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==Neolithic==
==Neolithic==
The change from [[Hunter-gatherer|hunting and gathering]] to agriculture during the [[Neolithic Revolution]] was a watershed in world history. The societies that first made the change to agriculture are believed to have lived in the Middle East around 10,000 BCE. Agriculture was introduced into Europe by migrating farmers from the Middle East.<ref>{{harvcoltxt|Brace|Seguchi|Quintyn|Fox|2006}}</ref> According to the [[demic diffusion]] model, these Middle Eastern farmers either replaced or interbred with the local hunter-gather populations that had been living in Europe since the Out of Africa migration.{{sfn|Cavalli-Sforza|1993}}
The change from [[Hunter-gatherer|hunting and gathering]] to agriculture during the [[Neolithic Revolution]] was a watershed in world history. The societies that first made the change to agriculture are believed to have lived in the Middle East around 10,000 BCE. Agriculture was introduced into Europe by migrating farmers from the Middle East.<ref>{{harvcoltxt|Brace|Seguchi|Quintyn|Fox|2006}}</ref> According to the [[demic diffusion]] model, these Middle Eastern farmers either replaced or interbred with the local hunter-gather populations that had been living in Europe since the Out of Africa migration.<!--{{sfn|Cavalli-Sforza|1993}}-->{{citation needed}}


It has been suggested that the first Middle Eastern farmers reflected North African influences.{{sfnp|Bar-Yosef|1987}} There have been suggestions that some genetic lineages found in the Middle East arrived there during this period.<ref name=underkivis>{{cite journal | vauthors = Underhill PA, Kivisild T | title = Use of y chromosome and mitochondrial DNA population structure in tracing human migrations | journal = Annual Review of Genetics | volume = 41 | issue = 1 | pages = 539–64 | year = 2007 | pmid = 18076332 | doi = 10.1146/annurev.genet.41.110306.130407}}</ref> The first agricultural societies in the Middle East are generally thought to have emerged after, and perhaps from, the [[Natufian culture]] between 12,000 and 10,000 BCE. The latter group was widely semi-sedentary even before the introduction of agriculture. An important migration from North Africa across the [[Sinai Peninsula|Sinai]] also appears to have occurred before the formation of the Natufian.{{citation needed|date=August 2015}}.
It has been suggested that the first Middle Eastern farmers reflected North African influences.{{sfnp|Bar-Yosef|1987}} There have been suggestions that some genetic lineages found in the Middle East arrived there during this period.<ref name=underkivis>{{cite journal | vauthors = Underhill PA, Kivisild T | title = Use of y chromosome and mitochondrial DNA population structure in tracing human migrations | journal = Annual Review of Genetics | volume = 41 | issue = 1 | pages = 539–64 | year = 2007 | pmid = 18076332 | doi = 10.1146/annurev.genet.41.110306.130407}}</ref> The first agricultural societies in the Middle East are generally thought to have emerged after, and perhaps from, the [[Natufian culture]] between 12,000 and 10,000 BCE. The latter group was widely semi-sedentary even before the introduction of agriculture. An important migration from North Africa across the [[Sinai Peninsula|Sinai]] also appears to have occurred before the formation of the Natufian.{{citation needed|date=August 2015}}.

Revision as of 14:05, 3 August 2021

Map of Sub-Saharan African and North African admixture in European populations
5 component admixture plots for European, West Asian, North African and West African populations
For the African ancestry of all non-African Paleolithic populations, see recent African origin, Southern Dispersal.

African admixture in Europe refers to the presence of admixture events attributable to dispersal of populations inhabiting Africa in the genetic history of Europe. Certain Y-DNA and mtDNA lineages are thought to have spread from Northeastern Africa to the Near East during the later Pleistocene, and from there to Europe with the Neolithic Revolution.[1][2] More recent, direct African admixture – primarily Berber admixture from North Africa – is associated with the Carthaginian period as well as Muslim conquests of the Early Middle Ages, and is primarily concentrated in the Iberian peninsula, averaging from ~11% in the west and south to ~3% in the northeast, dropping to close to 0% in the Basque region.[3][4][5][6][7] North African admixture has also been detected in the island of Sicily.[3][8][9][10][11][12]

Neolithic

The change from hunting and gathering to agriculture during the Neolithic Revolution was a watershed in world history. The societies that first made the change to agriculture are believed to have lived in the Middle East around 10,000 BCE. Agriculture was introduced into Europe by migrating farmers from the Middle East.[13] According to the demic diffusion model, these Middle Eastern farmers either replaced or interbred with the local hunter-gather populations that had been living in Europe since the Out of Africa migration.[citation needed]

It has been suggested that the first Middle Eastern farmers reflected North African influences.[14] There have been suggestions that some genetic lineages found in the Middle East arrived there during this period.[15] The first agricultural societies in the Middle East are generally thought to have emerged after, and perhaps from, the Natufian culture between 12,000 and 10,000 BCE. The latter group was widely semi-sedentary even before the introduction of agriculture. An important migration from North Africa across the Sinai also appears to have occurred before the formation of the Natufian.[citation needed].

Historical period

In historical times, there has been a period of north African influence in southern Europe, especially western-southern Iberia and parts of southern Italy (namely Sicily), during various Muslim conquests. The genetic effect of this period on modern European populations is the subject of discussion (see below). In more recent history, the peoples of Europe and Africa came into contact during the exploration and colonization of Africa and as a consequence of the Atlantic slave trade.[2]

Defining African admixture

Generally, markers and lineages used to characterize African admixture are those that are believed to be specific to Africa. There are also DNA polymorphisms that are shared between populations native to Europe, West Asia, North Africa and the Horn of Africa, such as the y-chromosomal haplogroup E1b1b and the mitochondrial haplogroup M1.[2]

With regard to the paternal haplogroup E1b1b and maternal haplogroup M1, derivatives of these clades have been observed in prehistoric human fossils excavated at the Ifri n'Amr or Moussa site in Morocco, which have been radiocarbon-dated to the Early Neolithic period (ca. 5,000 BC). Ancient DNA analysis of these specimens indicates that they carried paternal haplotypes related to the E1b1b1b1a (E-M81) subclade and the maternal haplogroups U6a and M1, all of which are frequent among present-day communities in the Maghreb. These ancient individuals also bore an autochthonous Maghrebi genomic component that peaks among modern Berbers, indicating that they were ancestral to populations in the area. Additionally, fossils excavated at the Kelif el Boroud site near Rabat were found to carry the broadly-distributed paternal haplogroup T-M184 as well as the maternal haplogroups K1, T2 and X2, the latter of which were common mtDNA lineages in Neolithic Europe and Anatolia. These ancient individuals likewise bore the Berber-associated Maghrebi genomic component. This altogether indicates that the Late Neolithic Kelif el Boroud inhabitants were ancestral to contemporary populations in the area, but also likely experienced gene flow from Europe.[16]

Other lineages that are now found in Africa and Europe may have a common origin in Asia (e.g. Y haplogroups R1, and some paternal haplogroup T and U subclades). One subclade of haplogroup U, namely U6a1, is known to have expanded from northern and eastern Africa back into Europe[17][18] even though haplogroup U6 is considered to have originated in the Middle East. Other lineages are known to have moved from Europe directly into Africa, for example mitochondrial haplogroups H1 and H3.[19] Such bidirectional migrations between Africa and Eurasia complicate the task of defining admixture.

Y-DNA

One proposed example of Holocene gene flow from North Africa to Europe, via the Middle East, is thought to be E1b1b, which is thought to have emerged about 40,000 years ago in north-east Africa, and branches of it are thought to have migrated to the Middle East by 14,000 years ago during the late Pleistocene period.[20][15][21]

Entering the late mesolithic Natufian culture, the E1b1b1a2 (E-V13) subclade has been associated with the spread of farming from the Middle East into Europe either during or just before the Neolithic transition. E1b1b1 lineages are found throughout Europe but are distributed along a south-to-north cline, with an E1b1b1a mode in the Balkans.[1][22][23][a][b]

In separate migrations, E lineages in the form of the E1b1b1b subclade appear to have entered Europe from Northwest Africa into Iberia. In a sample of European males, Cruciani et al. observed haplogroup E at a frequency of 7.2%. The timing of this movement has been given widely varying estimates.[24]

In much of Europe, frequencies of E lineages are very low, usually less than 1%. For example, Cruciani et al. (2004) report such lineages at 2% in southern Portugal, 4% in northern Portugal, 2.9% in Istanbul, and 4.3% among Turkish Cypriots. E1b1a is closely related to E1b1b, the most frequent clade in Europe. E lineages that are not E1b1a or E1b1b could therefore reflect either a recent expansion associated with E1b1a or ancient population movements associated with E1b1b. For example, haplogroup E1a lineages have been detected in Portugal (5/553 = 1%),[25] among Italians in Calabria (1/80=1.3%), and among Albanians in Calabria (2/68=2.9%).[22] According to a study by Gonçalves et al. (2005), the distribution of haplogroup E1a lineages in Portugal was independent of the distribution of the younger and more ubiquitous E1b1a. The authors suggest that this distribution is consistent with a prehistoric migration from Africa to Iberia, possibly alongside mtDNA haplogroup U6.

Haplogroups A and B are thought to have been the predominant haplogroups in central and southern Africa prior to the Bantu Expansion. Currently these haplogroups are less common than E lineages. In a sample of 5,000 African men, haplogroup A had a frequency of 5%. Haplogroup A has rare occurrences in Europe, but recently the haplogroup was detected in seven indigenous British males with the same Yorkshire surname.[26]

E3b1 and its haplogroups E-M81 and E-M78 in North Africa, and E-M123 in the Near East

The subclade E3b1 (probably originating in northeastern Africa) has a wide distribution in North Africa, the Horn of Africa, the Middle East, and Europe. This haplogroup, in Italy, is represented by E-M78, E-M123 and E-M81 (Figure 3)[27] and reaches a frequency of 8% in northern and central Italy and slightly higher, 11%, in the south of that country.[27]
It has also been argued that the European distribution of E3b1 is compatible with the Neolithic demic diffusion of agriculture; thus, two subclades—E3b1a-M78 and E3b1c-M123—present a higher occurrence in Anatolia, the Balkans, and the Italian peninsula. Another subclade, E3b1b-M81 is associated with Berber populations and is commonly found in regions that have had historical gene flow with northern Africa, such as the Iberian Peninsula, the Canary Islands, and Sicily.[27]

North African Y-DNA E-M81 was found at a total of 41.1% among "pasiegos" from Cantabria, Spain. That is the highest frequency observed in Europe to date.[1] In Sardinians, Sub-Saharan Y-DNA lineages A1b1b2b and E1a1 were found at a total of 1.0% (A1b1b2b 0.5% / E1a1 0.5%).[28]

In Majorcans, Sub-Saharan Y-DNA lineage E-V38 was found at a total of 3.2% (2/62).[29]

Sub-Saharan Y-DNA lineages E3a, E1, BC*, (xE3), and E3* are found between 1 and 5% in Portugal, Valencia, Majorca, Cantabria, Málaga, Seville, and Galicia (Spain).[30][31]

mtDNA

Haplogroup L lineages are relatively infrequent (1% or less) throughout Europe with the exception of Iberia (Spain and Portugal), where frequencies as high as 22% have been reported, and some regions of Southern Italy, where frequencies as high as 2% and 3% have been found. According to a study in 2012 by Cerezo et al., about 65% of the European L lineages most likely arrived in rather recent historical times (Romanization period, Arab conquest of the Iberian Peninsula and Sicily, Atlantic slave trade) and about 35% of L mtDNAs form European-specific subclades, revealing that there was gene flow from Sub-Saharan Africa toward Europe as early as 11,000 years ago.[32]

Map (in the link) showing the distribution of Sub-Saharan mtDNA (shown in red) in Europe
Map is From Cerezo et al. 2012[32]
Universidad de Santiago de Compostela
Iberia (Spain & Portugal) having the highest amount and strongest concentration of Sub-Saharan mtDNA in Europe.

In Iberia the mean frequency of haplogroup L lineages reaches 3.83%; the frequency is higher in Portugal (5.83%) than in Spain (2.9% average), and without parallel in the rest of Europe. In both countries, frequencies vary widely between regions, but with increased frequencies observed for Madeira (insular Portugal), southern Portugal, Córdoba (southern Spain), Huelva (southern Spain), Canary Islands (insular Spain), Extremadura (western Spain) and Leon (western Spain).[3] In the Autonomous regions of Portugal (i.e. Madeira and the Azores), L haplogroups constituted about 13% of the lineages in Madeira, significantly more than in the Azores.[33] In the Canary Islands, frequencies have been reported at 6.6%.[33] Regarding Iberia, current debates are concerned with whether these lineages are associated with prehistoric migrations, the Islamic occupation of Iberia, or the slave trade. Pereira, Prata & Amorim (2000) suggested that African lineages in Iberia were predominantly the result of the Atlantic slave trade. González et al. (2003) revealed that most of the L lineages in Iberia matched Northwest African L lineages rather than contemporary Sub-Saharan L lineages. The authors suggest that this pattern indicates that most of the Sub-Saharan L lineages entered Iberia in prehistoric times rather than during the slave trade. According to Pereira et al. (2005), the Sub-Saharan lineages found in Iberia matched lineages from diverse regions in Africa. They suggest this pattern is more compatible with a recent arrival of these lineages after slave trading began in the 15th century. According to the study, alternative scenarios that invoke much older and demographically more significant introductions (González et al. (2003)) or that claim a substantial role of the Roman and/or Islamic periods on the introduction of Sub-Saharan lineages seem unlikely. Casas et al. (2006) extracted DNA from human remains that were exhumed from old burial sites in Al-Andalus, Spain. The remains date to between the 12th and 13th centuries. The frequency of Sub-Saharan lineages detected in the medieval samples was 14.6% and 8.3% in the present population of Priego de Cordoba. The authors suggest the Muslim occupation and prehistoric migrations before the Muslim occupation would have been the source of these lineages. The highest frequencies of Sub-Saharan lineages found so far in Europe were observed by Álvarez et al. (2010) in the comarca of Sayago (18.2%) which is, according to the authors, "comparable to that described for the South of Portugal".[10][34]

In Italy, haplogroup L lineages are present at lower frequencies than in Iberia—between —between 2% and 3%— and are detected only in certain regions: Latium, Volterra,[35] Basilicata, and Sicily.[36]

In eastern Europe, haplogroup L lineages are present at very low frequencies. Though a high diversity of African mtDNA lineages have been detected, few lineages have accumulated enough mutations in Europe to form monophyletic clusters. Malyarchuk & Czarny (2005) detected only two monophyletic clusters, L1b and L3b, in Russians, with an estimated age no greater than 6,500 years. Malyarchuk et al. (2008) identified African L1b, L2a, L3b, L3d and M1 clades in Slavic populations at low frequencies. L1b, L3b and L3d had matches with West African populations, indicating that these lineages probably entered Europe through Iberia. One lineage, L2a1a, appeared to be much older, indicating that it may have entered Europe in prehistoric times. This clade was possibly related to L2a1 clades identified in ten individuals of Ashkenazi heritage from France, Germany, Poland, Romania, and Russia. L2a lineages are widespread throughout Africa; as a result, the origins of this lineage are uncertain.[37]

Haplogroup M1 is also found in Europe at low frequencies. In a study by González et al. (2007), haplogroup M1 had a frequency of 0.3%. The origins of haplogroup M1 have yet to be conclusively established.

A 2015 study found that a prehistoric episode would be the main contributor to the sub-Saharan presence in Mediterranean Europe.[9]

Frequencies of haplogroup L lineages

Country Region Number tested Study %
Italy Countrywide 583 Brisighelli et al. (2012) 1.20%[38]
Italy Countrywide 865 Boattini et al. (2013) 0.00%[39]
Italy Countrywide 240 Babalini et al. (2005) 0.40%[40]
Italy Tuscany 322 Achilli et al. (2007) 1.86%
Italy Tuscany 49 Plaza et al. (2003) 2.00%
Italy Latium 138 Achilli et al. (2007) 2.90%
Italy Marche 813 Achilli et al. (2007) 0.98%
Italy Central Italy 83 Plaza et al. (2003) 1.20%
Italy Lombardy 177 Achilli et al. (2007) 0.00%
Italy Piedmont 169 Achilli et al. (2007) 0.00%
Italy Sardinia 258 Pardo et al. (2012) 0.40%
Italy Sardinia 73 Plaza et al. (2003) 2.80%
Italy Sardinia 85 Sanna et al. (2011) 0.00%
Italy Sardinia (Ogliastra) 475 Fraumene C et al. (2003) 0.00%[41]
Italy Sardinia 96 Morelli et al. (1999) 0.00%
Italy Campania (South Italy) 313 Achilli et al. (2007) 0.32%
Italy Basilicata (South Italy) 92 Ottoni et al. (2009) 2.20%
Italy Apulia & Calabria (South Italy) 226 Achilli et al. (2007) 0.00%
Italy Southern Italy 115 Sarno et al. (2014) 0.00%
Italy Southern Italy 37 Plaza et al. (2003) 8.10%
Italy Sicily 106 Cali et al. (2003) 0.94%
Italy Sicily 105 Achilli et al. (2007) 1.90%
Italy Sicily 169 Plaza et al. (2003) 0.60%
Italy Sicily 198 Sarno et al. (2014) 1.01%
Italy Sicily 465 Romano et al. (2013) 0.65%[42]
South Iberia Spain & Portugal 310 Casas et al. (2006) 7.40%
Spain Countrywide 312 Álvarez et al. (2007) 2.90%
Spain Central Spain 50 Plaza et al. (2003) 4.00%
Spain North-West Spain 216 Achilli et al. (2007) 3.70%
Spain Galicia 92 Pereira et al. (2005) 3.30%
Spain Zamora 214 Álvarez et al. (2010) 4.70%
Spain Sayago 33 Álvarez et al. (2010) 18.18%
Spain Cordoba 108 Casas et al. (2006) 8.30%
Spain Huelva 135 Hernandez et al. (2014) 5.70%
Spain Catalonia 101 Álvarez-Iglesias et al. (2009) 2.97%
Spain Balearic Islands 231 Picornell et al. (2005) 2.20%
Spain Canary Islands 300 Brehm et al. (2003) 6.60%
Portugal Countrywide 594 Achilli et al. (2007) 6.90%
Portugal Countrywide 1429 Barral-Arca et al. (2016) 6.16%
Portugal Countrywide 549 Pereira et al. (2005) 5.83%
Portugal North 100 Pereira et al. (2010) 5.00%
Portugal Center 82 Pereira et al. (2010) 9.70%
Portugal Center 82 Plaza et al. (2003) 6.10%
Portugal South 195 Brehm et al. (2003) 11.30%
Portugal South 303 Achilli et al. (2007) 10.80%
Portugal Coruche 160 Pereira et al. (2010) 8.70%
Portugal Pias 75 Pereira et al. (2010) 3.90%
Portugal Alcácer do Sal 50 Pereira et al. (2010) 22.00%
Portugal Azores 179 Brehm et al. (2003) 3.40%
Portugal Madeira 155 Brehm et al. (2003) 12.90%
Portugal Madeira 153 Fernandes et al. (2006) 12.40%
Greece Crete 202 Achilli et al. (2007) 0.99%
Cyprus Cyprus 91 Irwin et al. (2008) 3.30%

A similar study by Auton et al. (2009)—which also contains an admixture analysis chart but no cluster membership coefficients—shows little to no Sub-Saharan African influence in a wide array of European samples, i.e. Albanians, Austrians, Belgians, Bosnians, Bulgarians, Croatians, Cypriots, Czechs, Danes, Finns, Frenchmen, Germans, Greeks, Hungarians, Irish, Italians, Kosovars, Lithuanians, Latvians, Macedonians, Netherlanders, Norwegians, Poles, Portuguese, Romanians, Russians, Scots, Serbians, Slovaks, Slovenians, Spaniards, Swedes, Swiss (German, French and Italian), Ukrainians, subjects of the United Kingdom, and Yugoslavians.[43]

Haplogroup U6, to which a North African origin has been attributed (Rando et al. 1998), is largely distributed among Mozabites (28.2%) and Mauritanians (20%). In other northwest Africans, the frequency of U6 ranges from 4.2% in Tunisians to 8% in Moroccan Arabs (Plaza et al. 2003). In Europe, U6 is most common in Spain and Portugal.[44][45]

Frequencies of haplogroup U6 lineages

Country Region Number tested Study %
Italy Countrywide 583 Brisighelli et al. (2012) 0.8%
Italy Mainland 411 Plaza et al. (2003) 0.0%
Italy Countrywide 865 Boattini et al. (2013) 0.35%
Italy Sicily 169 Plaza et al. (2003) 0.6%
Italy Sicily 106 Maca-Meyer et al. (2003) 0.94%
Italy Lazio 52 Babalini et al. (2005) 5.8%[40]
Italy Abruzzo (Molise) 73 Babalini et al. (2005) 0%[40]
Italy Campania 48 Babalini et al. (2005) 0%[40]
Italy Volterra (Tuscany) 114 Achilli et al. (2007) 0.00%
Italy Murlo (Tuscany) 86 Achilli et al. (2007) 1.20%
Italy Casentino (Tuscany) 122 Achilli et al. (2007) 0.80%
Italy Sicily 105 Achilli et al. (2007) 0.95%
Italy Latium 138 Achilli et al. (2007) 0.00%
Italy Lombardy 177 Achilli et al. (2007) 0.00%
Italy Piedmont 169 Achilli et al. (2007) 0.00%
Italy Marche 813 Achilli et al. (2007) 0.25%
Italy Campania 313 Achilli et al. (2007) 1.28%
Italy Apulia-Calabria 226 Achilli et al. (2007) 1.33%
Italy Sardinia 370 Achilli et al. (2007) 0.27%
Spain Central Spain 50 Plaza et al. (2003) 2.0%
Spain Galicia 103 Plaza et al. (2003) 1.9%
Spain Galicia 135 Maca-Meyer et al. (2003) 2.2%
Spain Catalonia 118 Maca-Meyer et al. (2003) 1.6%
Spain Huelva 135 Hernandez et al. (2014) 8.8%
Spain Maragatos 49 Maca-Meyer et al. (2003) 8.1%
Spain Canary Islands 300 Brehm et al. (2003) 14.0%
Portugal Countrywide 54 Plaza et al. (2003) 5.6%
Portugal North Portugal 184 Maca-Meyer et al. (2003) 4.3%
Portugal Central Portugal 161 Brehm et al. (2003) 1.9%
Portugal Madeira 155 Brehm et al. (2003) 3.9%
Portugal Madeira 153 Fernandes et al. (2006) 3.3%
Iberia Spain & Portugal 887 Plaza et al. (2003) 1.8%

Admixture

  • A 2009 study by Auton et al. found a north–south cline of HapMap Yoruba haplotypes (YRI) in Europe. The study determined that southern and southwestern subpopulations had the highest proportion of YRI. This distribution is indicative of recurrent gene flow into Europe from the southwest and the Middle East. The authors suggest that the haplotype sharing between Europe and the YRI are suggestive of gene flow from Africa, albeit from West Africa and not necessarily North Africa.[43]
  • A 2007 study conducted at Penn State University found low levels of African admixture(2.8-10%) that were distributed along a north–south cline. The authors suggest that the distribution of this African admixture mirrors the distribution of haplogroup E3b-M35(E1b1b).[c][47]
  • A principal component analysis of data from the Human Genome Diversity Project by Reich et al. detected a west-to-east gradient of Bantu-related ancestry across Eurasia. The authors suggest that after the Out of Africa migration, there was most likely a later Bantu-related gene flow into Europe.[48]
  • The most recent study regarding African admixture in Iberian populations was conducted in April 2013 using genome-wide SNP data for over 2,000 individuals."Southwestern European populations averaged between 4% and 20% of their genomes assigned to a Northwest African ancestral cluster, whereas this value did not exceed 2% in southeastern European populations". However, contrary to past autosomal studies and to what is inferred from Y-chromosome and mitochondrial haplotype frequencies (see below), it does not detect significant levels of Sub-Saharan ancestry in any European population outside the Canary Islands.[8]
  • A panel of 52 SNPs was genotyped in 435 Italian individuals according to Sánchez et al.[49] in order to estimate the proportion of ancestry from a three-way differentiation: Sub-Saharan Africa, Europe, and Asia. The study indicated an autosomal basal proportion of Sub-Saharan ancestry that is higher (9.2%, on average) than other central or northern European populations (1.5%, on average). The amount of African ancestry in Italians is however more comparable to (but slightly higher than) the average in other Mediterranean countries (7.1%).[27] In northwestern Spain and in Portugal, Sub-Saharan autosomal ancestry is on average 7.1%.[27]
  • A 2015 study found that a prehistoric episode would be the main contributor to the sub-Saharan presence in Mediterranean Europe and Iberia:[9]
  • Measures of genetic distance between Europe and Sub-Saharan are generally smaller than genetic distances between Africa and other continental populations. Cavalli-Sforza states that the relatively short genetic distance is likely due to prehistoric admixture.[50]
  • A 2011 study by Moorjani et al. found that many southern Europeans have inherited 1%–3% Sub-Saharan ancestry (3.2% in Portugal, 2.9% in Sardinia, 2.7% in southern Italy, 2.4% in Spain and 1.1% in northern Italy), although the percentages were lower (ranging from 0.2% in Sardinia and northern Italy to 2.1% in Portugal) when reanalyzed with the 'STRUCTURE' statistical model. An average mixture date of around 55 generations/1100 years ago was given, "consistent with North African gene flow at the end of the Roman Empire and subsequent Arab migrations".[4]
  • An autosomal study in 2011 found an average Northwest African influence of about 17% in Canary Islanders, with a wide interindividual variation ranging from 0% to 96%. According to the authors, the substantial Northwest African ancestry found for Canary Islanders supports the idea that, despite the aggressive conquest by the Spanish in the 15th century and the subsequent immigration, genetic footprints of the first settlers of the Canary Islands persist in the current inhabitants. Paralleling mtDNA findings, the largest average Northwest African contribution was found for the samples from La Gomera.[51]

GM immunoglobulin allotypes

Further studies have shown that the presence of haplotype GM*1,17 23' 5* in southern Europe. This haplotype is considered a genetic marker of Sub-Saharan Africa, where it shows frequencies of about 80%.[52] Whereas, in non-Mediterranean European populations, that value is about 0.3%, in Spain the average figure for this African haplotype is nearly eight times greater (though still at a low level) at 2.4%, and it shows a peak at 4.5% in Galicia.[53] Values of around 4% have also been found in Huelva and in the Aran valley in the Pyrenees.[54] According to Calderón et al. 2007, although some researchers have associated African traces in Iberia to Islamic conquest, the presence of GM*1,17 23' 5* haplotype in Iberia may in fact be due to more ancient processes as well as more recent ones through the introduction of genes from black slaves from Africa.[53]
In Sicily the North African haplotype Gm 5*;1;17; ranges from 1.56% at Valledolmo to 5.5% at Alia.[55] The hypothesis is that the presence of this haplotype suggests past contacts with people from North Africa. The introduction of African markers could be due to the Phoenician colonization at the end of the second millennium B.C. or to the more recent Arab conquest (8th–9th centuries A.D.).

Paleoanthropology

The migration of farmers from the Middle East into Europe is believed to have significantly influenced the genetic profile of present-day Europeans. Some recent studies have focused on corroborating current genetic data with the archeological evidence from Europe, the Middle East, and Africa.[24] The Natufian culture, which existed about 12,000 years ago, has been the subject of various archeological investigations, as it is generally believed to be the source of the European and North African Neolithic.

According to a hypothesis stated by Bar-Yosef (1987), the Natufian culture emerged from the mixing of two Stone Age cultures: (1) the Kebaran, a culture indigenous to the Levant, and (2) the Mushabian, a culture introduced into the Levant from North Africa†. It is suggested that the Mushabian culture originated in Africa, given that archeological sites with Mushabian industries in the Nile Valley predate those in the Levant†. The Mushabians would have then moved into the Sinai from the Nile Delta bringing with them their technologies†. Bar-Yosef (1987) states: "the population overflow from Northeast Africa played a definite role in the establishment of the Natufian adaptation, which in turn led to the emergence of agriculture as a new subsistence system".

A study by Brace et al. (2006) analysed human remains from the Natufian culture. According to the study, there is evidence of Sub-Saharan influences in the Natufian samples. They argue that these influences would have been diluted by the interbreeding of the Neolithic farmers from the Near East with the indigenous foragers in Europe. Ricaut & Waelkens (2008) associate the Sub-Saharan influences detected in the Natufian samples with the migration of E1b1b lineages from Northeast Africa to the Levant and then into Europe.

According to the most recent ancient DNA analyses conducted by Lazaridis et al. (2016) on Natufian skeletal remains from present-day northern Israel, the Natufians in fact shared no evident genetic affinity to sub-Saharan Africans. The authors also state that they were unable to test for affinity in the Natufians to early North African populations using present-day North Africans as a reference because present-day North Africans owe most of their ancestry to back-migration from Eurasia.[21][56] The Natufians carried the Y-DNA (paternal) haplogroups E1b1b1b2(xE1b1b1b2a,E1b1b1b2b) (2/5; 40%), CT (2/5; 40%), and E1b1(xE1b1a1,E1b1b1b1) (1/5; 20%).[21][57] In terms of autosomal DNA, these Natufians carried around 50% of the Basal Eurasian (BE) and 50% of Western Eurasian Unknown Hunter Gather (UHG) components. However, they were slightly distinct from the northern Anatolian populations that contributed to the peopling of Europe, who had higher Western Hunter-Gatherer (WHG) inferred ancestry. Natufians were strongly genetically differentiated[58] from Neolithic Iranian farmers from the Zagros Mountains, who were a mix of Basal Eurasians (up to 62%) and Ancient North Eurasians (ANE). This might suggest that different strains of Basal Eurasians contributed to Natufians and Zagros farmers,[59][60][61] as both Natufians and Zagros farmers descended from different populations of local hunter gatherers. Mating between Natufians, other Neolithic Levantines, Caucasus Hunter Gatherers (CHG), Anatolian and Iranian farmers is believed to have decreased genetic variability among later populations in the Middle East. The scientists suggest that the Levantine early farmers may have spread southward into East Africa, bringing along Western Eurasian and Basal Eurasian ancestral components separate from that which would arrive later in North Africa.

† The Mushabian industry is now known to have originated in the Levant from the previous lithic industries of the region of Lake Lisan.[62] The Mushabian industry was originally thought to have originated in Africa because the microburin technique was not yet known to be much older in the eastern Levant.[63] Currently there is no known industry to connect with the African migration that occurred 14,700 years ago,[1] but it no doubt caused a population expansion in the Negev and Sinai which would not have accommodated an increase in population with the meager resources of a steppe/desert climate.[14] Since all of the known cultures in the Levant at the time of the migration originated in the Levant and an archaeological culture cannot be associated with it, there must have been assimilation into a Levantine culture at the onset, most likely the Ramonian which was present in the Sinai 14,700 years ago.[64]

See also

Notes

  1. ^ Recently, it has been proposed that E3b originated in eastern Africa and expanded into the Near East and northern Africa at the end of the Pleistocene. E3b lineages would have then been introduced from the Near East into southern Europe by migrant farmers, during the Neolithic expansion.[1]
  2. ^ A Mesolithic population carrying Group III lineages with the M35/M215 mutation expanded northwards from sub-Saharan to north Africa and the Levant. The Levantine population of farmers that dispersed into Europe during and after the Neolithic carried these African Group III M35/M215 lineages, together with a cluster of Group VI lineages characterized by M172 and M201 mutations.[23]
  3. ^ We observed patterns of apportionment similar to those described previously using sex and autosomal markers, such as European admixture for African Americans (14.3%) and Mexicans (43.2%), European (65.5%) and East Asian affiliation (27%) for South Asians and low levels of African admixture (2.8–10.8%) mirroring the distribution of Y E3b haplogroups among various Eurasian populations.[46]

Footnotes

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References

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