Haplogroup E-M215 (Y-DNA)
|Possible time of origin||42,600|
|Possible place of origin||North Africa or Horn of Africa|
|Descendants||Haplogroups E-M35 and E-M281|
|Defining mutations||M215, most often found in conjunction with M35|
Human Y-chromosome DNA haplogroup E-M215, also referred to in the literature by other names such as E1b1b and E3b (see further discussion below), is a major Y chromosome haplogroup. It is a division of the macro haplogroup E-M96, which is defined by the single-nucleotide polymorphism (SNP) mutation M215. In other words, it is one of the major patrilineal lineages of humanity, linking from father-to-son back to a common male-line ancestor ("Y-chromosomal Adam"). It is a subject of discussion and study in genetics as well as genetic genealogy, archaeology, and historical linguistics.
E-M215 has two ancient branches that contain all known modern E-M215, E-M35 and E-M281 subclades. Of the latter two, the only branch that has been confirmed in a native population outside of Ethiopia is E-M35. E-M35 in turn has two known branches, haplogroup E-V68 and haplogroup E-Z827, which contain by far the majority of all modern E-M215 men. E-V68 and E-V257 have been found in highest numbers in North Africa and the Horn of Africa, but also in lower numbers in parts of the Middle East and Europe, and in isolated populations of Southern Africa.
- 1 Origins
- 2 Distribution
- 3 Subclades of E-M215
- 3.1 Family Tree
- 3.2 TMRCA of the major nodes in E-M215
- 3.3 E-V68 (E1b1b1a)
- 3.4 E-Z827 (E1b1b1b)
- 4 Phylogenetics
- 5 See also
- 6 Notes
- 7 References
- 8 Additional sources
- 9 External links
The origins of E-M215 were dated by Cruciani in 2007 to about 22,400 years ago in the Horn of Africa.[Note 1] E-M35 was dated by Batini in 2015 to between 15,400 and 20,500 years ago. In June 2015, Trombetta et al. reported a previously unappreciated large difference in the age between haplogroup E-M215 (38.6 kya; 95% CI 31.4-45.9 kya) and its sub-haplogroup E-M35 (25.0 kya; 95% CI 20.0-30.0 kya).
All major sub-branches of E-M35 are thought to have originated in the same general area as the parent clade: in North Africa, the Horn of Africa, or nearby areas of the Near East. Some branches of E-M35 are assumed to have left Africa thousands of years ago, whereas others may have arrived from the Near East. For example, Underhill (2002) associates the spread have the haplogroup with the Neolithic Revolution, believing that the structure and regional pattern of E-M35 subclades potentially give "reagents with which to infer specific episodes of population histories associated with the Neolithic agricultural expansion". Battaglia et al. (2007) also estimate that E-M78 (called E1b1b1a1 in that paper) has been in Europe longer than 10,000 years. Accordingly, human remains excavated in a Spanish funeral cave dating from approximately 7,000 years ago were shown to be in this haplogroup. Two more E-M78 have been found in the Neolitich Sopot and Lengyel cultures too.
Concerning European E-M35 within this scheme, Underhill & Kivisild (2007) have remarked that E-M215 seems to represent a late-Pleistocene migration from North Africa to Europe over the Sinai Peninsula in Egypt.[Note 2] While this proposal remains uncontested, it has more recently been proposed by Trombetta et al. (2011) that there is also evidence for additional migration of E-M215 carrying men directly from Africa to southwestern Europe, via a maritime route (see below.)
In Africa, E-M215 is distributed in highest frequencies in the Horn of Africa and North Africa, whence it has in recent millennia expanded as far south as South Africa, and northwards into Western Asia and Europe (especially the Mediterranean and the Balkans).
Almost all E-M215 men are also in E-M35. In 2004, M215 was found to be older than M35 when individuals were found who have the M215 mutation, but do not have M35 mutation. In 2013, one individual in Khorasan, North-East Iran, was found by Di Cristofaro et al. (2013) to be positive for M215 but negative for M35.
E-M215 and E-M35 are quite common among Afroasiatic speakers. The linguistic group and carriers of E-M35 lineage have a high probability to have arisen and dispersed together from the Afroasiatic Urheimat. Amongst populations with an Afro-Asiatic speaking history, a significant proportion of Jewish male lineages are E-M35. Haplogroup E-M35, which accounts for approximately 18% to 20% of Ashkenazi and 8.6% to 30% of Sephardi Y-chromosomes, appears to be one of the major founding lineages of the Jewish population.[Note 3]
The following table only includes sample populations with more than 1% E-M215 men with all known subclades as of June 2015 including the recent clade (E-V1515) which was defined by Trombetta et al. 2015 and includes all the sub-Saharan haplogroups (E-V42, E-M293, E-V92, E-V6) reported as E-M35 basal clades in a previous phylogeny.
|Moyen Atlas Berbers||69||Morocco||AA/Berber||81.2||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||10.1||0.0||71.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0|
|Egyptian Berbers from Siwa||93||Egypt||AA/Semitic||18.3||0.0||0.0||0.0||0.0||0.0||0.0||2.2||0.0||0.0||4.3||2.2||1.1||0.0||2.2||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||6.5||0.0|
|Egyptians from Baharia||41||Egypt||AA/Semitic||56.1||0.0||0.0||0.0||0.0||2.4||22.0||14.6||0.0||0.0||2.4||7.3||4.9||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||2.4||0.0|
|Egyptians from Gurna Oasis||34||Egypt||AA/Semitic||17.6||0.0||5.9||0.0||0.0||0.0||0.0||8.8||2.9||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0|
|Shuwa Arabs||5||Cameroon (North)||AA/Semitic||20.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||20.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0|
|Fulbe from Cameroon||76||Cameroon (North)||NC/Atlantic||1.3||1.3||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0|
|Nilotic Western Kenya||11||Kenya||NS/Sudanic||45.5||0.0||0.0||0.0||0.0||0.0||9.1||9.1||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||18.2||0.0||0.0||0.0||9.1||0.0|
|Pasiegos from Cantabria||56||Spain||IE||42.9||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||1.8||41.1||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0|
|Arabs||41||United Arab Emirate||AA/Semitic||7.3||0.0||0.0||0.0||0.0||0.0||2.4||0.0||0.0||0.0||0.0||0.0||0.0||0.0||4.9||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0||0.0|
Subclades of E-M215
The following phylogenetic tree is based on the YCC 2008 tree and subsequent published research as summarized by ISOGG. It includes all known subclades as of June 2015 (Trombetta et al. 2015)
- E-M215 (E1b1b)
- E-M215*. Rare or non-existent.
- E-M35 (E1b1b1)
- E-V68 (E1b1b1a)
- E-V2009. Found in individuals in Sardinia and Morocco.
- E-M78 (E1b1b1a1). North Africa, Horn of Africa, West Asia, Europe. (Formerly "E1b1b1a".)
- E-V1477. Found in Tunisian Jews.
- E-V1083*. Found only in Eritrea (1.1%) and Sardinia (0.3%).
- E-V259. Found in North Cameroon.
- E-Z827 (E1b1b1b)
- E-V257/L19 (L19, V257) - E1b1b1b1
- E-M81 (M81)
- E-Z830 (Z830) - E1b1b1b2
- E-M123 (M123)
- E-M34 (M34)
- E-M84 (M84)
- E-M136 (M136)
- E-M290 (M290)
- E-V23 (V23)
- E-L791 (L791,L792)
- E-M84 (M84)
- E-M34 (M34)
- E-V1515. E-V1515 and its subclades are mainly restricted to eastern Africa.
- E-M293 (M293)
- E-P72 (P72)
- E-V42 (V42)
- E-V6 (V6)
- E-M123 (M123)
- E-V257/L19 (L19, V257) - E1b1b1b1
- E-V16/E-M281 (E1b1b2). Rare. Found in individuals in Ethiopia, Yemen and Saudi Arabia.
- E-V68 (E1b1b1a)
Exceptional cases of men who are M215 positive but M35 negative ("E-M215*") have been discovered so far in two Amharas of Ethiopia and one Yemeni. At least some of these men, perhaps all, are known since early 2011 to be in a rare sibling clade to E-M35, known as E-V16 or E-M281. The discovery of M281 was announced by Semino et al. 2002, who found it in two Ethiopian Oromo. Trombetta et al. 2011 found 5 more Ethiopian individuals and an equivalent SNP to M281, V16. It was in the 2011 paper that the family tree position (M215+/M35-) was discovered as described above. The E-M215 derivative, E-M35 is defined by the M35 SNP. Individuals with no known subclade defining mutations are referred to as E-M35*. As of June 2015, there is an increasingly complex tree structure which divides most men in E-M35 into two branches: E-V68 and E-Z827.
The most frequently described subclades are E-M78, a part of E-V68, and E-M81, which is a branch of E-Z827. These two subclades represent the largest proportion of the modern E-M215 population. E-M78 is found over most of the range where E-M215 is found excluding Southern Africa. E-M81 is found mainly in North Africa. E-M123 is less common but widely scattered, with significant populations in specific parts of the Horn of Africa, the Levant, Arabia, Iberia, and Anatolia. 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 the E-V42, E-M293, E-V92 and E-V6 subclades, which were identified as E-M35 basal clades in a previous phylogeny.
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 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.
TMRCA of the major nodes in E-M215
|TMRCA (kya)||Trombetta 2015||YFull|
E-V68, is dominated by its longer-known subclade E-M78. Three "E-V68*" individuals who are in E-V68 but not E-M78 have been reported in Sardinia, by Trombetta et al. 2011, when announcing the discovery of V68. The authors noted that because E-V68* was not found in the Middle Eastern samples, this appears to be evidence of maritime migration from Africa to southwestern Europe. E-M78 is a commonly occurring subclade, widely distributed in North Africa, the Horn of Africa, West Asia, (the Middle East and Near East) "up to Southern Asia", and all of Europe. The European distribution has a frequency peak centered in parts of the Balkans (up to almost 50% in some areas) and Sicily, and declining frequencies evident toward western, central, and northeastern Europe.
Based on genetic STR variance data, Cruciani et al. 2007 suggests that E-M78 originated in the region of Egypt and Libya.[Note 4] about 18,600 years ago (17,300 - 20,000 years ago).[Note 5] Battaglia et al. 2008 describe Egypt as "a hub for the distribution of the various geographically localized M78-related subclades" and, based on archaeological data, they propose that the point of origin of E-M78 (as opposed to later dispersal from Egypt) may have been in a refugium which "existed on the border of present-day Sudan and Egypt, near Lake Nubia, until the onset of a humid phase around 8500 BC. The northward-moving rainfall belts during this period could have also spurred a rapid migration of Mesolithic foragers northwards in Africa, the Levant and ultimately onward to Asia Minor and Europe, where they each eventually differentiated into their regionally distinctive branches". Towards the south, Hassan et al. 2008 also explain evidence that some subclades of E-M78, specifically E-V12 and E-V22, "might have been brought to Sudan from North Africa after the progressive desertification of the Sahara around 6,000-8,000 years ago". And similarly, Cruciani et al. 2007 propose that E-M78 in Ethiopia, Somalia and surrounding areas, back-migrated to this region from the direction of Egypt after acquiring the E-M78 mutation.
Recently, E-M78 was dated by Trombetta et al. 2015. between 20,300 and 14,800 years ago.
Subclades of E-M78
Listed here are the main subclades of M78 as of June 2015. Within the E-M78 subclade, Trombetta et al. 2015 allocated most of the former E-M78* chromosomes to three new distinct branches: E-V1083*, E-V1477 and E-V259. The first is a paragroup sister to clades E-V22 and E-V13. The mutation V1477 defines a new basal branch that has been observed only in one northern African sample. Finally, a sister clade of E-V12 defined by V264 includes E-V65 and V259, a new lineage distributed in central Africa.
- E-M78 (E1b1b1a1). North Africa, Horn of Africa, West Asia, Europe. (Formerly "E1b1b1a".)
- E-M78* Found in southern Portugal, Spain and Iran (Tehran and Semnan provinces).
- E-V1477 Found in Tunisian Jews.
- E-V1083* Found only in Eritrea (1.1%) and Sardinia (0.3%).
- E-V13 This is the most common subclade of E-M215 found in Europe. It is especially common in the Balkans.
- E-V22. Concentrated in Northeast Africa and the Near East. Peaks among the Saho.
- E-V12. Found in Egypt, Sudan, and other places. Has an important subclade
- E-V12* Most common lineage among Southern Egyptians (74.5%).
- E-V32. Very common among Somalis, Tigre and Oromos.
- E-V259. Found in North Cameroon.
- E-V65. Associated with North Africa, but also found in Italy and Spain.
- E-V12. Found in Egypt, Sudan, and other places. Has an important subclade
- E-M521. Not mentioned by Trombetta et al.2015. Found in two individuals in Greece by Battaglia et al. 2008 and in one individual from the Eastern Alpine region of Italy by Coia et al. (2013)
In human genetics, E-Z827, is the name of a major human Y-chromosome DNA haplogroup abundantly found in North Africa, particularly the Maghreb, and to a lesser extent in Northeast Africa, the Near East and Europe.
E-V257* individuals in their samples who were E-V257, but not E-M81. A Borana from Kenya, a Marrakesh Berber, a Corsican, a Sardinian, a southern Spaniard and a Cantabrian. As mentioned above, Trombetta et al. 2011 propose that the absence of E-V257* in the Middle East makes a maritime movement from northern Africa to southern Europe the most plausible hypothesis so far to explain its distribution.
E-M81 is the most common Y-chromosome haplogroup in North Africa, dominated by its subclade E-M183. It is thought to have originated in the area of North Africa 14,200 years ago. This haplogroup reaches a mean frequency of 42% in North Africa, decreasing in frequency from approximately 80% or more in some Moroccan Berber populations, including Saharawis, to approximately 10% to the east of this range in Egypt. Because of its prevalence among these groups and also others such as Mozabite, Middle Atlas, Kabyle and other Berber groups, it is sometimes referred to as a genetic "Berber marker". Pereira et al. 2010 report high levels amongst Tuareg in two Saharan populations - 77.8% near Gorom-Gorom, in Burkina Faso, and 81.8% from Gosi in Mali. There was a much lower frequency of 11.1% in the vicinity of Tanut in Niger. E-M81 is also quite common among Maghrebi Arabic-speaking groups. It is generally found at frequencies around 45% in coastal cities of North Africa (Oran, Tizi Ouzou, Algiers, Tunis, Sousse).
In this key area from Egypt to the Atlantic Ocean, Arredi et al. 2004 report a pattern of decreasing microsatellite haplotype variation (implying greater 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 Libyan Desert in Egypt.
Arredi et al. 2004 believe the pattern of distribution and variance to be consistent with the hypothesis of a post Paleolithic "demic diffusion" from the East. The ancestral lineage of E-M81 in their hypothesis could have been linked with the spread of Neolithic food-producing technologies from the Fertile Crescent via the Nile, although pastoralism rather than agriculture. E-M81 may also have been carried into its currently most common region together with a form of the proto-Afroasiatic language. On the basis of these possible links, the men who brought E-M81 into northwestern Africa may therefore have come from Asia, or they may represent a "local contribution to the North African Neolithic transition". But 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 North Africa, or possibly as far south as the Horn of Africa.
In Europe, E-M81 is widespread but rare, in the Iberian Peninsula Spain shows an average frequency of 4.3% (49/1140) in the Iberian Peninsula with frequencies reaching 9% in Galicia, 10% in Western Andalusia and Northwest Castile. However this study 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, average for Iberian Peninsula is 4.9% (47/963), it is found at comparable levels to E-M78, with an average frequency of around 5%, and in some regions it is more common. Its frequencies are higher in the western half of the peninsula with frequencies reaching 8% in Extremadura and southern Portugal, 4% to 9% in Galicia, 14% in western Andalusia and 10% in northwest Castile and 9% to 17% in Cantabria. The highest frequencies of this clade found so far in Europe were observed in the Valles Pasiegos from Cantabria, ranging from 18% (8/45) to 41% (23/56). 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%).
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), in Sicily (approximately 2% overall, but up to 5% in Piazza Armerina), and in very much lower frequency near Lucera (1.7%), in continental Italy, possibly due to ancient migrations during the Islamic, Roman, and Carthaginian empires. E-M81 was also found in 2013 at 5.8% in a large sample of 1 204 Sardinians.
As a result of its old world distribution, this subclade is found throughout Latin America, for example 6.1% in Cuba, 5.4% in Brazil (Rio de Janeiro),[Note 6] and among Hispanic men from California and Hawaii 2.4%.
In smaller numbers, E-M81 men can be found in areas in contact with [North Africa, both around the Sahara, in places like Sudan, and around the Mediterranean in places like Lebanon, Turkey, and amongst Sephardi Jews.
There are two recognized subclades of E-M81, although one is much more important than the other.
E-M183 is extremely dominant within E-M81. Karafet et al. 2008 first described it as a subclade of E-M81. The known subclades of E-M183 include:
- E-M165 Underhill et al. 2000 found one example in Middle East.
- E-L351 Found in two related participants in The E-M35 Phylogeny Project.
This is a recently discovered subclade which has not yet been included in most haplogroup trees, E-Z830 includes the confirmed subclades of E-M123, E-V1515 (E-M293, E-V42, E-V6, E-V92), and E-Z830*, 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.
E-M123 is mostly known for its major subclade E-M34, which dominates this clade.[Note 7]
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.
E-M293 is a subclade of E-V1515. It was first 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 East Africa into Southern Africa. So far high levels have been found in specific ethnic groups in Tanzania and Southern Africa. Highest were the Datooga (43%), Khwe (Kxoe) (31%), Burunge (28%), and Sandawe peoples (24%). Henn (2008) in their study also found two Bantu-speaking Kenyan males with the M293 mutation.
Other E-M215 subclades are rare in Southern Africa. The authors state "Without information about M293 in the Maasai, Hema, and other populations in Kenya, Sudan, and Ethiopia, we cannot pinpoint the precise geographic source of M293 with greater confidence. However, the available evidence points to present-day Tanzania as an early and important geographic locus of M293 evolution.". They also say that "M293 is only found in sub-Saharan Africa, indicating a separate phylogenetic history for M35.1 * (former) samples further north".
E-P72 appears in Karafet (2008). Trombetta et al. 2011 announced that this is a subclade of E-M293.
Trombetta et al. 2011 announced the discovery of E-V42 in two Beta Israel persons. It was suggested that it may be restricted to the region around Ethiopia. However, further testing by commercial DNA testing companies confirmed many positive results for this subclade in Saudi Arabia , Kuwait and one person in Portugal who has a root from Arabia.
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 Somalis. Among the Ethiopian and Somali samples, the highest were 14.7% among the Amhara and 16.7% among the Wolayta.
To the south, Tishkoff et al. (2007) identified one V6+ man in a sample of 35 Datooga of Tanzania. And further to the north, Dugoujon et al. (2009) identified another 6 men in a sample of 93 from the Siwa Oasis, which is a Berber population
Trombetta et al. 2011 announced the discovery of E-V92 in two Amharas. Like E-V6 and E-V42 it possibly only exists in the area of Ethiopia.
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|
The following research teams per their publications were represented in the creation of the YCC Tree.
E-M215 and E1b1b1 are the currently accepted names found in the proposals of the Y Chromosome Consortium (YCC), for the clades defined by mutation M215 and M35 respectively, which can also be referred to as E-M215 and E-M35. The nomenclature E3b (E-M215) and E3b1 (E-M35) respectively were the YCC defined names used to designate the same haplogroups in older literature with E-M35 branching as a separate subclade of E-M215 in 2004. Prior to 2002 these haplogroups were not designated in a consistent way, and nor was their relationship to other related clades within haplogroup E and haplogroup DE. But in non-standard or older terminologies, E-M215 is for example approximately the same as "haplotype V", still used in publications such as Gérard et al. (2006).
|Wikiquote has quotations related to: Haplogroup E-M215 (Y-DNA)|
Y-DNA E Subclades
- Haplogroup E-L485
- Haplogroup E-M123
- Haplogroup E-M180
- Haplogroup E-M215
- Haplogroup E-M33
- Haplogroup E-M521
- Haplogroup E-M75
- Haplogroup E-M96
- Haplogroup E-P147
- Haplogroup E-P177
- Haplogroup E-P2
- Haplogroup E-V12
- Haplogroup E-V13
- Haplogroup E-V22
- Haplogroup E-V38
- Haplogroup E-V65
- Haplogroup E-V68
- Haplogroup E-Z820
- Haplogroup E-Z827
Y-DNA backbone tree
|Evolutionary tree of human Y-chromosome DNA haplogroups [χ 1][χ 2]|
|A00||A0-T [χ 3]|
|I||J||LT [χ 5]||K2|
|L||T||NO [χ 6]||K2b [χ 7]||K2c||K2d||K2e [χ 8]|
|N||O||K2b1 [χ 9]||P|
|M||S [χ 10]||Q||R|
- Cruciani et al. (2004) Semino et al. (2004) For E-M215 Cruciani et al. (2007) reduced their estimate to 22,400 from 25,600 in Cruciani et al. (2004), re-calibrating the same data.
- "Y chromosome data show a signal for a separate late-Pleistocene migration from Africa to Europe via Sinai as evidenced through the distribution of haplogroup E3b lineages, which is not manifested in mtDNA haplogroup distributions."Underhill & Kivisild (2007:547)
- "Paragroup E-M35 * and haplogroup J-12f2a* fit the criteria for major AJ founding lineages because they are widespread both in AJ populations and in Near Eastern populations, and occur at much lower frequencies in European non-Jewish populations." Behar et al. (2004)
- Cruciani et al. 2007 use the term Northeastern Africa to refer to Egypt and Libya, as shown in Table 1 of the study. Prior to Cruciani et al. 2007, Semino et al. 2004 East Africa as a possible place of origin of E-M78, based upon Ethiopian testing. This was because of the high frequency and diversity of E-M78 lineages in the region of Ethiopia. However, Cruciani et al. 2007 were able to study more data, including populations from North Africa who were not represented in the Semino et al. 2004 study, and found evidence that the E-M78 lineages which make up a significant proportion of some populations in that region, were relatively young branches (see E-V32 below). They therefore concluded that "Northeast Africa" was the likely place of origin of E-M78 based on "the peripheral geographic distribution of the most derived subhaplogroups with respect to northeastern Africa, as well as the results of quantitative analysis of UEP and microsatellite diversity". So according to Cruciani et al. 2007 E-M35, the parent clade of E-M78, originated in East Africa, subsequently spread to Northeast Africa, and then there was a "back migration" of E-M215 chromosomes that had acquired the E-M78 mutation. Cruciani et al. 2007 therefore note this as evidence for "a corridor for bidirectional migrations" between Northeast Africa (Egypt and Libya in their data) on the one hand and East Africa on the other. The authors believe there were "at least 2 episodes between 23.9–17.3 ky and 18.0–5.9 ky ago".
- Cruciani et al. 2007 use two calculation methods for estimating the age of E-M78 which give very different results. For the main 18,600 years ago, the ASD method is used, while for a second "ρ method", used as a check, gives 13.7kya with a standard deviation of 2.3kya, but the difference between the two methods is only large for the age estimation of E-M78, not its subclades. The authors state that the big difference is "attributable to the relevant departure from a star-like structure because of repeated founder effects"
- (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 9.6% in Portugal, quite similar to the frequency found in Rio de Janeiro (5.4%) among European contributors." Silva et al. 2006
- As of 11 November 2008 for example, the E-M35 phylogeny project had records of four E-M123* tests, compared to 93 test results with E-M34.
- Cruciani et al. (2004)
- Semino et al. (2004)
- ISOGG (2011) Cite error: Invalid
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- Karafet et al. (2008) Cite error: Invalid
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- Y Chromosome Consortium "YCC" (2002)
- Cruciani et al. (2007)
- Large-scale recent expansion of European patrilineages shown by population resequencing, Chiara Batini et al, nature.com, 2015
- Trombetta et al. 2015, Phylogeographic refinement and large scale genotyping of human Y chromosome haplogroup E provide new insights into the dispersal of early pastoralists in the African continent
- Henn et al. (2008) Cite error: Invalid
<ref>tag; name "Henn2008" defined multiple times with different content (see the help page).
- Hassan et al. (2008)
- Lacan et al. (2011)
- "Molecular genetic investigation of the Neolithic population history in the western Carpathian Basin" (PDF).
- Rosser et al. (2000)
- Firasat et al. (2006)
- Ehret et al. (2004)
- Keita & Boyce (2005)
- Keita 2008
- Behar et al. (2003)
- Behar et al. (2004)
- Shen et al. (2004)
- Adams et al. (2008)
- Nebel et al. (2001)
- ISOGG 2015
- Cadenas et al. 2007
- Trombetta et al. 2011
- Cruciani et al. 2006
- Peričic et al. 2005
- Cruciani et al. 2006
- "E-M81 YTree". www.yfull.com. Retrieved 2016-05-23.
- Arredi et al. 2004
- Alvarez et al. 2009
- Bosch et al. 2001
- Robino et al. 2008
- see table.
- Flores et al. 2005
- Beleza et al. 2006
- Capelli et al. 2009
- MacaMaca-Meyer et al. 2003
- Fregel et al. 2009, see table
- Ramos-Luisa et al. 2009
- Only men with French surname were analysed, in order to try to exclude more recent immigrants.
- Di Gaetano et al. 2009
- Francalacci et al. (2013), Low-Pass DNA Sequencing of 1200 Sardinians Reconstructs European Y-Chromosome Phylogeny
- (8 out of 132), Mendizabal et al. 2008
- (7 out of 295), Paracchini et al. 2003
- Adams, Susan M; Bosch, Elena; Balaresque, Patricia L.; Ballereau, Stéphane J.; Lee, Andrew C.; Arroyo, Eduardo; López-Parra, Ana M.; Aler, Mercedes; et al. (2008), "The Genetic Legacy of Religious Diversity and Intolerance: Paternal Lineages of Christians, Jews, and Muslims in the Iberian Peninsula", The American Journal of Human Genetics 83 (6): 725–36, doi:10.1016/j.ajhg.2008.11.007, PMC 2668061, PMID 19061982
- Alvarez; Santos, Cristina; Montiel, Rafael; Caeiro, Blazquez; Baali, Abdellatif; Dugoujon, Jean-Michel; Aluja, Maria Pilar (2009), "Y-chromosome variation in South Iberia: Insights into the North African contribution", American Journal of Human Biology 21 (3): 407–409, doi:10.1002/ajhb.20888, PMID 19213004
- Arredi, B; Poloni, E; Paracchini, S; Zerjal, T; Fathallah, D; Makrelouf, M; Pascali, V; Novelletto, A; Tylersmith, C (2004), "A Predominantly Neolithic Origin for Y-Chromosomal DNA Variation in North Africa", American Journal of Human Genetics 75 (2): 338–345, doi:10.1086/423147, PMC 1216069, PMID 15202071
- Badro, Danielle A.; Douaihy, Bouchra; Haber, Marc; Youhanna, Sonia C.; Salloum, Angélique; Ghassibe-Sabbagh, Michella; Johnsrud, Brian; Khazen, Georges; Matisoo-Smith, Elizabeth; Soria-Hernanz, David F.; Wells, R. Spencer; Tyler-Smith, Chris; Platt, Daniel E.; Zalloua, Pierre A. (2013), "Y-Chromosome and mtDNA Genetics Reveal Significant Contrasts in Affinities of Modern Middle Eastern Populations with European and African Populations", PLoS ONE 8 (1: e54616): e54616, doi:10.1371/journal.pone.0054616, PMC 3559847, PMID 23382925
- Battaglia, Vincenza; Fornarino, Simona; Al-Zahery, Nadia; Olivieri, Anna; Pala, Maria; Myres, Natalie M; King, Roy J; Rootsi, Siiri; et al. (2008), "Y-chromosomal evidence of the cultural diffusion of agriculture in southeast Europe", European Journal of Human Genetics 17 (6): 820–830, doi:10.1038/ejhg.2008.249, PMC 2947100, PMID 19107149
- Behar, Doron M.; Thomas, Mark G.; Skorecki, Karl; Hammer, Michael F.; Bulygina, Ekaterina; Rosengarten, Dror; Jones, Abigail L.; Held, Karen; et al. (October 2003), "Multiple Origins of Ashkenazi Levites: Y Chromosome Evidence for Both Near Eastern and European Ancestries", Am. J. Hum. Genet. 73 (4), pp. 768–779, doi:10.1086/378506, PMC 1180600, PMID 13680527. Also at http://www.ucl.ac.uk/tcga/tcgapdf/Behar-AJHG-03.pdf and http://www.familytreedna.com/pdf/400971.pdf
- Behar; Garrigan; Kaplan; Mobasher; Rosengarten (November 2004), "Contrasting patterns of Y chromosome variation in Ashkenazi Jewish and host non-Jewish European populations" (PDF), Hum. Genet., pp. 354–365
- Beleza, Sandra; Gusmao, Leonor; Lopes, Alexandra; Alves, Cintia; Gomes, Iva; Giouzeli, Maria; Calafell, Francesc; Carracedo, Angel; Amorim, Antonio (2006), "Micro-Phylogeographic and Demographic History of Portuguese Male Lineages", Annals of Human Genetics 70 (2): 181–194, doi:10.1111/j.1529-8817.2005.00221.x, PMID 16626329
- Bird, Steven (2007), "Haplogroup E3b1a2 as a Possible Indicator of Settlement in Roman Britain by Soldiers of Balkan Origin", Journal of Genetic Genealogy 3 (2)
- Bortolini; Thomas, Mark G.; Chikhi, Lourdes; Aguilar, Juan A.; Castro-De-Guerra, Dinorah; Salzano, Francisco M.; Ruiz-Linares, Andres (2004), "Ribeiro's typology, genomes, and Spanish colonialism, as viewed from Gran Canaria and Colombia" (PDF), Genetics and Molecular Biology 27 (1): 1–8, doi:10.1590/S1415-47572004000100001
- Bosch, Elena; Calafell, Francesc; Comas, David; Oefner, Peter J.; Underhill, Peter A.; Bertranpetit, Jaume (2001), "High-resolution analysis of human Y-chromosome variation shows a sharp discontinuity and limited gene flow between north-western Africa and the Iberian Peninsula", Am J Hum Genet 68 (4): 1019–1029, doi:10.1086/319521, PMC 1275654, PMID 11254456
- Bosch, E.; Calafell, F.; Gonzalez-Neira, A.; Flaiz, C.; Mateu, E.; Scheil, H.-G.; Huckenbeck, W.; Efremovska, L.; et al. (2006), "Paternal and maternal lineages in the Balkans show a homogeneous landscape over linguistic barriers, except for the isolated Aromuns", Annals of Human Genetics 70 (4): 459–487, doi:10.1111/j.1469-1809.2005.00251.x, PMID 16759179
- Cadenas; Zhivotovsky, Lev A; Cavalli-Sforza, Luca L; Underhill, Peter A; Herrera, Rene J (2007), "Y-chromosome diversity characterizes the Gulf of Oman", European Journal of Human Genetics 16 (3): 1–13, doi:10.1038/sj.ejhg.5201934, PMID 17928816
- Capelli, Cristian; Redhead, Nicola; Abernethy, Julia K.; Gratrix, Fiona; Wilson, James F.; Moen, Torolf; Hervig, Tor; Richards, Martin; et al. (2003), "A Y Chromosome Census of the British Isles", Current Biology 13 (11): 979–84, doi:10.1016/S0960-9822(03)00373-7, PMID 12781138 also at 
- Caratti; Gino, S.; Torre, C.; Robino, C. (2009), "Subtyping of Y-chromosomal haplogroup E-M78 (E1b1b1a) by SNP assay and its forensic application", International Journal of Legal Medicine 123 (4): 357–360, doi:10.1007/s00414-009-0350-y, PMID 19430804
- Capelli, Cristian; Onofri, Valerio; Brisighelli, Francesca; Boschi, Ilaria; Scarnicci, Francesca; Masullo, Mara; Ferri, Gianmarco; Tofanelli, Sergio; et al. (2009), "Moors and Saracens in Europe: estimating the medieval North African male legacy in southern Europe", European Journal of Human Genetics 17 (6): 848–852, doi:10.1038/ejhg.2008.258, PMC 2947089, PMID 19156170
- Cinnioğlu, Cengiz; King, Roy; Kivisild, Toomas; Kalfoglu, Ersi; Atasoy, Sevil; Cavalleri, Gianpiero L.; Lillie, Anita S.; Roseman, Charles C.; et al. (2004), "Excavating Y-chromosome haplotype strata in Anatolia" (PDF), Hum Genet 114 (2): 127–48, doi:10.1007/s00439-003-1031-4, PMID 14586639
- Contu, Daniela; Morelli, Daniela; Santoni, Federico; Foster, Jamie W.; Francalacci, Paolo; Cucca, Francesco (2008), "Y-Chromosome Based Evidence for Pre-Neolithic Origin of the Genetically Homogeneous but Diverse Sardinian Population: Inference for Association Scans" (PDF), PLoS ONE 3 (1): e1430, doi:10.1371/journal.pone.0001430, PMC 2174525, PMID 18183308
- Cruciani, Fulvio; Santolamazza, Piero; Shen, Peidong; MacAulay, Vincent; Moral, Pedro; Olckers, Antonel; Modiano, David; Holmes, Susan (2002), "A Back Migration from Asia to Sub-Saharan Africa Is Supported by High-Resolution Analysis of Human Y-Chromosome Haplotypes" (PDF), American Journal of Human Genetics 70 (5): 1197–1214, doi:10.1086/340257, PMC 447595, PMID 11910562
- Cruciani; La Fratta; Santolamazza; Sellitto (May 2004), "Phylogeographic Analysis of Haplogroup E3b (E-M215) Y Chromosomes Reveals Multiple Migratory Events Within and Out Of Africa" (PDF), American Journal of Human Genetics 74 (5): 1014–1022, doi:10.1086/386294, PMC 1181964, PMID 15042509
- Cruciani; La Fratta; Torroni; Underhill; Scozzari (2006), "Molecular Dissection of the Y Chromosome Haplogroup E-M78 (E3b1a): A Posteriori Evaluation of a Microsatellite-Network-Based Approach Through Six New Biallelic Markers" (PDF), Human Mutation 27 (8): 831–2, doi:10.1002/humu.9445, PMID 16835895
- Cruciani, F.; La Fratta, R.; Trombetta, B.; Santolamazza, P.; Sellitto, D.; Colomb, E. B.; Dugoujon, J.-M.; Crivellaro, F.; et al. (2007), "Tracing Past Human Male Movements in Northern/Eastern Africa and Western Eurasia: New Clues from Y-Chromosomal Haplogroups E-M78 and J-M12", Molecular Biology and Evolution 24 (6): 1300–1311, doi:10.1093/molbev/msm049, PMID 17351267 Also see Supplementary Data.
- Di Gaetano; Cerutti, Francesca; Crobu, Carlo; Robino (2009), "Differential Greek and northern African migrations to Sicily are supported by genetic evidence from the Y chromosome", European Journal of Human Genetics 17 (1): 91–99, doi:10.1038/ejhg.2008.120, PMC 2985948, PMID 18685561
- Dugoujon; Coudray; Torroni; Cruciani; Scozzari; Moral; Louali; Kossmann (2009), d'Errico; Hombert, eds., "The Berber and the Berbers: Genetic and linguistic diversities" (PDF), Becoming Eloquent Advances in the emergence of language, human cognition, and modern cultures
- Ehret, C.; Keita, SO; Newman, P (2004), "The Origins of Afroasiatic", Science 306 (5702): 1680, doi:10.1126/science.306.5702.1680c, PMID 15576591
- El-Sibai, Mirvat; Platt, Daniel E.; Haber, Marc; Xue, Yali; Youhanna, Sonia C.; Wells, R. Spencer; Izaabel, Hassan; Sanyoura, May F.; et al. (2009), "Geographical Structure of the Y-chromosomal Genetic Landscape of the Levant: A coastal-inland contrast", Annals of Human Genetics 73 (6): 568–581, doi:10.1111/j.1469-1809.2009.00538.x, PMC 3312577, PMID 19686289
- Firasat; Khaliq, Shagufta; Mohyuddin, Aisha; Papaioannou, Myrto; Tyler-Smith, Chris; Underhill, Peter A; Ayub, Qasim (2006), "Y-chromosomal evidence for a limited Greek contribution to the Pathan population of Pakistan", European Journal of Human Genetics 15 (1): 121–126, doi:10.1038/sj.ejhg.5201726, PMC 2588664, PMID 17047675
- Flores, Carlos; Maca-Meyer, Nicole; González, Ana M; Oefner, Peter J; Shen, Peidong; Pérez, Jose A; Rojas, Antonio; Larruga, Jose M; Underhill, Peter A (2004), "Reduced genetic structure of the Iberian peninsula revealed by Y-chromosome analysis: implications for population demography" (PDF), European Journal of Human Genetics 12 (10): 855–863, doi:10.1038/sj.ejhg.5201225, PMID 15280900
- Flores; Maca-Meyer, Nicole; Larruga, Jose M.; Cabrera, Vicente M.; Karadsheh, Naif; Gonzalez, Ana M. (2005), "Isolates in a corridor of migrations: a high-resolution analysis of Y-chromosome variation in Jordan", J Hum Genet 50 (9): 435–441, doi:10.1007/s10038-005-0274-4, PMID 16142507
- Francalacci, P.; Morelli, L.; Underhill, P.A.; Lillie, A.S.; Passarino, G.; Useli, A.; Madeddu, R.; Paoli, G.; et al. (2003), "Peopling of Three Mediterranean Islands (Corsica, Sardinia, and Sicily) Inferred by Y-Chromosome Biallelic Variability", American Journal of Physical Anthropology 121 (3): 270–279, doi:10.1002/ajpa.10265, PMID 12772214 replacement character in
|last10=at position 4 (help)
- Fregel, Rosa; Gomes, Verónica; Gusmão, Leonor; González, Ana M; Cabrera, Vicente M; Amorim, António; Larruga, Jose M (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
- Gérard; Berriche, S; Aouizérate, A; Diéterlen, F; Lucotte, G (2006), "North African Berber and Arab influences in the western Mediterranean revealed by Y-chromosome DNA haplotypes", Human Biology 78 (3): 307–316, doi:10.1353/hub.2006.0045, PMID 17216803
- Gonçalves, R; Freitas, A; Branco, M; Rosa, A; Fernandes, AT; Zhivotovsky, LA; Underhill, PA; Kivisild, T; Brehm, A (2005), "Y-chromosome Lineages from Portugal, Madeira and Açores Record Elements of Sephardim and Berber Ancestry", Annals of Human Genetics 69 (Pt 4): 443–454, doi:10.1111/j.1529-8817.2005.00161.x, PMID 15996172
- Hammer (2003), "Human population structure and its effects on sampling Y chromosome sequence variation", Genetics 164 (4): 1495–1509, PMC 1462677, PMID 12930755
- Hassan, Hisham Y.; Underhill, Peter A.; Cavalli-Sforza, Luca L.; Ibrahim, Muntaser E. (2008), "Y-Chromosome Variation Among Sudanese: Restricted Gene Flow, Concordance With Language, Geography, and History" (PDF), American Journal of Physical Anthropology 137 (3): 316–23, doi:10.1002/ajpa.20876, PMID 18618658
- Henn, B. M.; Gignoux, C.; Lin, Alice A; Oefner, Peter J.; Shen, P.; Scozzari, R.; Cruciani, F.; Tishkoff, S. A.; Mountain, J. L.; Underhill, P. A. (2008), "Y-chromosomal evidence of a pastoralist migration through Tanzania to southern Africa", PNAS 105 (31): 10693–8, doi:10.1073/pnas.0801184105, PMC 2504844, PMID 18678889. See comment on Dienekes blog, comment on the Spitoon blog and public release.
- ISOGG (2011), Y-DNA Haplogroup E and its Subclades - 2011, International Society of Genetic Genealogists "ISOGG"
- Jobling, M.A.; Tyler-Smith, C. (2000), "New uses for new haplotypes the human Y chromosome, disease and selection", Trends Genet. 16 (8): 356–362, doi:10.1016/S0168-9525(00)02057-6, PMID 10904265
- Karafet, T. M.; Mendez, F. L.; Meilerman, M. B.; Underhill, P. A.; Zegura, S. L.; Hammer, M. F. (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. Published online April 2, 2008. See also Supplementary Material.
- Keita, Shomarka (2008), "Geography, selected Afro-Asiatic families, and Y chromosome lineage variation", In Hot Pursuit of Language in Prehistory: Essays in the Four Fields of Anthropology : In Honor of Harold Crane Fleming, ISBN 978-90-272-3252-6
- Keita, S. O. Y.; Boyce, A. J. (Anthony J.) (2005), "Genetics, Egypt, and History: Interpreting Geographical Patterns of Y Chromosome Variation", History in Africa 32: 221–246, doi:10.1353/hia.2005.0013
- King, R. J.; Özcan, S. S.; Carter, T.; Kalfoğlu, E.; Atasoy, S.; Triantaphyllidis, C.; Kouvatsi, A.; Lin, A. A.; et al. (2008), "Differential Y-chromosome Anatolian Influences on the Greek and Cretan Neolithic" (PDF), Annals of Human Genetics 72 (2): 205–214, doi:10.1111/j.1469-1809.2007.00414.x, PMID 18269686
- King; Underhill (2002), "Congruent distribution of Neolithic painted pottery and ceramic figurines with Y-chromosome lineages", Antiquity 76: 707–14
- Kujanova; Pereira; Fernandes; Pereira; Cerný (2009), "Near Eastern Neolithic Genetic Input in a Small Oasis of the Egyptian Western Desert", American Journal of Physical Anthropology 140 (2): 336–346, doi:10.1002/ajpa.21078, PMID 19425100
- Lacan, Marie; Keyser, Christine; Ricaut, François-Xavier; Brucato, Nicolas; Tarrús, Josep; Bosch, Angel; Guilaine, Jean; Crubézy, Eric; Ludes, Bertrand (2011), "Ancient DNA suggests the leading role played by men in the Neolithic dissemination", PNAS 108 (45): 18255–9, doi:10.1073/pnas.1113061108, PMC 3215063, PMID 22042855
- Lancaster, Andrew (2009), "Y Haplogroups, Archaeological Cultures and Language Families: a Review of the Multidisciplinary Comparisons using the case of E-M35" (PDF), Journal of Genetic Genealogy 5 (1)
- Luis, J; Rowold, D; Regueiro, M; Caeiro, B; Cinnioglu, C; Roseman, C; Underhill, P; Cavallisforza, L; Herrera, R (2004), "The Levant versus the Horn of Africa: Evidence for Bidirectional Corridors of Human Migrations" (PDF), American Journal of Human Genetics 74 (3): 532–544, doi:10.1086/382286, PMC 1182266, PMID 14973781. (Also see Errata)
- Maca-Meyer N, Sánchez-Velasco P, Flores C, Larruga JM, González AM, Oterino A, Leyva-Cobián F, et al. (2003), "Y Chromosome and Mitochondrial DNA Characterization of Pasiegos, a Human Isolate from Cantabria (Spain)", Annals of Human Genetics 67 (Pt 4): 329–339, doi:10.1046/j.1469-1809.2003.00045.x, PMID 12914567.
- Martinez, Laisel; Underhill, Peter A; Zhivotovsky, Lev A; Gayden, Tenzin; Moschonas, Nicholas K; Chow, Cheryl-Emiliane T; Conti, Simon; Mamolini, Elisabetta; Cavalli-Sforza, L Luca; Herrera, Rene (April 1, 2007), "Paleolithic Y-haplogroup heritage predominates in a Cretan highland plateau", European Journal of Human Genetics 15 (4): 485–493, doi:10.1038/sj.ejhg.5201769, ISSN 1018-4813, PMID 17264870
- Mendizabal, Isabel; Sandoval, Karla; Berniell-Lee, Gemma; Calafell, Francesc; Salas, Antonio; Martinez-Fuentes, Antonio; Comas, David (2008), "Genetic origin, admixture, and asymmetry in maternal and paternal human lineages in Cuba", BMC Evol Biol. 8: 213, doi:10.1186/1471-2148-8-213, PMC 2492877, PMID 18644108
- Nebel; Filon, D; Brinkmann, B; Majumder, P; Faerman, M; Oppenheim, A (2001), "The Y Chromosome Pool of Jews as Part of the Genetic Landscape of the Middle East", American Journal of Human Genetics 69 (5): 1095–1112, doi:10.1086/324070, PMC 1274378, PMID 11573163
- Onofri, Valerio; Alessandrini, Federica; Turchi, Chiara; Pesaresi, Mauro; Buscemi, Loredana; Tagliabracci, Adriano (2006), "Development of multiplex PCRs for evolutionary and forensic applications of 37 human Y chromosome SNPs" (PDF), Forensic Science International 157 (1): 23–35, doi:10.1016/j.forsciint.2005.03.014, PMID 15896936
- Paracchini; Pearce, CL; Kolonel, LN; Altshuler, D; Henderson, BE; Tyler-Smith, C (2003), "A Y chromosomal influence on prostate cancer risk: the multi-ethnic cohort study", J Med Genet 40 (11): 815–819, doi:10.1136/jmg.40.11.815, PMC 1735314, PMID 14627670
- Pelotti; Ceccardi, S; Lugaresi, F; Trane, R; Falconi, M; Bini, C; Willuweit, S; Roewer, L (2007), "Microgeographic genetic variation of Y chromosome in a population sample of Ravenna's area in the Emilia-Romagna region (North of Italy)", Forensic Science International: Genetics Supplement Series 1 (1): 242–243, doi:10.1016/j.fsigss.2007.10.025
- Pereira, Luísa; Černý, Viktor; Cerezo, María; Silva, Nuno M; Hájek, Martin; Vašíková, Alžběta; Kujanová, Martina; Brdička, Radim; Salas, Antonio (2010), "Linking the sub-Saharan and West Eurasian gene pools: maternal and paternal heritage of the Tuareg nomads from the African Sahel" (PDF), European Journal of Human Genetics 18 (8): 915–923, doi:10.1038/ejhg.2010.21, PMC 2987384, PMID 20234393
- Peričic, M.; Lauc, LB; Klarić, IM; Rootsi, S; Janićijevic, B; Rudan, I; Terzić, R; Colak, I; et al. (2005), "High-resolution phylogenetic analysis of southeastern Europe traces major episodes of paternal gene flow among Slavic populations", Mol. Biol. Evol. 22 (10), pp. 1964–75, doi:10.1093/molbev/msi185, PMID 15944443.
- Pontikos D. "Phylogeographic refinement of haplogroup E" http://dienekes.blogspot.ru/2015/07/phylogeographic-refinement-of.html
- Ramos-Luisa, E.; Blanco-Verea, A.; Brión, M.; Van Huffel, V.; Carracedo, A.; Sánchez-Diz, P. (2009), "Phylogeography of French male lineages (unpublished data 23rd International ISFG Congress)" (PDF), Forensic Science International 2: 439–441, doi:10.1016/j.fsigss.2009.09.026
- Regueiro, M.; Cadenas, A.M.; Gayden, T.; Underhill, P.A.; Herrera, R.J. (2006), "Iran: Tricontinental Nexus for Y-Chromosome Driven Migration" (PDF), Hum Hered 61 (3): 132–143, doi:10.1159/000093774, PMID 16770078
- Robino, C.; Crobu, F.; Gaetano, C.; Bekada, A.; Benhamamouch, S.; Cerutti, N.; Piazza, A.; Inturri, S.; Torre, C. (2008), "Analysis of Y-chromosomal SNP haplogroups and STR haplotypes in an Algerian population sample", Journal International Journal of Legal Medicine 122 (3): 251–5, doi:10.1007/s00414-007-0203-5, PMID 17909833
- Rosa, Alexandra; Ornelas, Carolina; Jobling, Mark A; Brehm, António; Villems, Richard (2007), "Y-chromosomal diversity in the population of Guinea-Bissau: a multiethnic perspective" (PDF), BMC Evolutionary Biology 7: 124, doi:10.1186/1471-2148-7-124, PMC 1976131, PMID 17662131
- Rosser, Z; Zerjal, T; Hurles, M; Adojaan, M; Alavantic, D; Amorim, A; Amos, W; Armenteros, M; et al. (2000), "Y-Chromosomal Diversity in Europe Is Clinal and Influenced Primarily by Geography, Rather than by Language", American Journal of Human Genetics 67 (6): 1526–1543., doi:10.1086/316890, PMC 1287948, PMID 11078479
- Sanchez, Juan J; Hallenberg, Charlotte; Børsting, Claus; Hernandez, Alexis; Gorlin, RJ (2005), "High frequencies of Y chromosome lineages characterized by E3b1, DYS19-11, DYS392-12 in Somali males", European Journal of Human Genetics 13 (7): 856–866, doi:10.1038/sj.ejhg.5201390, PMID 15756297. Published online 9 March 2005
- Scozzari, Rosaria; Cruciani, F; Pangrazio, A; Santolamazza, P; Vona, G; Moral, P; Latini, V; Varesi, L; et al. (2001), "Human Y-Chromosome Variation in the Western Mediterranean Area: Implications for the Peopling of the Region" (PDF), Human Immunology 62 (9): 871–884, doi:10.1016/S0198-8859(01)00286-5, PMID 11543889
- Semino, O.; Passarino, G; Oefner, PJ; Lin, AA; Arbuzova, S; Beckman, LE; De Benedictis, G; Francalacci, P; et al. (2000), "The Genetic Legacy of Paleolithic Homo sapiens sapiens in Extant Europeans: A Y Chromosome Perspective" (PDF), Science 290 (5494), pp. 1155–59, doi:10.1126/science.290.5494.1155, PMID 11073453.
- Semino; Santachiara-Benerecetti, A. Silvana; Falaschi, Francesco; Cavalli-Sforza, L. Luca; Underhill, Peter A. (2002), "Ethiopians and Khoisan share the deepest clades of the human Y-chromosome phylogeny" (PDF), Am J Hum Genet 70 (1), pp. 265–268, doi:10.1086/338306, PMC 384897, PMID 11719903
- Semino, Ornella; Magri, Chiara; Benuzzi, Giorgia; Lin, Alice A.; Al-Zahery, Nadia; Battaglia, Vincenza; MacCioni, Liliana; Triantaphyllidis, Costas; et al. (2004), "Origin, Diffusion, and Differentiation of Y-Chromosome Haplogroups E and J: Inferences on the Neolithization of Europe and Later Migratory Events in the Mediterranean Area", American Journal of Human Genetics 74 (5), pp. 1023–1034, doi:10.1086/386295, PMC 1181965, PMID 15069642
- Shen, Peidong; Lavi, Tal; Kivisild, Toomas; Chou, Vivian; Sengun, Deniz; Gefel, Dov; Shpirer, Issac; Woolf, Eilon; et al. (2004), "Reconstruction of Patrilineages and Matrilineages of Samaritans and Other Israeli Populations From Y-Chromosome and Mitochondrial DNA Sequence Variation" (PDF), Human Mutation 24 (3), pp. 248–60, doi:10.1002/humu.20077, PMID 15300852
- Silva; Carvalho, Elizeu; Costa, Guilherme; Tavares, Lígia; Amorim, António; Gusmão, Leonor (2006), "Y-chromosome genetic variation in Rio De Janeiro population", American Journal of Human Biology 18 (6): 829–837, doi:10.1002/ajhb.20567, PMID 17039481
- Shlush; Behar, Doron M.; Yudkovsky, Guennady; Templeton, Alan; Hadid, Yarin; Basis, Fuad; Hammer, Michael; Itzkovitz, Shalev; Skorecki, Karl (2008), Gemmell, Neil John, ed., "The Druze: A Population Genetic Refugium of the Near East", PLoS ONE 3 (5): e2105, doi:10.1371/journal.pone.0002105, PMC 2324201, PMID 18461126
- Sykes, Bryan (2006), Blood of the Isles: Exploring the Genetic Roots of Our Tribal History, Bantam, ISBN 0-593-05652-3
- Tishkoff; Gonder; Henn; Mortensen; Knight (2007), "History of Click-Speaking Populations of Africa Inferred from mtDNA and Y Chromosome Genetic Variation", Mol Biol Evol 24 (10): 2180–2195, doi:10.1093/molbev/msm155, PMID 17656633
- Thomas; Stumpf, M. P.H; Harke, H. (2006), "Evidence for an apartheid-like social structure in early Anglo-Saxon England" (PDF), Proceedings of the Royal Society B 273 (1601): 2651–2657, doi:10.1098/rspb.2006.3627, PMC 1635457, PMID 17002951
- Trombetta, Beniamino; Cruciani, Fulvio; Sellitto, Daniele; Scozzari, Rosaria (2011), MacAulay, Vincent, ed., "A New Topology of the Human Y Chromosome Haplogroup E1b1 (E-P2) Revealed through the Use of Newly Characterized Binary Polymorphisms", PLoS ONE 6 (1): e16073, doi:10.1371/journal.pone.0016073, PMC 3017091, PMID 21253605
- Trombetta B. "Phylogeographic Refinement and Large Scale Genotyping of Human Y Chromosome Haplogroup E Provide New Insights into the Dispersal of Early Pastoralists in the African Continent" http://gbe.oxfordjournals.org/content/7/7/1940.long
- Underhill, Peter A.; Shen, Peidong; Lin, Alice A.; Jin, Li; Passarino, Giuseppe; Yang, Wei H.; Kauffman, Erin; Bonné-Tamir, Batsheva; et al. (2000), "Y chromosome sequence variation and the history of human populations", Nat Genet 26 (3), pp. 358–361, doi:10.1038/81685, PMID 11062480
- Underhill; Passarino, G.; Lin, A. A.; Shen, P.; Mirazon Lahr, M.; Foley, R. A.; Oefner, P. J.; Cavalli-Sforza, L. L. (2001), "The phylogeography of Y chromosome binary haplotypes and the origins of modern human populations" (PDF), Ann Hum Genet 65 (Pt 1), pp. 43–62, doi:10.1046/j.1469-1809.2001.6510043.x, PMID 11415522
- Underhill (2002), Bellwood and Renfrew, ed., Inference of Neolithic Population Histories using Y-chromosome Haplotypes, Cambridge: McDonald Institute for Archaeological Research, ISBN 1-902937-20-1
- Underhill, Peter A.; Kivisild, Toomas (2007), "Use of Y Chromosome and Mitochondrial DNA Population Structure in Tracing Human Migrations", Annu. Rev. Genet. 41: 539–64, doi:10.1146/annurev.genet.41.110306.130407, PMID 18076332
- Weale, M. E.; Weiss, D. A.; Jager, R. F.; Bradman, N.; Thomas, M. G. (2002), "Y Chromosome Evidence for Anglo-Saxon Mass Migration" (PDF), Mol. Biol. Evol. 19 (7), pp. 1008–1021, doi:10.1093/oxfordjournals.molbev.a004160, PMID 12082121.
- Weale; Shah, T; Jones, AL; Greenhalgh, J; Wilson, JF; Nymadawa, P; Zeitlin, D; Connell, BA; et al. (September 1, 2003), "Rare Deep-Rooting Y Chromosome Lineages in Humans: Lessons for Phylogeography", Genetics 165 (1), pp. 229–234, PMC 1462739, PMID 14504230
- Wood; Stover; Ehret; Destro-Bisol (2005), "Contrasting patterns of Y chromosome and mtDNA variation in Africa: evidence for sex-biased demographic processes", European Journal of Human Genetics 13: 867–876, doi:10.1038/sj.ejhg.5201408, PMID 15856073
- Y Chromosome Consortium "YCC" (2002), "A Nomenclature System for the Tree of Human Y-Chromosomal Binary Haplogroups", Genome Research 12 (2), pp. 339–348, doi:10.1101/gr.217602, PMC 155271, PMID 11827954
- Zalloua, Pierre A.; Xue, Yali; Khalife, Jade; Makhoul, Nadine; Debiane, Labib; Platt, Daniel E.; Royyuru, Ajay K.; Herrera, Rene J.; et al. (2008), "Y-Chromosomal Diversity in Lebanon Is Structured by Recent Historical Events", American Journal of Human Genetics 82 (4): 873–882, doi:10.1016/j.ajhg.2008.01.020, PMC 2427286, PMID 18374297
- Zalloua, Pierre A.; Platt, Daniel E.; El Sibai, Mirvat; Khalife, Jade; Makhoul, Nadine; Haber, Marc; Xue, Yali; Izaabel, Hassan; et al. (2008), "Identifying Genetic Traces of Historical Expansions: Phoenician Footprints in the Mediterranean" (PDF), The American Journal of Human Genetics 83 (5): 633–642, doi:10.1016/j.ajhg.2008.10.012, PMC 2668035, PMID 18976729
- Zerjal (1999), The use of Y-chromosomal DNA variation to investigate population history; in Papiha SS, Deka R, Chakraborty R (eds): Genomic diversity: applications in human population genetics, Kluwer Academic/Plenum Publishers, pp. 91–101
- Zhao; Khan, Faisal; Borkar, Minal; Herrera, Rene; Agrawal, Suraksha (2009), "Presence of three different paternal lineages among North Indians: A study of 560 Y chromosomes", Annals of Human Biology 36 (1): 1–14, doi:10.1080/03014460802558522, PMC 2755252, PMID 19058044
- E-M35 Phylogeny Project Wiki
- E-M35 Phylogeny Project (all labs site)
- E-M35 Phylogeny Project Public Forum
- Jewish E3b Project at FTDNA
- Map of E-M35 distribution in Europe
- Haplogroup E3B1 (E1b1b1) Facebook Group (M35 community)
- E1b1b1b*-A Project
- E1b1.org – International Y-DNA project of Haplogroup E1b1 and its Subclades