Fri . 20 Jul 2020
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African admixture in Europe

african admixture in europe
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 Africa to the Near East during the later Pleistocene, and from there to Europe with the Neolithic Revolution More recent, direct African admixture is associated with the Muslim conquests of the early medieval period including Berber admixture from North Africa, and is primarily concentrated in populations in the Iberian peninsula

Contents

  • 1 Neolithic
  • 2 Historical period
  • 3 Assessing African genetic contributions in non-Africans
    • 31 Defining African admixture
  • 4 Y-DNA
  • 5 mtDNA
    • 51 Frequencies of haplogroup L lineages
    • 52 Frequencies of Haplogroup U6 lineages
  • 6 Admixture
    • 61 GM immunoglobulin allotypes
    • 62 Sickle cell trait
  • 7 Paleoanthropology
  • 8 See also
  • 9 Notes
  • 10 Footnotes
  • 11 References

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 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

It has been suggested that the first Middle Eastern farmers reflected North African influences There have been suggestions that some genetic lineages found in the Middle East arrived there during this period 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

Historical period

In historical times, there has been a period of North African influence in southern Europe, especially Iberia and 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

Assessing African genetic contributions in non-Africans

The evolutionary forces that contribute to patterns of human genetic variation include new mutations, natural selection, sexual selection, genetic drift, population bottlenecks, founder effects, isolation by distance, genetic admixture and barriers to gene flow The most influential factors affecting human genetic variation are founder effects and isolation by distance

Founder effect occurs when a population is established by a small number of individuals who have departed from a much larger population Several generations after the population has expanded, individuals will still possess the limited gene pool of the founders Therefore, founder effects result in a loss of genetic diversity Genetic evidence suggests that the Out of Africa migration involved only small numbers of individuals The African migrants carried a small subset of the prehistoric African genetic diversity, resulting in a founder effect on the non-African population As humans spread across the globe populating Europe, Asia, Oceania, and the Americas, there were several founder effects As a result of these serial founder effects, genetic diversity tends to decrease with distance from Africa

The other major factor contributing to patterns of human genetic variation is "isolation by distance" According to this model, populations that live near one another are more likely to exchange mates than populations that live farther apart As a result, populations that live near one another are genetically more similar than populations that live far apart

Percentage genetic distances among major continents based on 120 classical polymorphisms
Africa Oceania East Asia Europe
Oceania 247
East Asia 206 10
Europe 166 135 97
America 226 146 89 95

Genetic distance is a measure used to compare the genetic relationship between populations It is based on the principle that populations that share similar frequencies of a trait are more closely related than populations that have different frequencies of a trait The genetic distance between two populations increases linearly with the geographic distance between them, due to isolation by distance and serial founder effects Genetic admixture increases the genetic diversity of a population When admixture occurs between populations, the genetic distance between the two populations is reduced

Cavalli-Sforza 1997 applied genetic distance measures to various populations around the world to infer phylogenetic relationships see Table on the right All non-African populations are more closely related to one another ie short genetic distance than they are to African populations This is consistent with a founder effect on the non-African population in that only a few individuals participated in the initial Out of Africa migration The largest genetic distances observed are between Africa and Oceania and between Africa and the Americas This is consistent with the isolation by distance and serial founder effects

Cavalli-Sforza 1997 suggests that the genetic distance between Sub-Saharan Africa and Europe is anomalously lower than it would be if the two populations had been evolving independently The study suggests that the lower genetic distance between Europe and Africa can be explained by genetic admixture

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

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

Other lineages that are now found in Africa and Europe may have a common origin in Asia eg 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 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 Such bidirectional migrations between Africa and Eurasia complicate the task of defining admixture

Y-DNA

One proposed example of Holocene gene flow from Africa to Europe, via the Middle East, is thought to be E1b1b1, which is thought to have emerged about 22,000 years ago in East Africa, and branches of it are thought to have migrated to the Middle East by 11,000 years ago during the late Pleistocene or early Neolithic periods

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

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 72% The timing of this movement has been given widely varying estimates

A major expansion of peoples throughout Sub-Saharan Africa occurred after the introduction of agriculture 5,000 years ago During the Bantu Expansion, people carrying haplogroup E not including E1b1b lineages dispersed across much of Sub-Saharan Africa from their homeland near the present-day border between Nigeria and Cameroon The haplogroup most often associated with this expansion is E1b1a, which is present in up to 48% of the African male gene pool The presence of E1b1a lineages outside Africa can typically be associated with events that occurred after the Bantu Expansion, such as the Moorish occupation of Iberia or the trade in African slaves In much of Europe, frequencies of E lineages which are not E1b1b 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, 29% in Istanbul, and 43% 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%, among Italians in Calabria 1/80=13%, and among Albanians in Calabria 2/68=29% 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

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 eastern 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 and reaches a frequency of 8% in northern and central Italy and slightly higher, 11%, in the south of that country
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

North African Y-DNA E-M81 was found at a total of 411% among "pasiegos" from Cantabria, Spain That is the highest frequency observed in Europe to date

In Sardinia, Sub-Saharan Y-DNA lineages A1b1b2b and E1a1 were found at a total of 10% A1b1b2b 05% / E1a1 05%

In Majorcans, Sub-Saharan Y-DNA lineage E-V38 was found at a total of 32% 2/62

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

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 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

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

In Iberia the mean frequency of haplogroup L lineages reaches 383%; the frequency is higher in Portugal 583% than in Spain 29% 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 In the Autonomous regions of Portugal ie Madeira and the Azores, L haplogroups constituted about 13% of the lineages in Madeira, significantly more than in the Azores In the Canary Islands, frequencies have been reported at 66% 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 et al 2000 suggested that African lineages in Iberia were predominantly the result of the Atlantic slave tarade 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 146% and 83% 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 182% which is, according to the authors, "comparable to that described for the South of Portugal" and by Pereira et al 2010 in Alcácer do Sal 22%

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, Basilicata, and Sicily

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 et al 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

Haplogroup M1 is also found in Europe at low frequencies In a study by González et al 2007, haplogroup M1 had a frequency of 03% 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

Frequencies of haplogroup L lineages

Country Region' Number tested Study %
Germany 333 Pliss et al 2005 120%
Italy Countrywide 583 Brisighelli et al 2012 120%
Italy Countrywide 865 Boattini et al 2013 000%
Italy Countrywide 240 Babalini et al 2005 040%
Italy Tuscany 322 Achilli et al 2007 186%
Italy Tuscany 49 Plaza et al 2003 200%
Italy Latium 138 Achilli et al 2007 290%
Italy Marche 813 Achilli et al 2007 098%
Italy Central Italy 83 Plaza et al 2003 120%
Italy Lombardy 177 Achilli et al 2007 000%
Italy Piedmont 169 Achilli et al 2007 000%
Italy Sardinia 258 Pardo et al 2012 040%
Italy Sardinia 73 Plaza et al 2003 280%
Italy Sardinia 85 Sanna et al 2011 000%
Italy Sardinia Ogliastra 475 Fraumene C et al 2003 000%
Italy Sardinia 96 Morelli et al 1999 000%
Italy Campania South Italy 313 Achilli et al 2007 032%
Italy Basilicata South Italy 92 Ottoni et al 2009 220%
Italy Apulia & Calabria South Italy 226 Achilli et al 2007 000%
Italy Southern Italy 115 Sarno et al 2014 000%
Italy Southern Italy 37 Plaza et al 2003 810%
Italy Sicily 106 Cali et al 2003 094%
Italy Sicily 105 Achilli et al 2007 190%
Italy Sicily 169 Plaza et al 2003 060%
Italy Sicily 198 Sarno et al 2014 101%
Italy Sicily 465 Romano et al 2013 065%
South Iberia Spain & Portugal 310 Casas et al 2006 740%
Spain Countrywide 312 Álvarez et al 2007 290%
Spain Central Spain 50 Plaza et al 2003 400%
Spain North-West Spain 216 Achilli et al 2007 370%
Spain Galicia 92 Pereira et al 2005 330%
Spain Zamora 214 Álvarez et al 2010 470%
Spain Sayago 33 Álvarez et al 2010 1818%
Spain Cordoba 108 Casas et al 2006 830%
Spain Huelva 135 Hernandez et al 2014 570%
Spain Catalonia 101 Álvarez-Iglesias et al 2009 297%
Spain Balearic Islands 231 Picornell et al 2005 220%
Spain Canary Islands 300 Brehm et al 2003 660%
Portugal Countrywide 594 Achilli et al 2007 690%
Portugal Countrywide 1429 Barral-Arca et al 2016 616%
Portugal Countrywide 549 Pereira et al 2005 583%
Portugal North 100 Pereira et al 2010 500%
Portugal Center 82 Pereira et al 2010 970%
Portugal Center 82 Plaza et al 2003 610%
Portugal South 195 Brehm et al 2003 1130%
Portugal South 303 Achilli et al 2007 1080%
Portugal Coruche 160 Pereira et al 2010 870%
Portugal Pias 75 Pereira et al 2010 390%
Portugal Alcácer do Sal 50 Pereira et al 2010 2200%
Portugal Azores 179 Brehm et al 2003 340%
Portugal Madeira 155 Brehm et al 2003 1290%
Portugal Madeira 153 Fernandes et al 2006 1240%
Greece Crete 202 Achilli et al 2007 099%
Cyprus Cyprus 91 Irwin et al 2008 330%

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, ie Albanians, Austrians, Belgians, Bosnians, Bulgarians, Croatians, Cypriots, Czechs, Danes, Finns, Frenchmen, Germans, Greeks, Hungarians, Irish, Italians, Kosovars, Latvians, Macedonians, Netherlanders, Norwegians, Poles, Portuguese, Romanians, Russians, Scots, Serbians, Slovakians, Slovenians, Spaniards, Swedes, Swiss German, French and Italian, Ukrainians, subjects of the United Kingdom, and Yugoslavians

Haplogroup U6, to which a North African origin has been attributed Rando et al 1998, is largely distributed among Mozabites 282% and Mauritanians 20% In other Northwest Africans, the frequency of U6 ranges from 42% in Tunisians to 8% in Moroccan Arabs Plaza et al 2003 In Europe, U6 is most common in Spain and Portugal

Frequencies of Haplogroup U6 lineages

Country Region Number tested Study %
Italy Countrywide 583 Brisighelli et al 2012 08%
Italy Mainland 411 Plaza et al 2003 00%
Italy Countrywide 865 Boattini et al 2013 035%
Italy Sicily 169 Plaza et al 2003 06%
Italy Sicily 106 Maca-Meyer et al 2003 094%
Italy Lazio 52 Babalini et al 2005 58%
Italy Abruzzo Molise 73 Babalini et al 2005 0%
Italy Campania 48 Babalini et al 2005 0%
Italy Volterra Tuscany 114 Achilli et al 2007 000%
Italy Murlo Tuscany 86 Achilli et al 2007 120%
Italy Casentino Tuscany 122 Achilli et al 2007 080%
Italy Sicily 105 Achilli et al 2007 095%
Italy Latium 138 Achilli et al 2007 000%
Italy Lombardy 177 Achilli et al 2007 000%
Italy Piedmont 169 Achilli et al 2007 000%
Italy Marche 813 Achilli et al 2007 025%
Italy Campania 313 Achilli et al 2007 128%
Italy Apulia-Calabria 226 Achilli et al 2007 133%
Italy Sardinia 370 Achilli et al 2007 027%
Spain Central Spain 50 Plaza et al 2003 20%
Spain Galicia 103 Plaza et al 2003 19%
Spain Galicia 135 Maca-Meyer et al 2003 22%
Spain Catalonia 118 Maca-Meyer et al 2003 16%
Spain Huelva 135 Hernandez et al 2014 88%
Spain Maragatos 49 Maca-Meyer et al 2003 81%
Spain Canary Islands 300 Brehm et al 2003 140%
Portugal Countrywide 54 Plaza et al 2003 56%
Portugal North Portugal 184 Maca-Meyer et al 2003 43%
Portugal Central Portugal 161 Brehm et al 2003 19%
Portugal Madeira 155 Brehm et al 2003 39%
Portugal Madeira 153 Fernandes et al 2006 33%
Iberia Spain & Portugal 887 Plaza et al 2003 18%

Admixture

  • A 2011 study by Moorjani et al found that almost all southern Europeans have inherited 1%–3% Sub-Saharan ancestry 32% in Portugal, 29% in Sardinia, 27% in southern Italy, 24% in Spain and 11% in northern Italy with an average mixture date of around 55 generations ago, "consistent with North African gene flow at the end of the Roman Empire and subsequent Arab migrations"

The finding of African ancestry in Southern Europe dating to ~55 generations ago, or ~1,600 years ago assuming 29 years per generation, needs to be placed in historical context The historical record documents multiple interactions of African and European populations over this period One potential opportunity for African gene flow was during the period of Roman occupation of North Africa that lasted until the early 5th century AD, and indeed tomb inscriptions and literary references suggest that trade relations continued even after that time North Africa was also a supplier of goods and products such as wine and olive oil to Italy, Spain and Gaul from 200–600 AD, and Morocco was a major manufacturer of the processed fish sauce condiment, garum, which was imported by Romans In addition, there was slave trading across the western Sahara during Roman times Another potential source of some of the African ancestry, especially in Spain and Portugal, is the invasion of Iberia by Moorish armies after 711 AD If the Moors already had some African ancestry when they arrived in Southern Europe, and then admixed with Iberians, we would expect the admixture date to be older than the date of the invasion, as we observe

— Moorjani et al 2011
  • 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
  • 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
  • A 2007 study conducted at Penn State University found low levels of African admixture28-10% that were distributed along a north/south cline The authors suggest that the distribution of this African admixture mirrors the distribution of haplogroup E3b-M35E1b1b
  • 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
  • The most recent study regarding African admixture in Iberian populations was conducted in April 2013 using genome-wide SNP data for over 2000 individuals This study concluded that Spain and Portugal hold significantly higher levels of North African heritage than the rest of the European continent Estimates of shared ancestry averaged between 4% and 20%, whereas these did not exceed 2% in other western or southern 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
  • A panel of 52 SNPs was genotyped in 435 Italian individuals according to Sánchez et al 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 92%, on average than other central or northern European populations 15%, on average The amount of African ancestry in Italians is however more comparable to but slightly higher than the average in other Mediterranean countries 71%
In northwestern Spain and in Portugal, Sub-Saharan autosomal ancestry is on average 71%
  • A 2009 autosomal study by Moorjani et al that used between 500 thousand and 15 million SNPs estimated that the proportion of Sub-Saharan African ancestry is 24% in Spain, and 19% in Greece According to the authors, this is consistent in the case of Spain, with the historically known movement of individuals of North African ancestry into Iberia, although it is possible that this estimate also reflects a wider range of mixture times Another study by the same author in 2011 that analyzed genome-wide polymorphism data from about 40 West Eurasian groups showed that "almost all Southern Europeans have inherited 1%–3% African ancestry with an average mixture date of around 55 generations ago, consistent with North African gene flow at the end of the Roman Empire and subsequent Arab invasion" According to the authors, application of f4 Ancestry Estimation, a method which produces accurate estimates of ancestry proportions, even in the absence of data from the true ancestral populations, suggests that the "highest proportion of African ancestry in Europe is in Iberia Portugal 32±03% and Spain 24±03%, consistent with inferences based on mitochondrial DNA and Y chromosomes and the observation by Auton et al that within Europe, the Southwestern Europeans have the highest haplotype-sharing with Africans" The authors found the following Sub-Saharan African admixture proportions in Europe using f4 Ancestry Estimation
  • A 2015 study found that a prehistoric episode would be the main contributor to the sub-Saharan presence in Mediterranean Europe and Iberia:
Population Sub-Saharan African >1%°
African American 794%
Portugal 32%
Spain 24%
S Italy 27%
Greece 19%
Sardinia 29%
N Italy 11%
Sephardic Jews 48%
Ashkenazi Jews 32%
From the same study, estimates of Sub-Saharan African admixture proportions in Europe:
Population Sub-Saharan African >01%°
African American 772%
Portugal 21%
Spain 11%
S Italy 17%
N Italy 02%
Sardinia 02%
Sephardic Jews 43%
Ashkenazi Jews 26%
  • The Spanish population may harbor some African-related admixture representing a fourth wave of migration into Europe, but affecting Spain much more than the other groups The Spanish population shows an African admixture of 148% 126% Mozabite and 22% Mbuti/Yoruba, confirming that gene flow from Sub-Saharan or North African populations has occurred in the Spanish sample
  • 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
Canary Islands and mainland Iberians N Average
NW African
ancestry
La Gomera 7 4250%
Fuerteventura 10 2160%
La Palma 7 2100%
El Hierro 7 1980%
Lanzarote 13 1640%
Tenerife 30 1430%
Gran Canaria 30 1240%
Total Canary Islanders 104 1740%
Total Canary Islanders from the Spanish DNA bank 15 1260%
Mainland Iberians 77 500%

GM immunoglobulin allotypes

Further studies have shown that the presence of haplotype GM1,17 23' 5 in southern Europe This haplotype is considered a genetic marker of Sub-Saharan Africa, where it shows frequencies of about 80% Whereas, in non-Mediterranean European populations, that value is about 03%, in Spain the average figure for this African haplotype is nearly eight times greater though still at a low level at 24%, and it shows a peak at 45% in Galicia Values of around 4% have also been found in Huelva and in the Aran valley in the Pyrenees According to Calderón et al 2007, although some researchers have associated African traces in Iberia to Islamic conquest, the presence of GM1,17 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
In Sicily the African haplotype Gm 5;1;17; ranges from 156% at Valledolmo to 55% at Alia 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 BC or to the more recent Arab conquest 8th–9th centuries AD

Sickle cell trait

Sickle cell genes of African origin have been detected in Europe, mostly in the Mediterranean region The sickle cell trait is associated with resistance to malaria Individuals with one copy of the sickle cell gene, heterozygotes, have higher resistance to malaria than individuals with no sickle cell genes In regions affected by malaria, the fertility of sickle cell heterozygotes will be higher than average Individuals with two copies of the sickle cell gene, homozygotes, will be affected by sickle cell disease and historically have had lower than average fertility The sickle cell trait is thus an example of heterozygote advantage which is subject to balancing selection When the sickle cell trait is introduced into a region affected by malaria, balancing selection will, on one hand, act to increase the frequency of the trait to counter malaria On the other hand, if the frequency of the trait in the population becomes high enough so that homozygotes with sickle cell disease become common, balancing selection will act to limit the spread of the trait Therefore, in regions affected by malaria, the sickle cell trait is maintained at intermediate frequencies relative to the incidence of malaria

In Africa, malaria is believed to be one of the most important factors that contributed to restricting population growth in prehistoric times Sickle cell mutations are believed to have occurred independently at least five times Four variants are of African origin and one of Indian/Arabian origin The African variants are referred to as Cameroon, Senegal, Benin and Bantu The emergence of the sickle cell trait would have contributed to population expansion, with the emergence of farming into tropical regions where malaria was endemic

Archeological and historical evidence suggests that malaria has been endemic in the Mediterranean regions of Europe in historical times In eastern Mediterranean regions—such as Italy, Greece, Albania, and Turkey—the Benin haplotype is the most frequent sickle cell variant The Benin haplotype is also the most frequent variant in the Middle East and has been observed in Syrians, Palestinians, Italians from Calabria and Sicily, Israeli Arabs, Israeli Jews and western Saudi Arabians This suggests that the Benin haplotype may have expanded from West Africa into North Africa and then into the Middle East and Europe The spread of the Benin haplotype to the Mediterranean region has been associated with various events, including Late Stone Age expansions from West Africa into North Africa, the trans-Saharan trade, and the Arab Slave Trade The occurrence of sickle cell trait is particularly high in Sicily, where frequencies of 13% have been reported Portugal is the only region in Europe where the Senegal and Bantu haplotypes are frequent These may be associated with Portuguese naval exploits, including the Atlantic Slave Trade and the colonization of various African countries

In Europe, the highest prevalence of the disease has been observed in France as a result of population growth in African-Caribbean regions of overseas France, and now immigration essentially from North and Sub-Saharan Africa to mainland France SCD has become the most common genetic disease in this country In 2010, 315% of all newborns in mainland France had parents originated from a region defined "at risk" mainly Africa and Overseas departments and territories of France and were screened for SCD The Paris metropolitan district Île-de-France is the region that accounts for the largest number of at-risk individuals Indeed, 60% of all newborns in this area in 2010 had parents originated from a region defined as "at-risk" and were screened for SCD The second largest number of at-risk individuals is in Provence-Alpes-Côte d'Azur, at nearly 432%, and the lowest number is in Brittany, at 55%

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 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 2005 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 et al 2008 associate the Sub-Saharan influences detected in the Natufian samples with the migration of E1b1b lineages from East Africa to the Levant and then into Europe

The Mushabian industry is now known to have originated in the Levant from the previous lithic industries of the region of Lake Lisan 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 Currently there is no known industry to connect with the African migration that occurred 14,700 years ago, 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 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

See also

  • Molecular Anthropology portal
  • Evolutionary biology portal
  • Europe portal
  • African immigration to Europe contemporary history
  • Archaeogenetics of the Near East
  • Genetic history of North Africa
  • Genetic history of Europe
  • Arab slave trade
  • Barbary slave trade
  • Abkhazians of African descent

Notes

  1. ^ 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:101086/386294 PMC 1181964  PMID 15042509 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 
  2. ^ Underhill PA, Passarino G, Lin AA, Shen P, Mirazón Lahr M, Foley RA, Oefner PJ, Cavalli-Sforza LL January 2001 "The phylogeography of Y chromosome binary haplotypes and the origins of modern human populations" Annals of Human Genetics 65 Pt 1: 43–62 doi:101046/j1469-180920016510043x PMID 11415522 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 
  3. ^ Halder I, Shriver M, Thomas M, Fernandez JR, Frudakis T May 2008 "A panel of ancestry informative markers for estimating individual biogeographical ancestry and admixture from four continents: utility and applications" Human Mutation 29 5: 648–58 doi:101002/humu20695 PMID 18286470 We observed patterns of apportionment similar to those described previously using sex and autosomal markers, such as European admixture for African Americans 143% and Mexicans 432%, European 655% and East Asian affiliation 27% for South Asians and low levels of African admixture 28–108% mirroring the distribution of Y E3b haplogroups among various Eurasian populations 

Footnotes

  1. ^ a b Cruciani et al 2004
  2. ^ a b c Malyarchuk et al 2005
  3. ^ http://wwwpnasorg/content/110/29/11791/F2largejpg
  4. ^ http://wwwpnasorg/content/110/29/11791full
  5. ^ Botigué LR, Henn BM, Gravel S, Maples BK, Gignoux CR, Corona E, Atzmon G, Burns E, Ostrer H, Flores C, Bertranpetit J, Comas D, Bustamante CD 2013 "Gene flow from North Africa contributes to differential human genetic diversity in southern Europe" Proc Natl Acad Sci USA 110 29: 11791–6 Bibcode:2013PNAS11011791B doi:101073/pnas1306223110 PMC 3718088  PMID 23733930 CS1 maint: Multiple names: authors list link
  6. ^ Pereira L, Cunha C, Alves C, Amorim A April 2005 "African female heritage in Iberia: a reassessment of mtDNA lineage distribution in present times" Hum Biol 77 2: 213–29 doi:101353/hub20050041 PMID 16201138 CS1 maint: Multiple names: authors list link
  7. ^ Hernández, Candela L; Soares, Pedro; Dugoujon, Jean M; Novelletto, Andrea; Rodríguez, Juan N; Rito, Teresa; et al 2015 "Early Holocenic and Historic mtDNA African Signatures in the Iberian Peninsula: The Andalusian Region as a Paradigm" PLOS One 10 10: e0139784 Bibcode:2015PLoSO1039784H doi:101371/journalpone0139784 PMC 4624789  PMID 26509580 
  8. ^ Alvarez L, Santos C, Ramos A, Pratdesaba R, Francalacci P, Aluja MP August 2010 "Mitochondrial DNA patterns in the Iberian Northern plateau: population dynamics and substructure of the Zamora province" Am J Phys Anthropol 142 4: 531–9 doi:101002/ajpa21252 PMID 20127843 
  9. ^ Alvarez L, Santos C, Ramos A, Pratdesaba R, Francalacci P, Aluja MP August 2010 "Mitochondrial DNA patterns in the Iberian Northern plateau: population dynamics and substructure of the Zamora province" Am J Phys Anthropol 142 4: 531–9 doi:101002/ajpa21252 PMID 20127843 As regards sub-Saharan Hgs L1b, L2b, and L3b, the high frequency found in the southern regions of Zamora, 182% in Sayago and 81% in Bajo Duero, is comparable to that described for the South of Portugal 
  10. ^ https://hrcaksrcehr/file/266318
  11. ^ Lazaridis et al, "Ancient human genomes suggest three ancestral populations for present-day Europeans", Nature, 5137518, 18 September 2014, 409–413, doi: 101038/nature13673 Supplemental Information
  12. ^ L Kovacevic et al, "Standing at the Gateway to Europe - The Genetic Structure of Western Balkan Populations Based on Autosomal and Haploid Markers", PLOS One, 22 August 2014, doi:101371/journalpone0105090
  13. ^ a b Botigué LR, Henn BM, Gravel S, Maples BK, Gignoux CR, Corona E, Atzmon G, Burns E, Ostrer H, Flores C, Bertranpetit J, Comas D, Bustamante CD July 2013 "Gene flow from North Africa contributes to differential human genetic diversity in southern Europe" Proceedings of the National Academy of Sciences of the United States of America 110 29: 11791–6 Bibcode:2013PNAS11011791B doi:101073/pnas1306223110 PMC 3718088  PMID 23733930 
  14. ^ Pereira L, Cunha C, Alves C, Amorim A April 2005 "African female heritage in Iberia: a reassessment of mtDNA lineage distribution in present times" Human Biology 77 2: 213–29 doi:101353/hub20050041 PMID 16201138 
  15. ^ Hernández CL, Soares P, Dugoujon JM, Novelletto A, Rodríguez JN, Rito T, Oliveira M, Melhaoui M, Baali A, Pereira L, Calderón R 2015 "Early Holocenic and Historic mtDNA African Signatures in the Iberian Peninsula: The Andalusian Region as a Paradigm" PLOS One 10 10: e0139784 Bibcode:2015PLoSO1039784H doi:101371/journalpone0139784 PMC 4624789  PMID 26509580  Alvarez L, Santos C, Ramos A, Pratdesaba R, Francalacci P, Aluja MP August 2010 "Mitochondrial DNA patterns in the Iberian Northern plateau: population dynamics and substructure of the Zamora province" American Journal of Physical Anthropology 142 4: 531–9 doi:101002/ajpa21252 PMID 20127843 As regards sub-Saharan Hgs L1b, L2b, and L3b, the high frequency found in the southern regions of Zamora, 182% in Sayago and 81% in Bajo Duero, is comparable to that described for the South of Portugal  https://hrcaksrcehr/file/266318
  16. ^ Brace et al 2005
  17. ^ Cavalli-Sforza 1993
  18. ^ Bar-Yosef O 1987 Pleistocene connections between Africa and Southwest Asia: an archaeological perspective The African Archaeological Review; Chapter 5, pg 29-38
  19. ^ a b Underhill PA, Kivisild T 2007 "Use of y chromosome and mitochondrial DNA population structure in tracing human migrations" Annual Review of Genetics 41 1: 539–64 doi:101146/annurevgenet41110306130407 PMID 18076332 
  20. ^ a b c Ramachandran S, Deshpande O, Roseman CC, Rosenberg NA, Feldman MW, Cavalli-Sforza LL November 2005 "Support from the relationship of genetic and geographic distance in human populations for a serial founder effect originating in Africa" Proceedings of the National Academy of Sciences of the United States of America 102 44: 15942–7 Bibcode:2005PNAS10215942R doi:101073/pnas0507611102 PMC 1276087  PMID 16243969 
  21. ^ a b c Handley LJ, Manica A, Goudet J, Balloux F September 2007 "Going the distance: human population genetics in a clinal world" Trends in Genetics 23 9: 432–9 doi:101016/jtig200707002 PMID 17655965 
  22. ^ Fregel; et al 2017 "Neolithization of North Africa involved the migration of people from both the Levant and Europe" bioRxiv 191569  
  23. ^ Rando JC, Cabrera VM, Larruga JM, Hernández M, González AM, Pinto F, Bandelt HJ September 1999 "Phylogeographic patterns of mtDNA reflecting the colonization of the Canary Islands" Annals of Human Genetics 63 Pt 5: 413–28 doi:101046/j1469-180919996350413x PMID 10735583 
  24. ^ González AM, Larruga JM, Abu-Amero KK, Shi Y, Pestano J, Cabrera VM July 2007 "Mitochondrial lineage M1 traces an early human backflow to Africa" BMC Genomics 8 1: 223 doi:101186/1471-2164-8-223 PMC 1945034  PMID 17620140 
  25. ^ Ennaffaa et al 2009
  26. ^ a b Semino et al 2004
  27. ^ Underhill PA, Passarino G, Lin AA, Shen P, Mirazón Lahr M, Foley RA, Oefner PJ, Cavalli-Sforza LL January 2001 "The phylogeography of Y chromosome binary haplotypes and the origins of modern human populations" PDF Annals of Human Genetics 65 Pt 1: 43–62 doi:101046/j1469-180920016510043x PMID 11415522 Archived from the original PDF on 2009-08-15 
  28. ^ a b Lancaster 2009
  29. ^ Gonçalves et al 2005
  30. ^ King TE, Parkin EJ, Swinfield G, Cruciani F, Scozzari R, Rosa A, Lim SK, Xue Y, Tyler-Smith C, Jobling MA March 2007 "Africans in Yorkshire The deepest-rooting clade of the Y phylogeny within an English genealogy" European Journal of Human Genetics 15 3: 288–93 doi:101038/sjejhg5201771 PMC 2590664  PMID 17245408 
  31. ^ a b c d e Brisighelli F, Álvarez-Iglesias V, Fondevila M, Blanco-Verea A, Carracedo A, Pascali VL, Capelli C, Salas A 2012 "Uniparental markers of contemporary Italian population reveals details on its pre-Roman heritage" PLOS One 7 12: e50794 Bibcode:2012PLoSO750794B doi:101371/journalpone0050794 PMC 3519480  PMID 23251386 
  32. ^ a b 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:101086/386294 PMC 1181964  PMID 15042509 
  33. ^ Francalacci P, Morelli L, Angius A, Berutti R, Reinier F, Atzeni R, Pilu R, Busonero F, Maschio A, Zara I, Sanna D, Useli A, Urru MF, Marcelli M, Cusano R, Oppo M, Zoledziewska M, Pitzalis M, Deidda F, Porcu E, Poddie F, Kang HM, Lyons R, Tarrier B, Gresham JB, Li B, Tofanelli S, Alonso S, Dei M, Lai S, Mulas A, Whalen MB, Uzzau S, Jones C, Schlessinger D, Abecasis GR, Sanna S, Sidore C, Cucca F August 2013 "Low-pass DNA sequencing of 1200 Sardinians reconstructs European Y-chromosome phylogeny" Science 341 6145: 565–9 Bibcode:2013Sci341565F doi:101126/science1237947 PMID 23908240 
  34. ^ Adams SM, Bosch E, Balaresque PL, Ballereau SJ, Lee AC, Arroyo E, López-Parra AM, Aler M, Grifo MS, Brion M, Carracedo A, Lavinha J, Martínez-Jarreta B, Quintana-Murci L, Picornell A, Ramon M, Skorecki K, Behar DM, Calafell F, Jobling MA December 2008 "The genetic legacy of religious diversity and intolerance: paternal lineages of Christians, Jews, and Muslims in the Iberian Peninsula" American Journal of Human Genetics 83 6: 725–36 doi:101016/jajhg200811007 PMC 2668061  PMID 19061982 
  35. ^ Adams SM, Bosch E, Balaresque PL, Ballereau SJ, Lee AC, Arroyo E, López-Parra AM, Aler M, Grifo MS, Brion M, Carracedo A, Lavinha J, Martínez-Jarreta B, Quintana-Murci L, Picornell A, Ramon M, Skorecki K, Behar DM, Calafell F, Jobling MA December 2008 "The genetic legacy of religious diversity and intolerance: paternal lineages of Christians, Jews, and Muslims in the Iberian Peninsula" American Journal of Human Genetics 83 6: 725–36 doi:101016/jajhg200811007 PMID 19061982 
  36. ^ Flores C, Maca-Meyer N, González AM, Oefner PJ, Shen P, Pérez JA, Rojas A, Larruga JM, Underhill PA October 2004 "Reduced genetic structure of the Iberian peninsula revealed by Y-chromosome analysis: implications for population demography" European Journal of Human Genetics 12 10: 855–63 doi:101038/sjejhg5201225 PMID 15280900 
  37. ^ Cerezo M, Achilli A, Olivieri A, Perego UA, Gómez-Carballa A, Brisighelli F, Lancioni H, Woodward SR, López-Soto M, Carracedo A, Capelli C, Torroni A, Salas A May 2012 "Reconstructing ancient mitochondrial DNA links between Africa and Europe" Genome Research 22 5: 821–6 doi:101101/gr134452111 PMC 3337428  PMID 22454235 
  38. ^ Cerezo M, Achilli A, Olivieri A, Perego UA, Gómez-Carballa A, Brisighelli F, Lancioni H, Woodward SR, López-Soto M, Carracedo A, Capelli C, Torroni A, Salas A May 2012 "Reconstructing ancient mitochondrial DNA links between Africa and Europe" Genome Research 22 5: 821–6 doi:101101/gr134452111 PMC 3337428  PMID 22454235 
  39. ^ Pereira L, Cunha C, Alves C, Amorim A April 2005 "African female heritage in Iberia: a reassessment of mtDNA lineage distribution in present times" Human Biology 77 2: 213–29 doi:101353/hub20050041 PMID 16201138 
  40. ^ a b Brehm A, Pereira L, Kivisild T, Amorim A December 2003 "Mitochondrial portraits of the Madeira and Açores archipelagos witness different genetic pools of its settlers" Human Genetics 114 1: 77–86 doi:101007/s00439-003-1024-3 PMID 14513360 
  41. ^ Alvarez L, Santos C, Ramos A, Pratdesaba R, Francalacci P, Aluja MP August 2010 "Mitochondrial DNA patterns in the Iberian Northern plateau: population dynamics and substructure of the Zamora province" American Journal of Physical Anthropology 142 4: 531–9 doi:101002/ajpa21252 PMID 20127843 
  42. ^ As regards sub-Saharan Hgs L1b, L2b, and L3b, the high frequency found in the southern regions of Zamora, 182% in Sayago and 81% in Bajo Duero, is comparable to that described for the South of Portugal, Álvarez et al 2010
  43. ^ Pereira V, Gomes V, Amorim A, Gusmão L, João Prata M September–October 2010 "Genetic characterization of uniparental lineages in populations from Southwest Iberia with past malaria endemicity" American Journal of Human Biology 22 5: 588–95 doi:101002/ajhb21049 PMID 20737604 
  44. ^ Achilli A, Olivieri A, Pala M, Metspalu E, Fornarino S, Battaglia V, Accetturo M, Kutuev I, Khusnutdinova E, Pennarun E, Cerutti N, Di Gaetano C, Crobu F, Palli D, Matullo G, Santachiara-Benerecetti AS, Cavalli-Sforza LL, Semino O, Villems R, Bandelt HJ, Piazza A, Torroni A April 2007 "Mitochondrial DNA variation of modern Tuscans supports the near eastern origin of Etruscans" American Journal of Human Genetics 80 4: 759–68 doi:101086/512822 PMC 1852723  PMID 17357081 
  45. ^ Ottoni C, Martinez-Labarga C, Vitelli L, Scano G, Fabrini E, Contini I, Biondi G, Rickards O 2009 "Human mitochondrial DNA variation in Southern Italy" Annals of Human Biology 36 6: 785–811 doi:103109/03014460903198509 PMID 19852679 
  46. ^ Behar DM, Metspalu E, Kivisild T, Achilli A, Hadid Y, Tzur S, Pereira L, Amorim A, Quintana-Murci L, Majamaa K, Herrnstadt C, Howell N, Balanovsky O, Kutuev I, Pshenichnov A, Gurwitz D, Bonne-Tamir B, Torroni A, Villems R, Skorecki K March 2006 "The matrilineal ancestry of Ashkenazi Jewry: portrait of a recent founder event" American Journal of Human Genetics 78 3: 487–97 doi:101086/500307 PMC 1380291  PMID 16404693 
  47. ^ a b Hernández CL, Soares P, Dugoujon JM, Novelletto A, Rodríguez JN, Rito T, Oliveira M, Melhaoui M, Baali A, Pereira L, Calderón R 2015 "Early Holocenic and Historic mtDNA African Signatures in the Iberian Peninsula: The Andalusian Region as a Paradigm" PLOS One 10 10: e0139784 Bibcode:2015PLoSO1039784H doi:101371/journalpone0139784 PMC 4624789  PMID 26509580 
  48. ^ Brisighelli F, Álvarez-Iglesias V, Fondevila M, Blanco-Verea A, Carracedo A, Pascali VL, Capelli C, Salas A 10 December 2012 "Uniparental markers of contemporary Italian population reveals details on its pre-Roman heritage" PLOS One 7 12: e50794 Bibcode:2012PLoSO750794B doi:101371/journalpone0050794 PMC 3519480  PMID 23251386 
  49. ^ Boattini A, Martinez-Cruz B, Sarno S, Harmant C, Useli A, Sanz P, Yang-Yao D, Manry J, Ciani G, Luiselli D, Quintana-Murci L, Comas D, Pettener D 2013 "Uniparental markers in Italy reveal a sex-biased genetic structure and different historical strata" PLOS One 8 5: e65441 Bibcode:2013PLoSO865441B doi:101371/journalpone0065441 PMC 3666984  PMID 23734255 
  50. ^ a b c d Babalini C, Martínez-Labarga C, Tolk HV, Kivisild T, Giampaolo R, Tarsi T, Contini I, Barać L, Janićijević B, Martinović Klarić I, Pericić M, Sujoldzić A, Villems R, Biondi G, Rudan P, Rickards O August 2005 "The population history of the Croatian linguistic minority of Molise southern Italy: a maternal view" European Journal of Human Genetics 13 8: 902–12 doi:101038/sjejhg5201439 PMID 15886710 
  51. ^ Fraumene C, Petretto E, Angius A, Pirastu M December 2003 "Striking differentiation of sub-populations within a genetically homogeneous isolate Ogliastra in Sardinia as revealed by mtDNA analysis" Human Genetics 114 1: 1–10 doi:101007/s00439-003-1008-3 PMID 13680359 
  52. ^ Romano V, Calì F, Ragalmuto A, D'Anna RP, Flugy A, De Leo G, Giambalvo O, Lisa A, Fiorani O, Di Gaetano C, Salerno A, Tamouza R, Charron D, Zei G, Matullo G, Piazza A January 2003 "Autosomal microsatellite and mtDNA genetic analysis in Sicily Italy" Annals of Human Genetics 67 Pt 1: 42–53 doi:101046/j1469-1809200300007x PMID 12556234 
  53. ^ a b c Auton A, Bryc K, Boyko AR, Lohmueller KE, Novembre J, Reynolds A, Indap A, Wright MH, Degenhardt JD, Gutenkunst RN, King KS, Nelson MR, Bustamante CD May 2009 "Global distribution of genomic diversity underscores rich complex history of continental human populations" Genome Research 19 5: 795–803 doi:101101/gr088898108 PMC 2675968  PMID 19218534 
  54. ^ Maca-Meyer N, González AM, Pestano J, Flores C, Larruga JM, Cabrera VM October 2003 "Mitochondrial DNA transit between West Asia and North Africa inferred from U6 phylogeography" BMC Genetics 4: 15 doi:101186/1471-2156-4-15 PMC 270091  PMID 14563219 
  55. ^ Moorjani P, Patterson N, Hirschhorn JN, Keinan A, Hao L, Atzmon G, Burns E, Ostrer H, Price AL, Reich D April 2011 McVean G, ed "The history of African gene flow into Southern Europeans, Levantines, and Jews" PLoS Genetics 7 4: e1001373 doi:101371/journalpgen1001373 PMC 3080861  PMID 21533020 
  56. ^ Bowcock AM, Kidd JR, Mountain JL, Hebert JM, Carotenuto L, Kidd KK, Cavalli-Sforza LL February 1991 "Drift, admixture, and selection in human evolution: a study with DNA polymorphisms" Proceedings of the National Academy of Sciences of the United States of America 88 3: 839–43 Bibcode:1991PNAS88839B doi:101073/pnas883839 PMC 50909  PMID 1992475 
  57. ^ Frudakis, Tony 2007 "West African ancestry in Southeastern Europe and the Middle East" Molecular photofitting: predicting ancestry and phenotype using DNA Amsterdam: Elsevier/Academic Press p 326 ISBN 0-12-088492-5 
  58. ^ Reich D, Price AL, Patterson N May 2008 "Principal component analysis of genetic data" Nature Genetics 40 5: 491–2 doi:101038/ng0508-491 PMID 18443580 
  59. ^ Sanchez JJ, Phillips C, Børsting C, Balogh K, Bogus M, Fondevila M, Harrison CD, Musgrave-Brown E, Salas A, Syndercombe-Court D, Schneider PM, Carracedo A, Morling N May 2006 "A multiplex assay with 52 single nucleotide polymorphisms for human identification" Electrophoresis 27 9: 1713–24 doi:101002/elps200500671 PMID 16586411 
  60. ^ Characterizing the history of sub-Saharan African gene flow into southern Europe, Moorjani et al 2009, Department of Genetics, Harvard Medical School
  61. ^ Moorjani P, Patterson N, Hirschhorn JN, Keinan A, Hao L, Atzmon G, Burns E, Ostrer H, Price AL, Reich D April 2011 McVean G, ed "The history of African gene flow into Southern Europeans, Levantines, and Jews" PLoS Genetics 7 4: e1001373 doi:101371/journalpgen1001373 PMC 3080861  PMID 21533020 
  62. ^ Reich D, Thangaraj K, Patterson N, Price AL, Singh L September 2009 "Reconstructing Indian population history" Nature 461 7263: 489–94 Bibcode:2009Natur461489R doi:101038/nature08365 PMC 2842210  PMID 19779445 
  63. ^ Lazaridis I, Patterson N, Mittnik A, Renaud G, Mallick S, Kirsanow K, et al September 2014 "Ancient human genomes suggest three ancestral populations for present-day Europeans" Nature 513 7518: 409–13 arXiv:13126639  Bibcode:2014Natur513409L bioRxiv 001552  doi:101038/nature13673 PMC 4170574  PMID 25230663 
  64. ^ Pino-Yanes M, Corrales A, Basaldúa S, Hernández A, Guerra L, Villar J, Flores C March 2011 O'Rourke D, ed "North African influences and potential bias in case-control association studies in the Spanish population" PLOS One 6 3: e18389 Bibcode:2011PLoSO6E8389P doi:101371/journalpone0018389 PMC 3068190  PMID 21479138 
  65. ^ Calderón R, Ambrosio B, Guitard E, González-Martín A, Aresti U, Dugoujon JM December 2006 "Genetic position of Andalusians from Huelva in relation to other European and North African populations: a study based on GM and KM allotypes" Human Biology 78 6: 663–79 doi:101353/hub20070008 PMID 17564246 
  66. ^ a b Calderón R, Lodeiro R, Varela TA, Fariña J, Ambrosio B, Guitard E, González-Martín A, Dugoujon JM June 2007 "GM and KM immunoglobulin allotypes in the Galician population: new insights into the peopling of the Iberian Peninsula" BMC Genetics 8 1: 37 doi:101186/1471-2156-8-37 PMC 1934380  PMID 17597520 
  67. ^ Giraldo MP, Esteban E, Aluja MP, Nogués RM, Backés-Duró C, Dugoujon JM, Moral P November 2001 "Gm and Km alleles in two Spanish Pyrenean populations Andorra and Pallars Sobirà: a review of Gm variation in the Western Mediterranean basin" Annals of Human Genetics 65 Pt 6: 537–48 doi:101046/j1469-180920016560537x PMID 11851984 
  68. ^ Cerutti N, Dugoujon JM, Guitard E, Rabino Massa E January 2004 "Gm and Km immunoglobulin allotypes in Sicily" Immunogenetics 55 10: 674–81 doi:101007/s00251-003-0628-z PMID 14652700 
  69. ^ Sallares R, Bouwman A, Anderung C July 2004 "The spread of malaria to Southern Europe in antiquity: new approaches to old problems" Medical History 48 3: 311–28 doi:101017/s0025727300007651 PMC 547919  PMID 16021928 
  70. ^ Rund D, Kornhendler N, Shalev O, Oppenheim A October 1990 "The origin of sickle cell alleles in Israel" Human Genetics 85 5: 521–4 doi:101007/BF00194229 PMID 1977682 
  71. ^ Bloom, Miriam 1995 Understanding Sickle Cell Disease Jackson: University Press of Mississippi ISBN 0-87805-745-5 
  72. ^ Ragusa A, Frontini V, Lombardo M, Amata S, Lombardo T, Labie D, Krishnamoorthy R, Nagel RL August 1992 "Presence of an African beta-globin gene cluster haplotype in normal chromosomes in Sicily" American Journal of Hematology 40 4: 313–5 doi:101002/ajh2830400413 PMID 1503087 
  73. ^ Ragusa A, Lombardo M, Sortino G, Lombardo T, Nagel RL, Labie D February 1988 "Beta S gene in Sicily is in linkage disequilibrium with the Benin haplotype: implications for gene flow" American Journal of Hematology 27 2: 139–41 doi:101002/ajh2830270214 PMID 2893541 
  74. ^ Monteiro C, Rueff J, Falcao AB, Portugal S, Weatherall DJ, Kulozik AE June 1989 "The frequency and origin of the sickle cell mutation in the district of Coruche/Portugal" Human Genetics 82 3: 255–8 doi:101007/BF00291165 PMID 2731937 
  75. ^ Bardakdjian J, Wajcman H September 2004 "" La Revue Du Praticien in French 54 14: 1531–3 PMID 15558961 
  76. ^ Bardakdjian-Michau J, Bahuau M, Hurtrel D, Godart C, Riou J, Mathis M, Goossens M, Badens C, Ducrocq R, Elion J, Perini JM January 2009 "Neonatal screening for sickle cell disease in France" Journal of Clinical Pathology 62 1: 31–3 doi:101136/jcp2008058867 PMID 19103855 
  77. ^ "Erreur / InVS / Informations générales / Accueil" Invssantefr Retrieved 21 May 2018 
  78. ^ Goring-Morris, Nigel et al 2009 The Dynamics of Pleistocene and Early Holocene Settlement Patterns in the Levant: An Overview In Transitions in Prehistory: Essays in Honor of Ofer Bar-Yosef eds John J Shea and Daniel E Lieberman Oxbow Books, 2009 ISBN 9781842173404
  79. ^ Olszewski, DI 2006 "Issues in the Levantine Epipaleolithic : The Madamaghan, Nebekian and Qalkhan Levant Epipaleolithic" Paléorient 32 1: 19–26 doi:103406/paleo20065168 
  80. ^ Bar-Yosef, O 1987 "Pleistocene connexions between Africa and Southwest Asia: an archaeological perspective" The African Archaeological Review 5 1: 29–38 doi:101007/BF01117080 
  81. ^ Richter, Tobias; et al 2011 "Interaction before Agriculture: Exchanging Material and Sharing Knowledge in the Final Pleistocene Levant" Cambridge Archaeological Journal 21 1: 95–114 doi:101017/S0959774311000060 

References

  • Moorjani P, Patterson N, Hirschhorn JN, Keinan A, Hao L, Atzmon G, Burns E, Ostrer H, Price AL, Reich D April 2011 McVean G, ed "The history of African gene flow into Southern Europeans, Levantines, and Jews" PLoS Genetics 7 4: e1001373 doi:101371/journalpgen1001373 PMC 3080861  PMID 21533020 
  • Brace CL, Seguchi N, Quintyn CB, Fox SC, Nelson AR, Manolis SK, Qifeng P January 2006 "The questionable contribution of the Neolithic and the Bronze Age to European craniofacial form" Proceedings of the National Academy of Sciences of the United States of America 103 1: 242–7 Bibcode:2006PNAS103242B doi:101073/pnas0509801102 PMC 1325007  PMID 16371462 
  • Bar-Yosef, O 1987 "Pleistocene connexions between Africa and Southwest Asia: an archaeological perspective" African Archaeological Review 5 1: 29–38 doi:101007/BF01117080 
  • Casas MJ, Hagelberg E, Fregel R, Larruga JM, González AM December 2006 "Human mitochondrial DNA diversity in an archaeological site in al-Andalus: genetic impact of migrations from North Africa in medieval Spain" American Journal of Physical Anthropology 131 4: 539–51 doi:101002/ajpa20463 PMID 16685727 
  • Cavalli-Sforza LL July 1997 "Genes, peoples, and languages" Proceedings of the National Academy of Sciences of the United States of America 94 15: 7719–24 Bibcode:1997PNAS947719C doi:101073/pnas94157719 PMC 33682  PMID 9223254 
  • Cherni L, Fernandes V, Pereira JB, Costa MD, Goios A, Frigi S, Yacoubi-Loueslati B, Amor MB, Slama A, Amorim A, El Gaaied AB, Pereira L June 2009 "Post-last glacial maximum expansion from Iberia to North Africa revealed by fine characterization of mtDNA H haplogroup in Tunisia" American Journal of Physical Anthropology 139 2: 253–60 doi:101002/ajpa20979 PMID 19090581 
  • 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:101086/386294 PMC 1181964  PMID 15042509 
  • Ennafaa H, Cabrera VM, Abu-Amero KK, González AM, Amor MB, Bouhaha R, Dzimiri N, Elgaaïed AB, Larruga JM February 2009 "Mitochondrial DNA haplogroup H structure in North Africa" BMC Genetics 10 1: 8 doi:101186/1471-2156-10-8 PMC 2657161  PMID 19243582 
  • Gonçalves R, Freitas A, Branco M, Rosa A, Fernandes AT, Zhivotovsky LA, Underhill PA, Kivisild T, Brehm A July 2005 "Y-chromosome lineages from Portugal, Madeira and Açores record elements of Sephardim and Berber ancestry" Annals of Human Genetics 69 Pt 4: 443–54 doi:101111/j1529-8817200500161x PMID 15996172 
  • González AM, Brehm A, Pérez JA, Maca-Meyer N, Flores C, Cabrera VM April 2003 "Mitochondrial DNA affinities at the Atlantic fringe of Europe" American Journal of Physical Anthropology 120 4: 391–404 doi:101002/ajpa10168 PMID 12627534 
  • 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
  • Maliarchuk BA, Czarny J 2005 "" Molekuliarnaia Biologiia in Russian 39 5: 806–12 doi:101007/s11008-005-0085-x PMID 16240714 
  • Malyarchuk BA, Derenko M, Perkova M, Grzybowski T, Vanecek T, Lazur J September 2008 "Reconstructing the phylogeny of African mitochondrial DNA lineages in Slavs" European Journal of Human Genetics 16 9: 1091–6 doi:101038/ejhg200870 PMID 18398433 
  • Ricaut FX, Waelkens M October 2008 "Cranial discrete traits in a Byzantine population and eastern Mediterranean population movements" Human Biology 80 5: 535–64 doi:103378/1534-6617-805535 PMID 19341322 
  • Pereira L, Prata MJ, Amorim A November 2000 "Diversity of mtDNA lineages in Portugal: not a genetic edge of European variation" Annals of Human Genetics 64 Pt 6: 491–506 doi:101046/j1469-180920006460491x PMID 11281213 
  • Pereira L, Cunha C, Alves C, Amorim A April 2005 "African female heritage in Iberia: a reassessment of mtDNA lineage distribution in present times" Human Biology 77 2: 213–29 doi:101353/hub20050041 PMID 16201138 
  • Semino O, Magri C, Benuzzi G, Lin AA, Al-Zahery N, Battaglia V, Maccioni L, Triantaphyllidis C, Shen P, Oefner PJ, Zhivotovsky LA, King R, Torroni A, Cavalli-Sforza LL, Underhill PA, Santachiara-Benerecetti AS May 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: 1023–34 doi:101086/386295 PMC 1181965  PMID 15069642 

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