthropology. 141 (3): 373–81. doi:10.1002/ajpa.21154. PMID 19918998.
  45. ^ Scozzari R, Cruciani F, Pangrazio A, Santolamazza P, Vona G, Moral P, Latini V, Varesi L, Memmi MM, Romano V, De Leo G, Gennarelli M, Jaruzelska J, Villems R, Parik J, Macaulay V, Torroni A (September 2001). "Human Y-chromosome variation in the western Mediterranean area: implications for the peopling of the region" (PDF). Human Immunology. 62 (9): 871–84. CiteSeerX doi:10.1016/S0198-8859(01)00286-5. PMID 11543889.
  46. ^ Brion M, Quintans B, Zarrabeitia M, Gonzalez-Neira A, Salas A, Lareu V, Tyler-Smith C, Carracedo A (March 2004). "Micro-geographical differentiation in Northern Iberia revealed by Y-chromosomal DNA analysis". Gene. 329: 17–25. doi:10.1016/j.gene.2003.12.035. PMID 15033525.
  47. ^ a b c d Santos C, Fregel R, Cabrera VM, Alvarez L, Larruga JM, Ramos A, LĂłpez MA, Pilar Aluja M, GonzĂĄlez AM (2014). "Mitochondrial DNA and Y-chromosome structure at the Mediterranean and Atlantic façades of the Iberian Peninsula". American Journal of Human Biology. 26 (2): 130–41. doi:10.1002/ajhb.22497. PMID 24375863. S2CID 205303141.
  48. ^ Regueiro M, Garcia-Bertrand R, Fadhlaoui-Zid K, Álvarez J, Herrera RJ (June 2015). "From Arabia to Iberia: A Y chromosome perspective". Gene. 564 (2): 141–52. doi:10.1016/j.gene.2015.02.042. PMID 25701402.
  49. ^ a b Ambrosio B, Dugoujon JM, HernĂĄndez C, De La Fuente D, GonzĂĄlez-MartĂ­n A, Fortes-Lima CA, Novelletto A, RodrĂ­guez JN, CalderĂłn R (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. S2CID 1667431.
  50. ^ Alvarez L, Ciria E, Marques SL, Santos C, Aluja MP (2014). "Y-chromosome analysis in a Northwest Iberian population: unraveling the impact of Northern African lineages". American Journal of Human Biology. 26 (6): 740–6. doi:10.1002/ajhb.22602. PMID 25123837. S2CID 205303372.
  51. ^ a b c d Ennafaa H, Fregel R, Khodjet-El-Khil H, GonzĂĄlez AM, Mahmoudi HA, Cabrera VM, Larruga JM, Benammar-ElgaaĂŻed A (October 2011). "Mitochondrial DNA and Y-chromosome microstructure in Tunisia". Journal of Human Genetics. 56 (10): 734–41. doi:10.1038/jhg.2011.92. PMID 21833004.
  52. ^ a b c d e Fadhlaoui-Zid K, Martinez-Cruz B, Khodjet-el-khil H, Mendizabal I, Benammar-Elgaaied A, Comas D (October 2011). "Genetic structure of Tunisian ethnic groups revealed by paternal lineages". American Journal of Physical Anthropology. 146 (2): 271–80. doi:10.1002/ajpa.21581. PMID 21915847.
  53. ^ "E-M35 Project Data". haplozone.net. Archived from the original on 2015-09-24. Retrieved 2012-05-22.
  54. ^ "E-M35 Project Data". haplozone.net. Archived from the original on 2015-09-24. Retrieved 2012-05-22.
  55. ^ "E-M35 Project Data". haplozone.net. Archived from the original on 2015-09-24. Retrieved 2012-05-22.
  56. ^ "E-M35 Project Data". haplozone.net. Archived from the original on 2015-09-24. Retrieved 2012-05-22.
  57. ^ "E-M35 phylogeny project". 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.[permanent dead link]
  58. ^ "E-CTS10880 YTree".
  59. ^ a b Henn BM, Gignoux C, Lin AA, Oefner PJ, Shen P, Scozzari R, Cruciani F, Tishkoff SA, Mountain JL, Underhill PA (August 2008). "Y-chromosomal evidence of a pastoralist migration through Tanzania to southern Africa". Proceedings of the National Academy of Sciences of the United States of America. 105 (31): 10693–8. Bibcode:2008PNAS..10510693H. doi:10.1073/pnas.0801184105. PMC 2504844. PMID 18678889.
  60. ^ "E-M35 Project Data". haplozone.net. Archived from the original on 2015-09-24. Retrieved 2013-01-16.
  61. ^ Jobling MA, Tyler-Smith C (August 2000). "New uses for new haplotypes the human Y chromosome, disease and selection". Trends in Genetics. 16 (8): 356–62. doi:10.1016/S0168-9525(00)02057-6. PMID 10904265.
  62. ^ Kalaydjieva L, Calafell F, Jobling MA, Angelicheva D, de Knijff P, Rosser ZH, Hurles ME, Underhill P, Tournev I, Marushiakova E, Popov V (February 2001). "Patterns of inter- and intra-group genetic diversity in the Vlax Roma as revealed by Y chromosome and mitochondrial DNA lineages". European Journal of Human Genetics. 9 (2): 97–104. doi:10.1038/sj.ejhg.5200597. PMID 11313742.
  63. ^ Underhill PA, Shen P, Lin AA, Jin L, Passarino G, Yang WH, et al. (November 2000). "Y chromosome sequence variation and the history of human populations". Nature Genetics. 26 (3): 358–61. doi:10.1038/81685. PMID 11062480. S2CID 12893406.
  64. ^ Hammer MF, Karafet TM, Redd AJ, Jarjanazi H, Santachiara-Benerecetti S, Soodyall H, Zegura SL (July 2001). "Hierarchical patterns of global human Y-chromosome diversity". Molecular Biology and Evolution. 18 (7): 1189–203. doi:10.1093/oxfordjournals.molbev.a003906. PMID 11420360.
  65. ^ Semino O, Passarino G, Oefner PJ, Lin AA, Arbuzova S, Beckman LE, et al. (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.
  66. ^ Su B, Xiao J, Underhill P, Deka R, Zhang W, Akey J, Huang W, Shen D, Lu D, Luo J, Chu J, Tan J, Shen P, Davis R, Cavalli-Sforza L, Chakraborty R, Xiong M, Du R, Oefner P, Chen Z, Jin L (December 1999). "Y-Chromosome evidence for a northward migration of modern humans into Eastern Asia during the last Ice Age". American Journal of Human Genetics. 65 (6): 1718–24. doi:10.1086/302680. PMC 1288383. PMID 10577926.
  67. ^ Capelli C, Wilson JF, Richards M, Stumpf MP, Gratrix F, Oppenheimer S, Underhill P, Pascali VL, Ko TM, Goldstein DB (February 2001). "A predominantly indigenous paternal heritage for the Austronesian-speaking peoples of insular Southeast Asia and Oceania". American Journal of Human Genetics. 68 (2): 432–43. doi:10.1086/318205. PMC 1235276. PMID 11170891.

Further reading

External links