Haplogroup E-Z827: Difference between revisions

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

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Revision as of 06:16, 26 May 2018

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.
— Trombetta (2011)

E-PF2431

File:E-PF2431.jpg

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), and Western Europe (United Kingdom (Derbyshire), Germany, Switzerland, Spain, Italy).

E-V257's dominant sub-clade E-M81 is thought to have originated in the area of North Africa 14,200 years ago.^[10]

E-M81

E-M81 is the most common subclade of haplogroup E-L19/V257. It is concentrated in the Tamazgha, 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.^[11]

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.^[12]^[13]^[14]^[15] 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". 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 reaches 75% among Moroccans Arabs,^[17] and is generally found at frequencies around 45% in coastal cities of the Maghreb (Oran, Tunis, Tizi Ouzou, Algiers).^[12]^[18]

In this key area from Egypt to the Atlantic Ocean,^[12] 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

The pattern of distribution and variance to be consistent with the hypothesis of a post Paleolithic "demic diffusion" from the East.^[12] 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.^[19]

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%).^[20]

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.^[21]^[22]^[23]^[24]^[25] The highest frequencies of this clade found so far in Europe were observed in the Pasiegos from Cantabria, ranging from 18% (8/45)^[25] 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%).^[26]

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),^[27] excluding recent immigration as only men with French surname were analysed in Sicily (approximately 2% overall, but up to 7% in Piazza Armerina),^[28] and in slightly lower frequencies in continental Italy (especially near Lucera)^[24] 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.^[29]

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, (8 out of 132),^[30]</ref> 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 (Beleza et al., (2006)), quite similar to the frequency found in Rio de Janeiro (5.4%) among European contributors."^[31] and among Hispanic men from California and Hawaii 2.4%.^[32]

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	^[33]
Algeria	Mozabite Berbers	20	80	^[17]
Algeria	Oran	102	45.1	^[34]
Algeria	Algiers	35	42.9	^[12]
Algeria	Kabyles from Tizi Ouzou	19	47.4	^[12]
Brazil	Rio de Janeiro	112	5.4	^[31]
Burkina Faso	Tuaregs	38	77.8	^[16]
Canary Islands	Fuerteventura	75	13.3	^[26]
Canary Islands	Gran Canaria	78	11.5	^[26]
Canary Islands	Tenerife	178	10.7	^[26]
Canary Islands	Lanzarote	97	6.2	^[26]
Canary Islands	La Palma	85	5.9	^[26]
Canary Islands	Gomera	92	4.4	^[26]
Canary Islands	Hierro	47	2.1	^[26]
Cuba		132	6.1	^[30]
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	^[22]
France		85	3.5	^[17]
France	Auvergne	89	5.6	^[27]
France	Île-de-France	91	5.5	^[27]
France	Nord-Pas-de-Calais	68	4.4	^[27]
France	Provence-Alpes-Côte d'Azur	45	2.2	^[27]
France	Midi-Pyrénées	67	1.5	^[27]
Iberia	Spain, Portugal	655	5.2	^[26]
Iberia	Spain, Portugal	1140	4.3	Adams et al. (2008)
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	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)^[29]
Jordania		101	4	^[22]
Lebanon		104	1.9	^[22]
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	^[16]
Morocco	Marrakesh Berbers	29	72.4	^[17]
Morocco	Southern Moroccan Berbers	187	98.5	Ahmed Reguig et al. (2014^[35]
Morocco	Moyen Atlas Berbers	69	71	^[17]
Morocco	Moroccan Arabs	54	31.5	^[17]
Morocco	Marrakesh (Amizmiz Valley)	33	84.8	Alvarez et al. (2009)
Morocco	Northern Moroccans (Beni Snassen)	67	79.1	^[33]
Morocco	Northern Moroccans (Rhiraya)	54	79.6	^[33]
Morocco	Immigrants resident in Italy	51	54.9	Onofri et al. (2008)
Morocco	Arabs and Berbers	221	65	^[26], from Bosh et al. 2001
Niger	Tuaregs	22	9.1	^[17]
Niger	Tuaregs	31	11.1	^[16]
North Africa	Sahara	89	59.6	^[26]
North Africa	Algeria, Tunisia	202	39.1	^[26]
Portugal	North	109	5.5	Flores et al. (2004)
Portugal	South	49	12.2	^[17]
Portugal	North	50	4	^[17]
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	^[16]
Portugal	Pias	46	4.3	^[16]
Portugal	Alcacer do Sal	21	4.8	^[16]
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	^[22]
Spain	Pasiegos from Cantabria	19	36.8	Scozzari et al. (2001)
Spain	Pasiegos from Cantabria	56	41.1	^[17]
Spain	Pasiegos from Cantabria	45	17.8	Maca-Meyer et al. (2003)
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	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	^[12]
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	^[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.^[36]^[37]^[38]^[39]

E-M123

E-M123 is mostly known for its major subclade E-M34, which dominates this clade.^{[Note 2]}

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.^[2]

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.^[40] 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.^[41] 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.^[42]

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.

α Jobling and Tyler-Smith 2000 and Kaladjieva 2001
β Underhill et al. 2000
γ Hammer 2001
δ Karafet et al. 2008
ε Semino et al. 2000
ζ Su 1999
η Capelli et al. 2001

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.
^ 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.

Ftdna E-PF2431 Project

References

^ https://www.yfull.com/tree/E-Z827/
^ ^a ^b 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.
^ 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-PF2431 YTree".
^ "E-M81 YTree". www.yfull.com. Retrieved 2016-07-09.
^ "E-M81 YTree". www.yfull.com. Retrieved 2016-07-10.
^ Solé-Morata N, García-Fernández 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. doi:10.1038/s41598-017-16271-y. PMID 29162904.
^ ^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.
^ Alvarez et al. (2009)
^ Bosch et al. (2006)
^ ^a ^b Kujanová M, Pereira L, Fernandes V, Pereira JB, Cerný 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.
^ ^a ^b ^c ^d ^e ^f ^g Pereira L, Cerný V, Cerezo M, Silva NM, Hájek M, Vasíková A, Kujanová 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 : EJHG. 18 (8): 915–23. doi:10.1038/ejhg.2010.21. PMC 2987384. PMID 20234393.
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o ^p ^q ^r ^s ^t ^u ^v ^w ^x 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.
^ Robino (2008)
^ Keita (2008). "In Hot Pursuit of Language". {{cite journal}}: |chapter= ignored (help); Cite journal requires |journal= (help); External link in |chapterurl= (help); Unknown parameter |chapterurl= ignored (|chapter-url= suggested) (help)
^ Ordóñez AC, Fregel R, Trujillo-Mederos A, Hervella M, de-la-Rúa 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.
^ Adams et al. (2008)
^ ^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.
^ Beleza et al. (2006)
^ ^a ^b Capelli et al. (2009)
^ ^a ^b Maca-Meyer N, Sánchez-Velasco P, Flores C, Larruga JM, González AM, Oterino A, Leyva-Cobián 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. doi:10.1046/j.1469-1809.2003.00045.x. PMID 12914567.{{cite journal}}: CS1 maint: postscript (link)
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l Fregel R, Gomes V, Gusmão L, González 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.{{cite journal}}: CS1 maint: unflagged free DOI (link)
^ ^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.
^ Di Gaetano et al. (2009)
^ ^a ^b Francalacci et al. (2013), Low-Pass DNA Sequencing of 1200 Sardinians Reconstructs European Y-Chromosome Phylogeny
^ ^a ^b Mendizabal I, Sandoval K, Berniell-Lee G, Calafell F, Salas A, Martínez-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.{{cite journal}}: CS1 maint: unflagged free DOI (link)
^ ^a ^b Silva DA, Carvalho E, Costa G, Tavares L, Amorim A, Gusmão L (2006). "Y-chromosome genetic variation in Rio de Janeiro population". American Journal of Human Biology : the Official Journal of the Human Biology Council. 18 (6): 829–37. doi:10.1002/ajhb.20567. PMID 17039481.
^ (7 out of 295), Paracchini et al. (2003)
^ ^a ^b ^c Dugoujon JM, Philippson G (2005). "The Berbers: Linguistic and genetic diversity" (PDF).
^ 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.
^ 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.
^ "E-M35 Project Data". haplozone.net.
^ "E-M35 Project Data". haplozone.net.
^ "E-M35 Project Data". haplozone.net.
^ "E-M35 Project Data". haplozone.net.
^ Henn et al. (2008)
^ 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.
^ "E-M35 Project Data". haplozone.net.

External links

[vanOven2014-45] Van Oven M, Van Geystelen A, Kayser M, Decorte R, Larmuseau HD (2014). "Seeing the wood for the trees: a minimal reference phylogeny for the human Y chromosome". Human Mutation. 35 (2): 187–91. doi:10.1002/humu.22468. PMID 24166809. S2CID 23291764.

[ISOGG2015-46] International Society of Genetic Genealogy (ISOGG; 2015), Y-DNA Haplogroup Tree 2015. (Access date: 1 February 2015.)

[A0-T-47] Haplogroup A0-T is also known as A-L1085 (and previously as A0'1'2'3'4).

[A1-48] Haplogroup A1 is also known as A1'2'3'4.

[LT-49] Haplogroup LT (L298/P326) is also known as Haplogroup K1.

[K2-50] Between 2002 and 2008, Haplogroup T-M184 was known as "Haplogroup K2". That name has since been re-assigned to K-M526, the sibling of Haplogroup LT.

[K2b-51] Haplogroup K2b (M1221/P331/PF5911) is also known as Haplogroup MPS.

[K-M2313-52] K-M2313*, which as yet has no phylogenetic name, has been documented in two living individuals, who have ethnic ties to India and South East Asia. In addition, K-Y28299, which appears to be a primary branch of K-M2313, has been found in three living individuals from India. See: Poznik op. cit.; YFull YTree v5.08, 2017, "K-M2335", and; PhyloTree, 2017, "Details of the Y-SNP markers included in the minimal Y tree" (Access date of these pages: 9 December 2017)

[K2b1-53] Haplogroup K2b1 (P397/P399) is also known as Haplogroup MS, but has a broader and more complex internal structure.

[P-54] Haplogroup P (P295) is also klnown as K2b2.

[S-55] Haplogroup S, as of 2017, is also known as K2b1a. (Previously the name Haplogroup S was assigned to K2b1a4.)

[M-56] Haplogroup M, as of 2017, is also known as K2b1b. (Previously the name Haplogroup M was assigned to K2b1d.)

[21] 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.

[41] 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.

[1] ttps://www.yfull.com/tree/E-Z827/

[Trombetta_2015-2] 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

[isogg-4] ISOGG (2008)

[Karafet2008-5] Karafet et al. (2008)

[YCC-6] Y Chromosome Consortium "YCC" (2002)

[ISOGG_2015-7] ISOGG 2015

[8] "E-PF2431 YTree".

[9] "E-M81 YTree". www.yfull.com. Retrieved 2016-07-09.

[10] "E-M81 YTree". www.yfull.com. Retrieved 2016-07-10.

[11] Solé-Morata N, García-Fernández 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. doi:10.1038/s41598-017-16271-y. PMID 29162904.

[Arredi_2004-12] ^ ^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.

[:0-13] Alvarez et al. (2009)

[14] Bosch et al. (2006)

[Kujanová_2009-15] Kujanová M, Pereira L, Fernandes V, Pereira JB, Cerný 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.

[Pereira_2010-16] ^ ^a ^b ^c ^d ^e ^f ^g Pereira L, Cerný V, Cerezo M, Silva NM, Hájek M, Vasíková A, Kujanová 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 : EJHG. 18 (8): 915–23. doi:10.1038/ejhg.2010.21. PMC 2987384. PMID 20234393.

[Cruciani_2004-17] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o ^p ^q ^r ^s ^t ^u ^v ^w ^x 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] Robino (2008)

[19] Keita (2008). "In Hot Pursuit of Language". {{cite journal}}: |chapter= ignored (help); Cite journal requires |journal= (help); External link in |chapterurl= (help); Unknown parameter |chapterurl= ignored (|chapter-url= suggested) (help)

[20] Ordóñez AC, Fregel R, Trujillo-Mederos A, Hervella M, de-la-Rúa 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.

[Adams2008-22] Adams et al. (2008)

[Flores_2005-23] 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] Beleza et al. (2006)

[Harvcoltxt|Capelli_et_al.|2009-25] Capelli et al. (2009)

[MacaMeyer2003-26] Maca-Meyer N, Sánchez-Velasco P, Flores C, Larruga JM, González AM, Oterino A, Leyva-Cobián 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. doi:10.1046/j.1469-1809.2003.00045.x. PMID 12914567.{{cite journal}}: CS1 maint: postscript (link)

[Fregel_2009-27] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l Fregel R, Gomes V, Gusmão L, González 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.{{cite journal}}: CS1 maint: unflagged free DOI (link)

[Ramos-Luis_2009-28] ^ ^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] Di Gaetano et al. (2009)

[sciencemag.org-30] Francalacci et al. (2013), Low-Pass DNA Sequencing of 1200 Sardinians Reconstructs European Y-Chromosome Phylogeny

[Mendizabal_2008-31] Mendizabal I, Sandoval K, Berniell-Lee G, Calafell F, Salas A, Martínez-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.{{cite journal}}: CS1 maint: unflagged free DOI (link)

[Silva_2006-32] Silva DA, Carvalho E, Costa G, Tavares L, Amorim A, Gusmão L (2006). "Y-chromosome genetic variation in Rio de Janeiro population". American Journal of Human Biology : the Official Journal of the Human Biology Council. 18 (6): 829–37. doi:10.1002/ajhb.20567. PMID 17039481.

[33] (7 out of 295), Paracchini et al. (2003)

[Dugoujon_2005-34] Dugoujon JM, Philippson G (2005). "The Berbers: Linguistic and genetic diversity" (PDF).

[Robino_2008-35] 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.

[Reguig_2014-36] 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.

[37] "E-M35 Project Data". haplozone.net.

[38] "E-M35 Project Data". haplozone.net.

[39] "E-M35 Project Data". haplozone.net.

[40] "E-M35 Project Data". haplozone.net.

[Henn2008-42] Henn et al. (2008)

[43] 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.

[44] "E-M35 Project Data". haplozone.net.

[1]

[2]

[3]

[4]

[5]

[6]

[7]

[8]

[9]

[10]

[11]

[12]

[13]

[14]

[15]

[16]

[17]

[18]

[19]

[20]

[Note 1]

[21]

[22]

[23]

[24]

[25]

[26]

[27]

[28]

[29]

[30]

[31]

[32]

[33]

[34]

[35]

[36]

[37]

[38]

[39]

[Note 2]

[40]

[41]

[42]

[χ 1]

[χ 2]

[χ 3]

[χ 4]

[χ 5]

[χ 6]

[χ 7]

[χ 8]

[χ 9]

[χ 10]

[χ 11]

[χ 12]

@@ Line 96: / Line 96: @@
 '''E-M81''' is the most common subclade of haplogroup E-L19/V257. It is concentrated in the [[Tamazgha]], 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.<ref>{{cite journal | vauthors = Solé-Morata N, García-Fernández C, Urasin V, Bekada A, Fadhlaoui-Zid K, Zalloua P, Comas D, Calafell F | title = Whole Y-chromosome sequences reveal an extremely recent origin of the most common North African paternal lineage E-M183 (M81) | journal = Scientific Reports | volume = 7 | issue = 1 | pages = 15941 | date = November 2017 | pmid = 29162904 | doi = 10.1038/s41598-017-16271-y | url = http://www.nature.com/articles/s41598-017-16271-y }}</ref>
-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.<ref name="Arredi_2004" /><ref name=":0">{{Harvcoltxt|Alvarez et al.|2009}}</ref><ref>{{Harvcoltxt|Bosch et al.|2006}}</ref><ref>{{cite journal | vauthors = Kujanová M, Pereira L, Fernandes V, Pereira JB, Cerný V | title = Near eastern neolithic genetic input in a small oasis of the Egyptian Western Desert | journal = American Journal of Physical Anthropology | volume = 140 | issue = 2 | pages = 336–46 | date = October 2009 | pmid = 19425100 | doi = 10.1002/ajpa.21078 }}</ref> Because of its prevalence among these groups and also others such as [[Mozabite people|Mozabite]], [[Middle Atlas]], [[Kabyle people|Kabyle]] and other [[Berber people|Berber]] groups, it is sometimes referred to as a genetic "[[Berber people|Berber]] marker". {{Harvcoltxt|Pereira et al.|2010}} report high levels among two [[Tuareg people|Tuareg]] populations inhabiting the [[Sahara]]: 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]].
+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.<ref name="Arredi_2004" /><ref name=":0">{{Harvcoltxt|Alvarez et al.|2009}}</ref><ref>{{Harvcoltxt|Bosch et al.|2006}}</ref><ref name = "Kujanová_2009">{{cite journal | vauthors = Kujanová M, Pereira L, Fernandes V, Pereira JB, Cerný V | title = Near eastern neolithic genetic input in a small oasis of the Egyptian Western Desert | journal = American Journal of Physical Anthropology | volume = 140 | issue = 2 | pages = 336–46 | date = October 2009 | pmid = 19425100 | doi = 10.1002/ajpa.21078 }}</ref> Because of its prevalence among these groups and also others such as [[Mozabite people|Mozabite]], [[Middle Atlas]], [[Kabyle people|Kabyle]] and other [[Berber people|Berber]] groups, it is sometimes referred to as a genetic "[[Berber people|Berber]] marker". High levels among two [[Tuareg people|Tuareg]] populations inhabiting the [[Sahara]]: 77.8% near [[Gorom-Gorom]], in [[Burkina Faso]], and 81.8% from [[Gossi]] in [[Mali]] are observed.<ref name="Pereira_2010">{{cite journal | vauthors = Pereira L, Cerný V, Cerezo M, Silva NM, Hájek M, Vasíková A, Kujanová M, Brdicka R, Salas A | title = Linking the sub-Saharan and West Eurasian gene pools: maternal and paternal heritage of the Tuareg nomads from the African Sahel | journal = European Journal of Human Genetics : EJHG | volume = 18 | issue = 8 | pages = 915–23 | date = August 2010 | pmid = 20234393 | pmc = 2987384 | doi = 10.1038/ejhg.2010.21 }}</ref> 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 [[Maghrebi Arabic|Arabic]]-speaking groups. It reaches 75% among Moroccans Arabs,<ref>{{cite journal | vauthors = 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 | title = Phylogeographic analysis of haplogroup E3b (E-M215) y chromosomes reveals multiple migratory events within and out of Africa | journal = American Journal of Human Genetics | volume = 74 | issue = 5 | pages = 1014–22 | date = May 2004 | pmid = 15042509 | pmc = 1181964 | doi = 10.1086/386294 }}</ref>  and is generally found at frequencies around 45% in coastal cities of the Maghreb ([[Oran]], [[Tunis]], [[Tizi Ouzou]], [[Algiers]]).<ref name="Arredi_2004" /><ref>Robino (2008)</ref>
+The E-M81 subclade is also quite common among North African [[Maghrebi Arabic|Arabic]]-speaking groups. It reaches 75% among Moroccans Arabs,<ref name = "Cruciani_2004">{{cite journal | vauthors = 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 | title = Phylogeographic analysis of haplogroup E3b (E-M215) y chromosomes reveals multiple migratory events within and out of Africa | journal = American Journal of Human Genetics | volume = 74 | issue = 5 | pages = 1014–22 | date = May 2004 | pmid = 15042509 | pmc = 1181964 | doi = 10.1086/386294 }}</ref>  and is generally found at frequencies around 45% in coastal cities of the Maghreb ([[Oran]], [[Tunis]], [[Tizi Ouzou]], [[Algiers]]).<ref name="Arredi_2004" /><ref>Robino (2008)</ref>
 In this key area from Egypt to the [[Atlantic Ocean]],<ref name="Arredi_2004" /> report a pattern of decreasing [[Short tandem repeat|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, {{Harvcoltxt|Kujanova et al.|2009}} found M81 in 28.6% (10 out of 35 men) in [[El-Hayez]] in the [[Libyan Desert|Western desert]] in Egypt
@@ Line 107: / Line 107: @@
 =====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<ref group="Note">{{Harvcoltxt|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 (Spain)|Galicia]], 10% in Western [[Andalusia]] and Northwest [[Castile (historical region)|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) , [http://www.cell.com/AJHG/image/S0002-9297(08)00592-2?imageId=gr1&imageType=large see table].</ref> 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 (Spain)|Galicia]], 10% in Western [[Andalusia]] and Northwest [[Castile (historical region)|Castile]] and 9% to 17% in [[Cantabria]].<ref name = "Adams2008">{{Harvcoltxt|Adams et al.|2008}}</ref><ref>{{Harvcoltxt|Flores et al.|2005}}</ref><ref>{{Harvcoltxt|Beleza et al.|2006}}</ref><ref name="Harvcoltxt|Capelli et al.|2009">{{Harvcoltxt|Capelli et al.|2009}}</ref><ref name = "MacaMeyer2003" /> The highest frequencies of this clade found so far in Europe were observed in the [[Valles Pasiegos|Pasiegos]] from [[Cantabria]], ranging from 18% (8/45)<ref name = "MacaMeyer2003">{{cite journal | vauthors = Maca-Meyer N, Sánchez-Velasco P, Flores C, Larruga JM, González AM, Oterino A, Leyva-Cobián F | title = Y chromosome and mitochondrial DNA characterization of Pasiegos, a human isolate from Cantabria (Spain) | journal = Annals of Human Genetics | volume = 67 | issue = Pt 4 | pages = 329–39 | date = July 2003 | pmid = 12914567 | doi = 10.1046/j.1469-1809.2003.00045.x | postscript = . }}</ref> to 41% (23/56).<ref>{{Harvcoltxt|Cruciani et al.|2004}}</ref> 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%).<ref>Fregel et al. (2009), [https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2728732/?tool=pubmed Demographic history of Canary Islands male gene-pool: replacement of native lineages by European], [https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2728732/table/T1/ see table]</ref>
+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<ref group="Note">{{Harvcoltxt|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 (Spain)|Galicia]], 10% in Western [[Andalusia]] and Northwest [[Castile (historical region)|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) , [http://www.cell.com/AJHG/image/S0002-9297(08)00592-2?imageId=gr1&imageType=large see table].</ref> 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 (Spain)|Galicia]], 10% in Western [[Andalusia]] and Northwest [[Castile (historical region)|Castile]] and 9% to 17% in [[Cantabria]].<ref name = "Adams2008">{{Harvcoltxt|Adams et al.|2008}}</ref><ref name="Flores_2005">{{cite journal | vauthors = Flores C, Maca-Meyer N, Larruga JM, Cabrera VM, Karadsheh N, Gonzalez AM | title = Isolates in a corridor of migrations: a high-resolution analysis of Y-chromosome variation in Jordan | journal = Journal of Human Genetics | volume = 50 | issue = 9 | pages = 435–41 | date = 2005 | pmid = 16142507 | doi = 10.1007/s10038-005-0274-4 }}</ref><ref>{{Harvcoltxt|Beleza et al.|2006}}</ref><ref name="Harvcoltxt|Capelli et al.|2009">{{Harvcoltxt|Capelli et al.|2009}}</ref><ref name = "MacaMeyer2003" /> The highest frequencies of this clade found so far in Europe were observed in the [[Valles Pasiegos|Pasiegos]] from [[Cantabria]], ranging from 18% (8/45)<ref name = "MacaMeyer2003">{{cite journal | vauthors = Maca-Meyer N, Sánchez-Velasco P, Flores C, Larruga JM, González AM, Oterino A, Leyva-Cobián F | title = Y chromosome and mitochondrial DNA characterization of Pasiegos, a human isolate from Cantabria (Spain) | journal = Annals of Human Genetics | volume = 67 | issue = Pt 4 | pages = 329–39 | date = July 2003 | pmid = 12914567 | doi = 10.1046/j.1469-1809.2003.00045.x | postscript = . }}</ref> to 41% (23/56).<ref name = "Cruciani_2004" /> 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%).<ref name="Fregel_2009">{{cite journal | vauthors = Fregel R, Gomes V, Gusmão L, González AM, Cabrera VM, Amorim A, Larruga JM | title = Demographic history of Canary Islands male gene-pool: replacement of native lineages by European | journal = BMC Evolutionary Biology | volume = 9 | issue = | pages = 181 | date = August 2009 | pmid = 19650893 | pmc = 2728732 | doi = 10.1186/1471-2148-9-181 | url = }}</ref>
+E-M81 is also found in [[France]], 2.70% (15/555) overall with frequencies surpassing 5% in [[Auvergne (region)|Auvergne]] (5/89) and [[Île-de-France (region)|Île-de-France]] (5/91),<ref name = "Ramos-Luis_2009">{{cite journal|vauthors = Ramos-Luis E, Blanco-Verea A, Brión M, Van Huffel V, Carracedo A, Sánchez-Diz P | title = Phylogeography of French male lineages | journal = Forensic Science International: Genetics Supplement Series | date = December 2009 | volume = 2 | issue = 1 |pages = 439–441 | doi = 10.1016/j.fsigss.2009.09.026 }}</ref> excluding recent immigration as only men with French surname were analysed in [[Sicily]] (approximately 2% overall, but up to 7% in [[Piazza Armerina]]),<ref>{{Harvcoltxt|Di Gaetano et al.|2009}}</ref> and in slightly lower frequencies in continental [[Italy]] (especially near [[Lucera]])<ref name="Harvcoltxt|Capelli et al.|2009"/>  due to historic colonization during the [[Islam in Europe|Islamic]], [[Roman Empire|Roman]], and [[Carthage|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 [[Sardinia]]ns.<ref name="sciencemag.org">Francalacci et al. (2013), [http://www.sciencemag.org/content/341/6145/565.abstract Low-Pass DNA Sequencing of 1200 Sardinians Reconstructs European Y-Chromosome Phylogeny]</ref>
-E-M81 is also found in [[France]], 2.70% (15/555) overall with frequencies surpassing 5% in [[Auvergne (region)|Auvergne]] (5/89) and [[Île-de-France (region)|Île-de-France]] (5/91),<ref>{{Harvcoltxt|Ramos-Luisa et al.|2009}}</ref><ref>excluding recent immigration as only men with French surname were analysed</ref> in [[Sicily]] (approximately 2% overall, but up to 7% in [[Piazza Armerina]]),<ref>{{Harvcoltxt|Di Gaetano et al.|2009}}</ref> and in slightly lower frequencies in continental [[Italy]] (especially near [[Lucera]])<ref name="Harvcoltxt|Capelli et al.|2009"/>  due to historic colonization during the [[Islam in Europe|Islamic]], [[Roman Empire|Roman]], and [[Carthage|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 [[Sardinia]]ns.<ref name="sciencemag.org">Francalacci et al. (2013), [http://www.sciencemag.org/content/341/6145/565.abstract Low-Pass DNA Sequencing of 1200 Sardinians Reconstructs European Y-Chromosome Phylogeny]</ref>
 =====Latin America=====
-As a result of [[Spanish colonization of the Americas|Spanish]] and [[Portuguese colonization of the Americas|Portuguese]] colonization of the Americas, this sub-clade is found throughout [[Latin America]], for example 6.1% in [[Cuba]],<ref>(8 out of 132), {{Harvcoltxt|Mendizabal et al.|2008}}</ref> 5.4% in [[Brazil]] (Rio de Janeiro), <ref group="Note">(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]] ({{Harvcoltxt|Beleza et al.,|2006}}), quite similar to the frequency found in [[Rio de Janeiro]] (5.4%) among European contributors." {{Harvcoltxt|Silva et al.|2006}}</ref> and among [[Hispanic]] men from [[California]] and [[Hawaii]] 2.4%.<ref>(7 out of 295), {{Harvcoltxt|Paracchini et al.|2003}}</ref>
+As a result of [[Spanish colonization of the Americas|Spanish]] and [[Portuguese colonization of the Americas|Portuguese]] colonization of the Americas, this sub-clade is found throughout [[Latin America]], for example 6.1% in [[Cuba]], (8 out of 132),<ref name="Mendizabal_2008">{{cite journal | vauthors = Mendizabal I, Sandoval K, Berniell-Lee G, Calafell F, Salas A, Martínez-Fuentes A, Comas D | title = Genetic origin, admixture, and asymmetry in maternal and paternal human lineages in Cuba | journal = BMC Evolutionary Biology | volume = 8 | issue = | pages = 213 | date = July 2008 | pmid = 18644108 | pmc = 2492877 | doi = 10.1186/1471-2148-8-213 }}</ref></ref> 5.4% in [[Brazil]] (Rio de Janeiro) (6 out of 112), "The presence of chromosomes of North African origin (E3b1b-M81;<ref name = "Cruciani_2004" /> can also be explained by a Portuguese-mediated influx, since this haplogroup reaches a frequency of 5.6% in [[Portugal]] ({{Harvcoltxt|Beleza et al.,|2006}}), quite similar to the frequency found in [[Rio de Janeiro]] (5.4%) among European contributors."<ref name="Silva_2006">{{cite journal | vauthors = Silva DA, Carvalho E, Costa G, Tavares L, Amorim A, Gusmão L | title = Y-chromosome genetic variation in Rio de Janeiro population | journal = American Journal of Human Biology : the Official Journal of the Human Biology Council | volume = 18 | issue = 6 | pages = 829–37 | date = 2006 | pmid = 17039481 | doi = 10.1002/ajhb.20567 | url = }}</ref> and among [[Hispanic]] men from [[California]] and [[Hawaii]] 2.4%.<ref>(7 out of 295), {{Harvcoltxt|Paracchini et al.|2003}}</ref>
 =====Others=====
@@ Line 124: / Line 125: @@
 !style="background:#f0f0f0;" align="center" | Country/Region!!Sampling!!n!!%E-M81!!Source
 |-
+| Algeria||Mozabite Berbers||67||86.6||<ref name = "Dugoujon_2005">{{cite web | url = http://web.archive.org/web/20120325132014/http://www.ddl.ish-lyon.cnrs.fr/Fulltext/philippson/AUSSOIS_2005_final.pdf | title = The Berbers: Linguistic and genetic diversity | vauthors = Dugoujon JM, Philippson G | date = 2005 }}</ref>
-| Algeria||Mozabite Berbers||67||86.6||Dugoujon et al. (2009)
 |-
-| Algeria||Mozabite Berbers||20||80||Cruciani et al. (2004)
+| Algeria||Mozabite Berbers||20||80||<ref name = "Cruciani_2004" />
 |-
+| Algeria||Oran||102||45.1||<ref name="Robino_2008">{{cite journal | vauthors = Robino C, Crobu F, Di Gaetano C, Bekada A, Benhamamouch S, Cerutti N, Piazza A, Inturri S, Torre C | title = Analysis of Y-chromosomal SNP haplogroups and STR haplotypes in an Algerian population sample | journal = International Journal of Legal Medicine | volume = 122 | issue = 3 | pages = 251–5 | date = May 2008 | pmid = 17909833 | doi = 10.1007/s00414-007-0203-5 }}</ref>
-| Algeria||Oran||102||45.1||Robino et al. (2008)
 |-
 | Algeria||Algiers||35||42.9||<ref name="Arredi_2004">{{cite journal | vauthors = Arredi B, Poloni ES, Paracchini S, Zerjal T, Fathallah DM, Makrelouf M, Pascali VL, Novelletto A, Tyler-Smith C | title = A predominantly neolithic origin for Y-chromosomal DNA variation in North Africa | journal = American Journal of Human Genetics | volume = 75 | issue = 2 | pages = 338–45 | date = August 2004 | pmid = 15202071 | pmc = 1216069 | doi = 10.1086/423147 }}</ref>
@@ Line 134: / Line 135: @@
 | Algeria||Kabyles from Tizi Ouzou||19||47.4||<ref name="Arredi_2004" />
 |-
-| Brazil||Rio de Janeiro||112||5.4||Silva et al. (2006)
+| Brazil||Rio de Janeiro||112||5.4||<ref name="Silva_2006" />
 |-
-| Burkina Faso||Tuaregs ||38||77.8||Pereira et al. (2010)
+| Burkina Faso||Tuaregs ||38||77.8||<ref name="Pereira_2010" />
 |-
-| Canary Islands||Fuerteventura||75||13.3||Fregel et al. (2009)
+| Canary Islands||Fuerteventura||75||13.3||<ref name="Fregel_2009" />
 |-
-| Canary Islands||Gran Canaria||78||11.5||Fregel et al. (2009)
+| Canary Islands||Gran Canaria||78||11.5||<ref name="Fregel_2009" />
 |-
-| Canary Islands||Tenerife||178||10.7||Fregel et al. (2009)
+| Canary Islands||Tenerife||178||10.7||<ref name="Fregel_2009" />
 |-
-| Canary Islands||Lanzarote||97||6.2||Fregel et al. (2009)
+| Canary Islands||Lanzarote||97||6.2||<ref name="Fregel_2009" />
 |-
-| Canary Islands||La Palma||85||5.9||Fregel et al. (2009)
+| Canary Islands||La Palma||85||5.9||<ref name="Fregel_2009" />
 |-
-| Canary Islands||Gomera||92||4.4||Fregel et al. (2009)
+| Canary Islands||Gomera||92||4.4||<ref name="Fregel_2009" />
 |-
-| Canary Islands||Hierro||47||2.1||Fregel et al. (2009)
+| Canary Islands||Hierro||47||2.1||<ref name="Fregel_2009" />
 |-
-| Cuba||||132||6.1||Mendizabal et al. (2008)
+| Cuba||||132||6.1||<ref name="Mendizabal_2008" />
 |-
-| Cyprus||Turkish Cypriots||46||8.7||Cruciani et al. (2004)
+| Cyprus||Turkish Cypriots||46||8.7||<ref name = "Cruciani_2004" />
 |-
-| Egypt||Northern Egyptians||21||4.8||Cruciani et al. (2004)
+| Egypt||Northern Egyptians||21||4.8||<ref name = "Cruciani_2004" />
 |-
-| Egypt||Western Desert ||35||28.6||Kujanová et al. (2009)
+| Egypt||Western Desert ||35||28.6||<ref name = "Kujanová_2009" />
 |-
-| Egypt||||147||8.2||Flores et al. (2005)
+| Egypt||||147||8.2||<ref name="Flores_2005" />
 |-
-| France||||85||3.5||Cruciani et al. (2004)
+| France||||85||3.5||<ref name = "Cruciani_2004" />
 |-
-| France||Auvergne||89||5.6|| Ramos-Luisa et al. (2009)
+| France||Auvergne||89||5.6||<ref name = "Ramos-Luis_2009" />
 |-
-| France||Île-de-France||91||5.5|| Ramos-Luisa et al. (2009)
+| France||Île-de-France||91||5.5||<ref name = "Ramos-Luis_2009" />
 |-
-| France||Nord-Pas-de-Calais||68||4.4|| Ramos-Luisa et al. (2009)
+| France||Nord-Pas-de-Calais||68||4.4||<ref name = "Ramos-Luis_2009" />
 |-
-| France|| Provence-Alpes-Côte d'Azur||45||2.2|| Ramos-Luisa et al. (2009)
+| France|| Provence-Alpes-Côte d'Azur||45||2.2||<ref name = "Ramos-Luis_2009" />
 |-
-| France|| Midi-Pyrénées||67||1.5|| Ramos-Luisa et al. (2009)
+| France|| Midi-Pyrénées||67||1.5||<ref name = "Ramos-Luis_2009" />
 |-
-| Iberia||Spain, Portugal||655||5.2||Fregel et al. (2009)
+| Iberia||Spain, Portugal||655||5.2||<ref name="Fregel_2009" />
 |-
 | Iberia||Spain, Portugal||1140||4.3||Adams et al. (2008)
 |-
-| Israel||Bedouins||28||3.6||Cruciani et al. (2004)
+| Israel||Bedouins||28||3.6||<ref name = "Cruciani_2004" />
 |-
-| Italy||Central Italians||89||2.2||Cruciani et al. (2004)
+| Italy||Central Italians||89||2.2||<ref name = "Cruciani_2004" />
 |-
-| Italy||Northern Italians||67||1.5||Cruciani et al. (2004)
+| Italy||Northern Italians||67||1.5||<ref name = "Cruciani_2004" />
 |-
 | Italy||North-West Apulia||46||4.3||Capelli et al. (2009)
@@ Line 198: / Line 199: @@
 | Italy||Sardinia||1204||5.8||Francalacci et al. (2013)<ref name="sciencemag.org"/>
 |-
-| Jordania||||101||4||Flores et al. (2005)
+| Jordania||||101||4||<ref name="Flores_2005" />
 |-
-| Lebanon||||104||1.9||Flores et al. (2005)
+| Lebanon||||104||1.9||<ref name="Flores_2005" />
 |-
 | Lebanon||||914||1.2||Zalloua et al. (2008)
@@ Line 208: / Line 209: @@
 | Libya||Arabs||215||35.9||Fadhlaoui-Zid et al. (2013)
 |-
-| Mali||Tuaregs (Gozi)||21||81.8||Pereira et al. (2010)
+| Mali||Tuaregs (Gozi)||21||81.8||<ref name="Pereira_2010" />
 |-
-| Morocco||Marrakesh Berbers||29||72.4||Cruciani et al. (2004)
+| Morocco||Marrakesh Berbers||29||72.4||<ref name = "Cruciani_2004" />
 |-
 |Morocco
@@ Line 218: / Line 219: @@
 |Ahmed Reguig et al. (2014<ref name="Reguig_2014">{{cite journal | vauthors = Reguig A, Harich N, Barakat A, Rouba H | title = Phylogeography of E1b1b1b-M81 haplogroup and analysis of its subclades in Morocco | journal = Human Biology | volume = 86 | issue = 2 | pages = 105–12 | date = 2014 | pmid = 25397701 | doi = 10.3378/027.086.0204 }}</ref>
 |-
-| Morocco||Moyen Atlas Berbers||69||71||Cruciani et al. (2004)
+| Morocco||Moyen Atlas Berbers||69||71||<ref name = "Cruciani_2004" />
 |-
-| Morocco||Moroccan Arabs||54||31.5||Cruciani et al. (2004)
+| Morocco||Moroccan Arabs||54||31.5||<ref name = "Cruciani_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 (Beni  Snassen) ||67||79.1||<ref name = "Dugoujon_2005" />
 |-
-| Morocco||Northern Moroccans (Rhiraya) ||54||79.6||Dugoujon et al. (2005)<ref>{{cite web|url=http://www.ddl.ish-lyon.cnrs.fr/Fulltext/philippson/AUSSOIS_2005_final.pdf|format=PDF|title=The Berbers Linguistic and enetic diversity|website=ddl.ish-lyon.cnrs.fr}}</ref>
+| Morocco||Northern Moroccans (Rhiraya) ||54||79.6||<ref name = "Dugoujon_2005" />
 |-
 | 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
+| Morocco|| Arabs and Berbers||221||65||<ref name="Fregel_2009" />,  from Bosh et al. 2001
 |-
-| Niger||Tuaregs||22||9.1||Cruciani et al. (2004)
+| Niger||Tuaregs||22||9.1||<ref name = "Cruciani_2004" />
 |-
-| Niger||Tuaregs||31||11.1||Pereira et al. (2010)
+| Niger||Tuaregs||31||11.1||<ref name="Pereira_2010" />
 |-
-| North Africa||Sahara||89||59.6||Fregel et al. (2009)
+| North Africa||Sahara||89||59.6||<ref name="Fregel_2009" />
 |-
-| North Africa||Algeria, Tunisia||202||39.1||Fregel et al. (2009)
+| North Africa||Algeria, Tunisia||202||39.1||<ref name="Fregel_2009" />
 |-
 | Portugal||North ||109||5.5||Flores et al. (2004)
 |-
-| Portugal||South||49||12.2||Cruciani et al. (2004)
+| Portugal||South||49||12.2||<ref name = "Cruciani_2004" />
 |-
-| Portugal||North||50||4||Cruciani et al. (2004)
+| Portugal||North||50||4||<ref name = "Cruciani_2004" />
 |-
 | Portugal||South||78||7.7||Adams et al. (2008)
@@ Line 278: / Line 279: @@
 | Portugal||Alentejo||65||7.7||Beleza  et al. (2006)
 |-
-| Portugal||Coruche||64||9.4||Pereira et al. (2010)
+| Portugal||Coruche||64||9.4||<ref name="Pereira_2010" />
 |-
-| Portugal||Pias||46||4.3||Pereira et al. (2010)
+| Portugal||Pias||46||4.3||<ref name="Pereira_2010" />
 |-
-| Portugal||Alcacer do Sal||21||4.8||Pereira et al. (2010)
+| Portugal||Alcacer do Sal||21||4.8||<ref name="Pereira_2010" />
 |-
 | Portugal||Tras-os-Montes (Jews)||57||5.3||Nogueiro et al. (2010)
@@ Line 288: / Line 289: @@
 | Portugal||Tras-os-Montes (Non Jews)||30||10||Nogueiro et al. (2010)
 |-
-| Somalia||||201||1.5||Flores et al. (2005)
+| Somalia||||201||1.5||<ref name="Flores_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||56||41.1||<ref name = "Cruciani_2004" />
 |-
 | Spain||Pasiegos from Cantabria||45||17.8||Maca-Meyer et al. (2003)
 |-
-| Spain||Spanish Basques||55||3.6||Cruciani et al. (2004)
+| Spain||Spanish Basques||55||3.6||<ref name = "Cruciani_2004" />
 |-
-| Spain||Asturians||90||2.2||Cruciani et al. (2004)
+| Spain||Asturians||90||2.2||<ref name = "Cruciani_2004" />
 |-
-| Spain||Southern Spaniards||62||1.6||Cruciani et al. (2004)
+| Spain||Southern Spaniards||62||1.6||<ref name = "Cruciani_2004" />
 |-
 | Spain||Castile, NorthWest||100||10||Adams et al. (2008)
@@ Line 392: / Line 393: @@
 | Tunisia||Cosmopolitan Tunis||33||54.4||Fadhlaoui-Zid et al. (2011)
 |-
-| Turkey||Istanbul Turkish||35||5.7||Cruciani et al. (2004)
+| Turkey||Istanbul Turkish||35||5.7||<ref name = "Cruciani_2004" />
 |-
-| Turkey||Sephardi Turkish||19||5.3||Cruciani et al. (2004)
+| Turkey||Sephardi Turkish||19||5.3||<ref name = "Cruciani_2004" />
 |-
-| Turkey||Southwestern Turkish||40||2.5||Cruciani et al. (2004)
+| Turkey||Southwestern Turkish||40||2.5||<ref name = "Cruciani_2004" />
 |-
-| Turkey||Northeastern Turkish||41||2.4||Cruciani et al. (2004)
+| Turkey||Northeastern Turkish||41||2.4||<ref name = "Cruciani_2004" />
 |-
 |}