African admixture in Europe

{{See also|Genetic history of Europe}}

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 Western Asia around 10,000 BCE. Agriculture was introduced into Europe c. 7000 BC by migrating farmers from Anatolia, known as Anatolian Neolithic Farmers in the genetics literature.{{cite journal |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC5003663/ |journal=Nature |volume=536 |issue=7617 |date=2016 |title=Genomic insights into the origin of farming in the ancient Near East |last1=Lazaridis |first1=Iosif |display-authors=etal |pages=419–424 |doi=10.1038/nature19310|hdl=10230/35986 |hdl-access=free }}{{cite journal |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC5069350/?report=classic |journal=Current Biology |volume=26 |issue=19 |date=2016 |title=The Demographic Development of the First Farmers in Anatolia |last1=Kılınç |first1=Gülşah Merve |display-authors=etal |pages=2659–2666 |doi=10.1016/j.cub.2016.07.057|hdl=11511/37135 |hdl-access=free }} These farmers either replaced or interbred with local European hunter-gather populations that had been living in Europe since the Paleolithic.{{cite journal | vauthors = Cavalli-Sforza LL, Piazza A | title = Human genomic diversity in Europe: a summary of recent research and prospects for the future | journal = European Journal of Human Genetics | volume = 1 | issue = 1 | pages = 3–18 | year = 1993 | pmid = 7520820 | doi = 10.1159/000472383 | s2cid = 25475102 }} Early Neolithic farmers in Anatolia and Europe were similar genetically to modern-day southern European populations.{{cite journal |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC5069350/?report=classic |journal=Current Biology |volume=26 |issue=19 |date=2016 |title=The Demographic Development of the First Farmers in Anatolia |last1=Kılınç |first1=Gülşah Merve |display-authors=etal |pages=2659–2666 |doi=10.1016/j.cub.2016.07.057|hdl=11511/37135 |hdl-access=free }}{{cite journal |url=https://pubmed.ncbi.nlm.nih.gov/26748850/ |journal=Current Biology |volume=26 |issue=2 |date=2016 |title=Genomic Evidence Establishes Anatolia as the Source of the European Neolithic Gene Pool |last1=Omrak |first1=Ayça |display-authors=etal |pages=270-275 |doi=10.1016/j.cub.2015.12.019|doi-access=free }}{{cite journal |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC9166250/ |journal=Cell |volume=185 |issue=11 |date=2022 |title=The genomic origins of the world’s first farmers |last1=Marchi |first1=Nina |display-authors=etal |pages=1842–1859.e18 |doi=10.1016/j.cell.2022.04.008|hdl=20.500.11850/563063 |hdl-access=free }} European Neolithic farmers later introduced agriculture to Northwest Africa c. 5400 BC from southwestern Europe, contributing significantly to the genetic formation of the ancient (and modern) population of North Africa.{{cite journal |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC10266975/ |journal=Nature |volume=618 |issue=7965 |date=2023 |title=Northwest African Neolithic initiated by migrants from Iberia and Levant |last1=Simões |first1=Luciana G |display-authors=etal |pages=550–556 |doi=10.1038/s41586-023-06166-6|hdl=10272/23423 |hdl-access=free }}{{cite journal |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC6042094/ |journal=Proc Natl Acad Sci USA |volume=115 |issue=26 |date=2018 |title=Ancient genomes from North Africa evidence prehistoric migrations to the Maghreb from both the Levant and Europe |last1=Fregel |first1=Rosa |display-authors=etal |pages=6774–6779 |doi=10.1073/pnas.1800851115|pmc=6042094 }}{{cite web |url=https://oxfordre.com/africanhistory/display/10.1093/acrefore/9780190277734.001.0001/acrefore-9780190277734-e-143 |website=Oxford Research Encyclopedia of African History |title=Genetics and the African Past |date=2022 |last1=Prendergast |first1=Mary E. |display-authors=etal |doi=10.1093/acrefore/9780190277734.013.143}} A later expansion from Europe into North Africa by the Late Neolithic/Early Bronze Age Bell Beaker culture further contributed to the North African genepool.{{cite journal |url= https://academic.oup.com/hmg/article/30/R1/R64/6028725?login=false |journal=Human Molecular Genetics |volume=30 |issue=R1 |date=2020 |title=The demography of the Canary Islands from a genetic perspective |last1=Fregel |first1=Rosa |display-authors=etal |pages=R64–R71 |doi=10.1093/hmg/ddaa262 |pmid=33295602 |doi-access=free}}{{cite journal |last1=Serrano |first1=J. G. |display-authors=etal |title=The genomic history of the indigenous people of the Canary Islands |journal=Nature Communications |volume=14 |issue=4641 |date=2023 |at="Supplementary Information" section, p. 13 |doi=10.1038/s41467-023-40198-w |doi-access=free |pmid=37582830 |pmc=10427657 |bibcode=2023NatCo..14.4641S |hdl=10553/124288 |hdl-access=free}}

It has been suggested that the first farmers in West Asia, descended from Natufian hunter-gatherers, reflected some North African influences or vice versa.{{Cite journal|vauthors=Bar-Yosef O|author-link=Ofer Bar-Yosef|title=Pleistocene connexions between Africa and Southwest Asia: an archaeological perspective |doi=10.1007/BF01117080|journal=African Archaeological Review |year=1987 |volume=5|issue=1|pages=29–38|s2cid=132865471}} There have been suggestions that some genetic lineages found in the Middle East arrived there during this period or earlier.{{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}} The first agricultural societies in the Middle East are generally thought to have emerged from the Natufian culture between 12,000 and 10,000 BCE. The Natufians were widely semi-sedentary even before the introduction of agriculture.

In historical times, there has been a period of north African influence in southern Europe, especially in 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.{{cite journal | vauthors = Malyarchuk BA, Czarny J | title = [African DNA lineages in mitochondrial gene pool of Europeans] | language = ru | journal = Molekuliarnaia Biologiia | volume = 39 | issue = 5 | pages = 806–12 | year = 2005 | pmid = 16240714 | doi = 10.1007/s11008-005-0085-x | s2cid = 2527074}}

File:Overview of the genetic structure and global ancestry inference in a selection of European, Near Eastern, and African populations.png

• Hernandez et al. (2020) identified 11.17 ± 1.87% North African ancestry in southern Portuguese samples (from a population similar to modern northern Moroccans and Algerians), 9.28 ± 1.79% of such ancestry in western Andalusians, and an average of 1.41 ± 0.72% sub-Saharan ancestry in southern Iberians (using Yoruba as a proxy source). Substantially lower levels of North African admixture were further detected in Northern Italians (0.77%) and Tuscans (1%).

File:NorthAfricanAdmixtureIberianPeninsula.jpg

• Bycroft et al. (2019) identified regionally varying fractions of Northwest African ancestry in modern Iberians, ranging from 0–12%. This ancestry was found to be from a source population similar to modern Northwest Africans. The admixture was dated to 860–1120 CE, associated with the Muslim conquest and subsequent Reconquista. The highest levels of Northwest African admixture were identified in western Iberia whilst the lowest levels were found in the Basque region and an area in the North East roughly corresponding to the 14th-century Crown of Aragon. They also found some evidence for a second admixture event in Portuguese and Southern Spanish groups involving a second North African population within which a small sub-Saharan African component was detected. This admixture event was dated to approximately 1300 CE.

File:Map of African admixture in European populations.png

• Botigué et al. (2013) analysed genome-wide SNP data from over 2,000 modern individuals from Iberia. They estimated an average of 4% to 12% Northwest African admixture in modern Iberians (with low or zero levels in Basques), whereas populations in southeastern Europe had less than 2% of such admixture. Sub-Saharan African ancestry was detected at less than 1% in Europe, with the exception of the Canary Islands.{{cite journal |vauthors=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 |date=July 2013 |title=Gene flow from North Africa contributes to differential human genetic diversity in southern Europe |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=110 |issue=29 |pages=11791–6 |bibcode=2013PNAS..11011791B |doi=10.1073/pnas.1306223110 |pmc=3718088 |pmid=23733930 |doi-access=free}}{{cite web |url=http://www.huffingtonpost.es/2013/06/03/los-espanoles-somos-los-e_n_3379814.html |website=Huffington Post |title=Los españoles somos los europeos con más genes magrebíes |date=2013 |quote=Comprobaron que entre un 4% y un 20% del genoma de los españoles es compartido con los norteafricanos. 'La cifra del 20% sólo se da en Canarias, para el resto del país oscila entre el 10% y 12%', explica Comas. Sólo los vascos de la muestra no presentan ese influjo norteafricano." English translation: "They found that between 4% and 20% of the genome of Spaniards is shared with North Africans. “The figure of 20% only occurs in the Canary Islands, for the rest of the country it ranges between 10% and 12%,” explains Comas. Only the Basques in the sample do not show this North African influence.}}

• Lazarids et al. (2014) detected 12.6 ± 2% North African ancestry and an average of 1.5 ± 0.2% sub-Saharan ancestry in the Spanish population.{{Cite journal |last=Lazaridis |first=Iosif |last2=Patterson |first2=Nick |last3=Mittnik |first3=Alissa |last4=Renaud |first4=Gabriel |last5=Mallick |first5=Swapan |last6=Kirsanow |first6=Karola |last7=Sudmant |first7=Peter H. |last8=Schraiber |first8=Joshua G. |last9=Castellano |first9=Sergi |last10=Lipson |first10=Mark |last11=Berger |first11=Bonnie |last12=Economou |first12=Christos |last13=Bollongino |first13=Ruth |last14=Fu |first14=Qiaomei |last15=Bos |first15=Kirsten I. |date=17 September 2014 |title=Ancient human genomes suggest three ancestral populations for present-day Europeans |url=https://pmc.ncbi.nlm.nih.gov/articles/instance/4170574/bin/NIHMS613260-supplement-supplement_1.pdf |journal=Nature |language=en |volume=513 |issue=7518 |pages=409–413 |doi=10.1038/nature13673 |issn=1476-4687}}

• Moorjani et al. (2011) estimated that some Southern Europeans have inherited 1%–3% sub-Saharan ancestry (approximately 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". This admixture was not identified in Northern Europeans.{{cite journal |vauthors=Moorjani P, Patterson N, Hirschhorn JN, Keinan A, Hao L, Atzmon G, Burns E, Ostrer H, Price AL, Reich D |date=April 2011 |title=The history of African gene flow into Southern Europeans, Levantines, and Jews |journal=PLOS Genetics |volume=7 |issue=4 |pages=e1001373 |doi=10.1371/journal.pgen.1001373 |pmc=3080861 |pmid=21533020 |doi-access=free |veditors=McVean G}}

• Pino-Yanes et al. (2011) found that from an autosomal analysis, the average Northwest African influence is about 17% in Canary Islanders, with a wide interindividual variation ranging from 0% to 96%. 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.{{cite journal |vauthors=Pino-Yanes M, Corrales A, Basaldúa S, Hernández A, Guerra L, Villar J, Flores C |date=March 2011 |title=North African influences and potential bias in case-control association studies in the Spanish population |journal=PLOS ONE |volume=6 |issue=3 |pages=e18389 |bibcode=2011PLoSO...618389P |doi=10.1371/journal.pone.0018389 |pmc=3068190 |pmid=21479138 |doi-access=free |veditors=O'Rourke D}}

• Ricardo Rodríguez-Varela et al. (2024), studied 37 individuals from the 7th to 11th Century in Las Gobas, a necropolis in northern Spain. They found there was no significant increase of North African (Berber) or Middle East ancestries over time, before and after the Umayyad expansion. The authors proposed it may have been due to the communities isolation. However, North African admixture was detected in Visigothic individuals from southern Spain, hypothesized to be due to Punic movement into Iberia or pre-Islamic contact between southern Spain and North Africa.{{Cite journal |last1=Rodríguez-Varela |first1=Ricardo |last2=Yaka |first2=Reyhan |last3=Pochon |first3=Zoé |last4=Sanchez-Pinto |first4=Iban |last5=Solaun |first5=José Luis |last6=Naidoo |first6=Thijessen |last7=Guinet |first7=Benjamin |last8=Pérez-Ramallo |first8=Patxi |last9=Lagerholm |first9=Vendela Kempe |last10=de Anca Prado |first10=Violeta |last11=Valdiosera |first11=Cristina |last12=Krzewińska |first12=Maja |last13=Herrasti |first13=Lourdes |last14=Azkarate |first14=Agustín |last15=Götherström |first15=Anders |date=2024-08-30 |title=Five centuries of consanguinity, isolation, health, and conflict in Las Gobas: A Northern Medieval Iberian necropolis |journal=Science Advances |language=en |volume=10 |issue=35 |pages=eadp8625 |doi=10.1126/sciadv.adp8625 |issn=2375-2548 |pmc=11352919 |pmid=39196943}}
• MacRoberts, Rebecca Anne et al. (2024) studied remains of the 8th-10th century AD in Portugal. Ancient DNA analysis conducted on three individuals revealed maternal (mtDNA) and paternal (Y-chromosome) lineages showing possible North African origin for some of the individuals. The mobility of females in this population was said to be higher than males, possibly because of the patriarchal structures practiced by Berber and Arab communities.{{Cite journal |last1=MacRoberts |first1=Rebecca Anne |last2=Liberato |first2=Marco |last3=Roca-Rada |first3=Xavier |last4=Valente |first4=Maria João |last5=Relvado |first5=Claudia |last6=Fernandes |first6=Teresa Matos |last7=Dias |first7=Cristina Barrocas |last8=Llamas |first8=Bastien |last9=Vilar |first9=Hermínia Vasconcelos |last10=Schöne |first10=Bernd R. |last11=Ribeiro |first11=Sara |last12=Santos |first12=José Francisco |last13=Teixeira |first13=João C. |last14=Maurer |first14=Anne-France |date=2024-03-06 |title=Shrouded in history: Unveiling the ways of life of an early Muslim population in Santarém, Portugal (8th– 10th century AD) |journal=PLOS ONE |language=en |volume=19 |issue=3 |pages=e0299958 |doi=10.1371/journal.pone.0299958 |doi-access=free |issn=1932-6203 |pmc=10917335 |pmid=38446809|bibcode=2024PLoSO..1999958M }}
• Silva, M. et al. (2021), analyzed the genome for an individual who was buried in an Islamic necropolis at Segorbe, Spain. The sample's uniparentals pointed to North African origin, but on the autosomal level he displays both European and Northern African-related ancestries.{{Cite journal |last1=Silva |first1=Marina |last2=Oteo-García |first2=Gonzalo |last3=Martiniano |first3=Rui |last4=Guimarães |first4=João |last5=von Tersch |first5=Matthew |last6=Madour |first6=Ali |last7=Shoeib |first7=Tarek |last8=Fichera |first8=Alessandro |last9=Justeau |first9=Pierre |last10=Foody |first10=M. George B. |last11=McGrath |first11=Krista |last12=Barrachina |first12=Amparo |last13=Palomar |first13=Vicente |last14=Dulias |first14=Katharina |last15=Yau |first15=Bobby |date=2021-09-13 |title=Biomolecular insights into North African-related ancestry, mobility and diet in eleventh-century Al-Andalus |journal=Scientific Reports |language=en |volume=11 |issue=1 |pages=18121 |doi=10.1038/s41598-021-95996-3 |pmid=34518562 |pmc=8438022 |bibcode=2021NatSR..1118121S |issn=2045-2322}}

• Olalde et al. (2019) found evidence for 'sporadic contacts' between Iberia and North Africa in the Copper Age and Bronze Age. A male sample from central Iberia, dating from 2473–2030 cal BCE, was found to cluster with modern and ancient North Africans, characterised by ancestry from both Late Pleistocene North Africans and Early Neolithic Europeans. Another Bronze Age sample had 25% such North African ancestry. However, North African ancestry only became widespread in Iberia in the past ~2000 years, associated with the Roman Empire or earlier Punic presence and the later period of Muslim rule. The study analysed 45 samples from southeastern Spain dating from the 3rd-16th centuries CE, all of which fell outside the genetic variation of preceding Iberian Iron Age populations, harbouring ancestry from both southern European and North African populations, as well as additional Levantine-related ancestry. 2 samples out of 23 dating from the 10th to 16th centuries were also found to have partial sub-Saharan ancestry, which was not identified in earlier samples. Present-day southern Iberians harbour less African ancestry than Muslim period samples, likely due to subsequent population expulsions and repopulation from the north, as supported by historical sources and genetic analysis of present-day groups.
• Gleize, Yves et al. (2016), discovered that the paternal lineages of Muslim cemeteries in France had 3 individuals with North African ancestry. The analyses was said to confirm the Berber origin of some of the first Islamic troops spreading through Europe, and also indicated the co-existence of communities practicing Christian and Muslim funerary customs.{{Cite journal |last1=Gleize |first1=Yves |last2=Mendisco |first2=Fanny |last3=Pemonge |first3=Marie-Hélène |last4=Hubert |first4=Christophe |last5=Groppi |first5=Alexis |last6=Houix |first6=Bertrand |last7=Deguilloux |first7=Marie-France |last8=Breuil |first8=Jean-Yves |date=2016-02-24 |title=Early Medieval Muslim Graves in France: First Archaeological, Anthropological and Palaeogenomic Evidence |journal=PLOS ONE |language=en |volume=11 |issue=2 |pages=e0148583 |doi=10.1371/journal.pone.0148583 |doi-access=free |issn=1932-6203 |pmc=4765927 |pmid=26910855|bibcode=2016PLoSO..1148583G }}

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, as well as Early European Farmer ancestry. This altogether indicates that the Late Neolithic Kelif el Boroud inhabitants were ancestral to contemporary populations in the area, but also experienced substantial gene flow from Europe.{{cite bioRxiv| vauthors = Fregel R, Méndez FL, Bokbot Y, Martín-Socas D, Camalich-Massieu MD, Ávila-Arcos MC, Underhill PA, Shapiro B, Wojcik G, Rasmussen M, Soares AE |display-authors = 6 |year=2017 |title=Neolithization of North Africa involved the migration of people from both the Levant and Europe|biorxiv=10.1101/191569}}

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{{cite journal | vauthors = Rando JC, Cabrera VM, Larruga JM, Hernández M, González AM, Pinto F, Bandelt HJ | title = Phylogeographic patterns of mtDNA reflecting the colonization of the Canary Islands | journal = Annals of Human Genetics | volume = 63 | issue = Pt 5 | pages = 413–28 | date = September 1999 | pmid = 10735583 | doi = 10.1046/j.1469-1809.1999.6350413.x| s2cid = 25089862 }}{{cite journal | vauthors = González AM, Larruga JM, Abu-Amero KK, Shi Y, Pestano J, Cabrera VM | title = Mitochondrial lineage M1 traces an early human backflow to Africa | journal = BMC Genomics | volume = 8 | issue = 1 | pages = 223 | date = July 2007 | pmid = 17620140 | pmc = 1945034 | doi = 10.1186/1471-2164-8-223 | doi-access = free }} 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.{{cite journal | vauthors = Ennafaa H, Cabrera VM, Abu-Amero KK, González AM, Amor MB, Bouhaha R, Dzimiri N, Elgaaïed AB, Larruga JM | title = Mitochondrial DNA haplogroup H structure in North Africa | journal = BMC Genetics | volume = 10 | issue = 1 | pages = 8 | date = February 2009 | pmid = 19243582 | pmc = 2657161 | doi = 10.1186/1471-2156-10-8 | doi-access = free }} Such bidirectional migrations between Africa and Eurasia complicate the task of defining admixture.

One proposed example of Holocene gene flow from North Africa to Europe, via the Middle East, is thought to be E1b1b, which is considered to have emerged about 40,000 years ago in the Horn of Africa, and branches of it would have then migrated to the Middle East by 15,000 years ago during the late Pleistocene period.{{cite journal | vauthors = Trombetta B, D'Atanasio E, Massaia A, Ippoliti M, Coppa A, Candilio F, Coia V, Russo G, Dugoujon JM, Moral P, Akar N, Sellitto D, Valesini G, Novelletto A, Scozzari R, Cruciani F | display-authors = 6 | title = Phylogeographic Refinement and Large Scale Genotyping of Human Y Chromosome Haplogroup E Provide New Insights into the Dispersal of Early Pastoralists in the African Continent | journal = Genome Biology and Evolution | volume = 7 | issue = 7 | pages = 1940–1950 | date = June 2015 | pmid = 26108492 | pmc = 4524485 | doi = 10.1093/gbe/evv118 }}

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.{{cite journal |vauthors=Cruciani F, La Fratta R, Santolamazza P, Sellitto D, Pascone R, Moral P, Watson E, Guida V, Colomb EB, Zaharova B, Lavinha J, Vona G, Aman R, Cali F, Akar N, Richards M, Torroni A, Novelletto A, Scozzari R |date=May 2004 |title=Phylogeographic analysis of haplogroup E3b (E-M215) y chromosomes reveals multiple migratory events within and out of Africa |journal=American Journal of Human Genetics |volume=74 |issue=5 |pages=1014–22 |doi=10.1086/386294 |pmc=1181964 |pmid=15042509}}{{cite journal | vauthors = 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 | title = 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 | journal = American Journal of Human Genetics | volume = 74 | issue = 5 | pages = 1023–34 | date = May 2004 | pmid = 15069642 | pmc = 1181965 | doi = 10.1086/386295}}{{cite journal | vauthors = Underhill PA, Passarino G, Lin AA, Shen P, Mirazón Lahr M, Foley RA, Oefner PJ, Cavalli-Sforza LL | title = The phylogeography of Y chromosome binary haplotypes and the origins of modern human populations | journal = Annals of Human Genetics | volume = 65 | issue = Pt 1 | pages = 43–62 | date = January 2001 | pmid = 11415522 | doi = 10.1046/j.1469-1809.2001.6510043.x | doi-access = free }}{{efn|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.}}{{efn|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.}}

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, haplogroup E was observed at a frequency of 7.2%. The timing of this movement has been given widely varying estimates.{{Cite journal|vauthors=Lancaster A|year=2009|url=http://www.jogg.info/51/files/Lancaster.pdf|journal=Journal of Genetic Genealogy|volume=5|issue=1|title=Y Haplogroups, Archaeological Cultures and Language Families: a Review of the Multidisciplinary Comparisons using the case of E-M35|access-date=2009-09-03|archive-date=2016-05-06|archive-url=https://web.archive.org/web/20160506150956/http://www.jogg.info/51/files/Lancaster.pdf|url-status=dead}}{{MEDRS|date=August 2010}} In much of Europe, frequencies of E lineages are very low, usually less than 1%. For example, the frequency of such lineages are 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 Northeast Africa, Northwest Africa and 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=1.3%), and among Albanians in Calabria (2/68=2.9%). The distribution of haplogroup E1a lineages in Portugal was independent of the distribution of the younger and more ubiquitous E1b1a.{{cite journal | vauthors = Gonçalves R, Freitas A, Branco M, Rosa A, Fernandes AT, Zhivotovsky LA, Underhill PA, Kivisild T, Brehm A | title = Y-chromosome lineages from Portugal, Madeira and Açores record elements of Sephardim and Berber ancestry | journal = Annals of Human Genetics | volume = 69 | issue = Pt 4 | pages = 443–54 | date = July 2005 | pmid = 15996172 | doi = 10.1111/j.1529-8817.2005.00161.x| hdl = 10400.13/3018 | s2cid = 3229760 | hdl-access = free }} This distribution is consistent with a prehistoric migration from Africa to Iberia, possibly alongside mtDNA haplogroup U6. In Majorcans, Sub-Saharan Y-DNA lineage E-V38 was found at a total of 3.2% (2/62).{{cite journal | vauthors = 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 | title = The genetic legacy of religious diversity and intolerance: paternal lineages of Christians, Jews, and Muslims in the Iberian Peninsula | journal = American Journal of Human Genetics | volume = 83 | issue = 6 | pages = 725–36 | date = December 2008 | pmid = 19061982 | pmc = 2668061 | doi = 10.1016/j.ajhg.2008.11.007 | url = }} 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).{{cite journal | vauthors = Flores C, Maca-Meyer N, González AM, Oefner PJ, Shen P, Pérez JA, Rojas A, Larruga JM, Underhill PA | title = Reduced genetic structure of the Iberian peninsula revealed by Y-chromosome analysis: implications for population demography | journal = European Journal of Human Genetics | volume = 12 | issue = 10 | pages = 855–63 | date = October 2004 | pmid = 15280900 | doi = 10.1038/sj.ejhg.5201225 | doi-access = free }} In Sardinians, Sub-Saharan Y-DNA lineages A1b1b2b and E1a1 were found at a total of 1.0% (A1b1b2b 0.5% / E1a1 0.5%).{{cite journal | vauthors = 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 | title = Low-pass DNA sequencing of 1200 Sardinians reconstructs European Y-chromosome phylogeny | journal = Science | volume = 341 | issue = 6145 | pages = 565–9 | date = August 2013 | pmid = 23908240 | doi = 10.1126/science.1237947 | bibcode = 2013Sci...341..565F | pmc = 5500864 }}

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.{{cite journal | vauthors = King TE, Parkin EJ, Swinfield G, Cruciani F, Scozzari R, Rosa A, Lim SK, Xue Y, Tyler-Smith C, Jobling MA | title = Africans in Yorkshire? The deepest-rooting clade of the Y phylogeny within an English genealogy | journal = European Journal of Human Genetics | volume = 15 | issue = 3 | pages = 288–93 | date = March 2007 | pmid = 17245408 | pmc = 2590664 | doi = 10.1038/sj.ejhg.5201771}}

File:E1b1bRoute.png

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) 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 41.1% among "pasiegos" from Cantabria, Spain. That is the highest frequency observed in Europe to date.{{cite journal | vauthors = Di Giacomo F, Luca F, Popa LO, Akar N, Anagnou N, Banyko J, Brdicka R, Barbujani G, Papola F, Ciavarella G, Cucci F, Di Stasi L, Gavrila L, Kerimova MG, Kovatchev D, Kozlov AI, Loutradis A, Mandarino V, Mammi' C, Michalodimitrakis EN, Paoli G, Pappa KI, Pedicini G, Terrenato L, Tofanelli S, Malaspina P, Novelletto A | display-authors = 6 | title = Y chromosomal haplogroup J as a signature of the post-neolithic colonization of Europe | journal = Human Genetics | volume = 115 | issue = 5 | pages = 357–371 | date = October 2004 | pmid = 15322918 | doi = 10.1007/s00439-004-1168-9 | s2cid = 18482536 }} From an analysis of the Y-chromosome with 659 samples from Southern Portugal, 680 from Northern Spain, 37 samples from Andalusia, 915 samples from mainland Italy, and 93 samples from Sicily found significantly higher levels of North African male ancestry in Portugal, Spain and Sicily (7.1%, 7.7% and 7.5% respectively) than in peninsular Italy (1.7%).{{cite journal | vauthors = Capelli C, Onofri V, Brisighelli F, Boschi I, Scarnicci F, Masullo M, Ferri G, Tofanelli S, Tagliabracci A, Gusmao L, Amorim A, Gatto F, Kirin M, Merlitti D, Brion M, Verea AB, Romano V, Cali F, Pascali V | display-authors = 6 | title = Moors and Saracens in Europe: estimating the medieval North African male legacy in southern Europe | journal = European Journal of Human Genetics | volume = 17 | issue = 6 | pages = 848–852 | date = June 2009 | pmid = 19156170 | pmc = 2947089 | doi = 10.1038/ejhg.2008.258 }} Considering both some E-M78 subhaplogroups and the E-M81 haplogroup, the contribution of northern African lineages to the entire male gene pool of Iberia (barring Pasiegos), continental Italy, and Sicily can be estimated as 5.6%, 3.6%, and 6.6%, respectively.{{cite journal | vauthors = Cruciani F, La Fratta R, Trombetta B, Santolamazza P, Sellitto D, Colomb EB, Dugoujon JM, Crivellaro F, Benincasa T, Pascone R, Moral P, Watson E, Melegh B, Barbujani G, Fuselli S, Vona G, Zagradisnik B, Assum G, Brdicka R, Kozlov AI, Efremov GD, Coppa A, Novelletto A, Scozzari R | display-authors = 6 | title = Tracing past human male movements in northern/eastern Africa and western Eurasia: new clues from Y-chromosomal haplogroups E-M78 and J-M12 | journal = Molecular Biology and Evolution | volume = 24 | issue = 6 | pages = 1300–1311 | date = June 2007 | pmid = 17351267 | doi = 10.1093/molbev/msm049 | doi-access = free }} Y-DNA lineages E-V12 and E-V22 have been associated with a Levantine source (represented by modern Lebanese), while North African haplogroup E-M81 shows an average frequency of 1.53% in the current Sicilian and Southern Italian genetic pool, but the typical Maghrebin core haplotype 13-14-30-24-9-11-13 has been found in only two out of the five E-M81 individuals. These results, along with the negligible contribution from North-African populations revealed by the admixture-like plot analysis, suggest only a marginal impact of trans-Mediterranean gene flows on the current Sicilian and Southern Italian genetic pool.{{cite journal | vauthors = Sarno S, Boattini A, Carta M, Ferri G, Alù M, Yao DY, Ciani G, Pettener D, Luiselli D | display-authors = 6 | title = An ancient Mediterranean melting pot: investigating the uniparental genetic structure and population history of sicily and southern Italy | journal = PLOS ONE | volume = 9 | issue = 4 | pages = e96074 | date = 2014-04-30 | pmid = 24788788 | pmc = 4005757 | doi = 10.1371/journal.pone.0096074 | doi-access = free | bibcode = 2014PLoSO...996074S }}

Auton et al. (2009) found that South West Europe had the highest proportion in Europe of haplotypes that are shared with sub-Saharan Africa (represented by Yoruba), and significantly more relative to South East Europe.{{cite journal |vauthors=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 |date=May 2009 |title=Global distribution of genomic diversity underscores rich complex history of continental human populations |journal=Genome Research |volume=19 |issue=5 |pages=795–803 |doi=10.1101/gr.088898.108 |pmc=2675968 |pmid=19218534}}

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. 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.{{cite journal | vauthors = 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 | title = Reconstructing ancient mitochondrial DNA links between Africa and Europe | journal = Genome Research | volume = 22 | issue = 5 | pages = 821–6 | date = May 2012 | pmid = 22454235 | pmc = 3337428 | doi = 10.1101/gr.134452.111 }}

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).{{cite journal | vauthors = Pereira L, Cunha C, Alves C, Amorim A | title = African female heritage in Iberia: a reassessment of mtDNA lineage distribution in present times | journal = Human Biology | volume = 77 | issue = 2 | pages = 213–29 | date = April 2005 | pmid = 16201138 | doi = 10.1353/hub.2005.0041| hdl = 10216/109268 | s2cid = 20901589 | hdl-access = free }} 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.{{cite journal | vauthors = Brehm A, Pereira L, Kivisild T, Amorim A | title = Mitochondrial portraits of the Madeira and Açores archipelagos witness different genetic pools of its settlers | journal = Human Genetics | volume = 114 | issue = 1 | pages = 77–86 | date = December 2003 | pmid = 14513360 | doi = 10.1007/s00439-003-1024-3 | s2cid = 8870699| hdl = 10400.13/3046 | hdl-access = free }} In the Canary Islands, frequencies have been reported at 6.6%. Regarding Iberia, current debates are concerned with whether these lineages are associated with prehistoric migrations, the Islamic occupation of Iberia, or the slave trade. African lineages in Iberia were predominantly the result of the Atlantic slave trade.{{cite journal | vauthors = Pereira L, Prata MJ, Amorim A | s2cid = 10478774 | title = Diversity of mtDNA lineages in Portugal: not a genetic edge of European variation | journal = Annals of Human Genetics | volume = 64 | issue = Pt 6 | pages = 491–506 | date = November 2000 | pmid = 11281213 | doi = 10.1046/j.1469-1809.2000.6460491.x| doi-access = free }} Most of the L lineages in Iberia matched Northwest African L lineages rather than contemporary Sub-Saharan L lineages.{{cite journal | vauthors = González AM, Brehm A, Pérez JA, Maca-Meyer N, Flores C, Cabrera VM | title = Mitochondrial DNA affinities at the Atlantic fringe of Europe | journal = American Journal of Physical Anthropology | volume = 120 | issue = 4 | pages = 391–404 | date = April 2003 | pmid = 12627534 | doi = 10.1002/ajpa.10168}} This pattern indicates that most of the Sub-Saharan L lineages entered Iberia in prehistoric times rather than during the slave trade. According to Sub-Saharan lineages found in Iberia matched lineages from diverse regions in Africa. This pattern is more compatible with a recent arrival of these lineages after slave trading began in the 15th century. Alternative scenarios that invoke much older and demographically more significant introductions have been proposed or a substantial role of the Roman and/or Islamic periods on the introduction of Sub-Saharan lineages seem unlikely. 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.{{cite journal | vauthors = Casas MJ, Hagelberg E, Fregel R, Larruga JM, González AM | title = Human mitochondrial DNA diversity in an archaeological site in al-Andalus: genetic impact of migrations from North Africa in medieval Spain | journal = American Journal of Physical Anthropology | volume = 131 | issue = 4 | pages = 539–51 | date = December 2006 | pmid = 16685727 | doi = 10.1002/ajpa.20463}} 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 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 in the comarca of Sayago (18.2%) which is "comparable to that described for the South of Portugal".{{cite journal | vauthors = Alvarez L, Santos C, Ramos A, Pratdesaba R, Francalacci P, Aluja MP | title = Mitochondrial DNA patterns in the Iberian Northern plateau: population dynamics and substructure of the Zamora province | journal = American Journal of Physical Anthropology | volume = 142 | issue = 4 | pages = 531–539 | date = August 2010 | pmid = 20127843 | doi = 10.1002/ajpa.21252 | quote = As regards sub-Saharan Hgs (L1b, L2b, and L3b), the high frequency found in the southern regions of Zamora, 18.2% in Sayago and 8.1% in Bajo Duero, is comparable to that described for the South of Portugal }}

In Italy, haplogroup L lineages are present at lower frequencies than in Iberia and are detected only in certain regions: Latium, Volterra,{{cite journal | vauthors = 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 | title = Mitochondrial DNA variation of modern Tuscans supports the near eastern origin of Etruscans | journal = American Journal of Human Genetics | volume = 80 | issue = 4 | pages = 759–68 | date = April 2007 | pmid = 17357081 | pmc = 1852723 | doi = 10.1086/512822 }} Basilicata, and Sicily.{{cite journal | vauthors = Ottoni C, Martinez-Labarga C, Vitelli L, Scano G, Fabrini E, Contini I, Biondi G, Rickards O | title = Human mitochondrial DNA variation in Southern Italy | journal = Annals of Human Biology | volume = 36 | issue = 6 | pages = 785–811 | year = 2009 | pmid = 19852679 | doi = 10.3109/03014460903198509 | s2cid = 1788055 }}

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. The monophyletic clusters L1b and L3b have an estimated age no greater than 6,500 years. African L1b, L2a, L3b, L3d and M1 clades in Slavic populations have been identified at low frequencies.{{cite journal | vauthors = Malyarchuk BA, Derenko M, Perkova M, Grzybowski T, Vanecek T, Lazur J | title = Reconstructing the phylogeny of African mitochondrial DNA lineages in Slavs | journal = European Journal of Human Genetics | volume = 16 | issue = 9 | pages = 1091–6 | date = September 2008 | pmid = 18398433 | doi = 10.1038/ejhg.2008.70| doi-access = free }} L1b, L3b and L3d had matches with West African populations, indicating that these lineages probably entered Europe through Iberia. One lineage, L2a1a, found in Czechs and Slovaks, appeared to be much older, indicating that it may have entered Europe in prehistoric times. This clade is distinct from the branch of L2a1 called L2a1l2a that is found in individuals of Ashkenazi heritage from central and eastern Europe{{cite book |last=Brook |first=Kevin Alan |pages=77–78 |date=2022 |title=The Maternal Genetic Lineages of Ashkenazic Jews |publisher=Academic Studies Press | isbn=978-1644699843 | doi=10.2307/j.ctv33mgbcn |s2cid=254519342 }} and less frequently in non-Jewish Poles.{{cite journal | author1=Marta Mielnik-Sikorska | author2=Patrycja Daca | author3=Boris Malyarchuk | author4=Miroslava Derenko | author5=Katarzyna Skonieczna | author6=Maria Perkova | author7=Tadeusz Dobosz | author8=Tomasz Grzybowski | title = The history of Slavs inferred from complete mitochondrial genome sequences | journal = PLOS ONE | volume = 8 | issue = 1 | pages = e54360 | date = 14 January 2013 | pmid = 23342138 | doi = 10.1371/journal.pone.0054360 | pmc=3544712 | bibcode=2013PLoSO...854360M | doi-access = free }} L2a lineages are widespread throughout Africa; as a result, the origins of this lineage are uncertain.{{cite journal | vauthors = 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 | title = The matrilineal ancestry of Ashkenazi Jewry: portrait of a recent founder event | journal = American Journal of Human Genetics | volume = 78 | issue = 3 | pages = 487–97 | date = March 2006 | pmid = 16404693 | pmc = 1380291 | doi = 10.1086/500307}}

Haplogroup M1 is also found in Europe at low frequencies. Haplogroup M1 had a frequency of 0.3%. The origins of haplogroup M1 have yet to be conclusively established.

A prehistoric episode from the Early Holocene is likely to be a contributor to some of the U6 and L lingeages present in Mediterranean Europe.{{cite journal | vauthors = Hernández CL, Soares P, Dugoujon JM, Novelletto A, Rodríguez JN, Rito T, Oliveira M, Melhaoui M, Baali A, Pereira L, Calderón R | title = Early Holocenic and Historic mtDNA African Signatures in the Iberian Peninsula: The Andalusian Region as a Paradigm | journal = PLOS ONE | volume = 10 | issue = 10 | pages = e0139784 | year = 2015 | pmid = 26509580 | pmc = 4624789 | doi = 10.1371/journal.pone.0139784 | bibcode = 2015PLoSO..1039784H | doi-access = free }}

In an analysis which also contains an admixture data 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.

Haplogroup U6, to which a North African origin has been attributed, is largely distributed among Mozabites (28.2%) and Mauritanians (20%).{{cite journal | vauthors = Rando JC, Pinto F, González AM, Hernández M, Larruga JM, Cabrera VM, Bandelt HJ | title = Mitochondrial DNA analysis of northwest African populations reveals genetic exchanges with European, near-eastern, and sub-Saharan populations | journal = Annals of Human Genetics | volume = 62 | issue = Pt 6 | pages = 531–50 | date = November 1998 | pmid = 10363131 | doi = 10.1046/j.1469-1809.1998.6260531.x | s2cid = 2925153 | url = | doi-access = free }} In other northwest Africans, the frequency of U6 ranges from 4.2% in Tunisians to 8% in Moroccan Arabs. In Europe, U6 is most common in Spain and Portugal.{{cite journal | vauthors = Maca-Meyer N, González AM, Pestano J, Flores C, Larruga JM, Cabrera VM | title = Mitochondrial DNA transit between West Asia and North Africa inferred from U6 phylogeography | journal = BMC Genetics | volume = 4 | pages = 15 | date = October 2003 | pmid = 14563219 | pmc = 270091 | doi = 10.1186/1471-2156-4-15 | doi-access = free }}{{cite journal | vauthors = Plaza S, Calafell F, Helal A, Bouzerna N, Lefranc G, Bertranpetit J, Comas D | title = Joining the pillars of Hercules: mtDNA sequences show multidirectional gene flow in the western Mediterranean | journal = Annals of Human Genetics | volume = 67 | issue = Pt 4 | pages = 312–328 | date = July 2003 | pmid = 12914566 | doi = 10.1046/j.1469-1809.2003.00039.x | s2cid = 11201992 }}

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%.{{cite journal | vauthors = Calderón R, Ambrosio B, Guitard E, González-Martín A, Aresti U, Dugoujon JM | title = Genetic position of Andalusians from Huelva in relation to other European and North African populations: a study based on GM and KM allotypes | journal = Human Biology | volume = 78 | issue = 6 | pages = 663–79 | date = December 2006 | pmid = 17564246 | doi = 10.1353/hub.2007.0008 | s2cid = 38748780}} 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.{{cite journal | vauthors = Calderón R, Lodeiro R, Varela TA, Fariña J, Ambrosio B, Guitard E, González-Martín A, Dugoujon JM | title = GM and KM immunoglobulin allotypes in the Galician population: new insights into the peopling of the Iberian Peninsula | journal = BMC Genetics | volume = 8 | issue = 1 | pages = 37 | date = June 2007 | pmid = 17597520 | pmc = 1934380 | doi = 10.1186/1471-2156-8-37 | doi-access = free }} Values of around 4% have also been found in Huelva and in the Aran valley in the Pyrenees.{{cite journal | vauthors = Giraldo MP, Esteban E, Aluja MP, Nogués RM, Backés-Duró C, Dugoujon JM, Moral P | title = Gm and Km alleles in two Spanish Pyrenean populations (Andorra and Pallars Sobirà): a review of Gm variation in the Western Mediterranean basin | journal = Annals of Human Genetics | volume = 65 | issue = Pt 6 | pages = 537–48 | date = November 2001 | pmid = 11851984 | doi = 10.1046/j.1469-1809.2001.6560537.x | doi-access = free }} 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 slaves sold from Africa.

In Sicily the North African haplotype Gm 5*;1;17; ranges from 1.56% at Valledolmo to 5.5% at Alia.{{cite journal | vauthors = Cerutti N, Dugoujon JM, Guitard E, Rabino Massa E | title = Gm and Km immunoglobulin allotypes in Sicily | journal = Immunogenetics | volume = 55 | issue = 10 | pages = 674–81 | date = January 2004 | pmid = 14652700 | doi = 10.1007/s00251-003-0628-z | s2cid = 9663858}} 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.).

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 or more, 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 one hypothesis, 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†. The overpopulation in Northeast Africa contributed to the development of the Natufian adaptation, which resulted in agriculture becoming a new way of sustenance.

From an analysis of human remains from the Natufian culture, there is evidence of Sub-Saharan influences in the Natufian samples.{{cite journal | vauthors = Brace CL, Seguchi N, Quintyn CB, Fox SC, Nelson AR, Manolis SK, Qifeng P | title = The questionable contribution of the Neolithic and the Bronze Age to European craniofacial form | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 103 | issue = 1 | pages = 242–7 | date = January 2006 | pmid = 16371462 | pmc = 1325007 | doi = 10.1073/pnas.0509801102| bibcode = 2006PNAS..103..242B | doi-access = free }}

These influences would have been diluted by the interbreeding of the Neolithic farmers from the Near East are associated with the indigenous foragers in Europe. 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.{{cite journal | vauthors = Ricaut FX, Waelkens M | title = Cranial discrete traits in a Byzantine population and eastern Mediterranean population movements | journal = Human Biology | volume = 80 | issue = 5 | pages = 535–64 | date = October 2008 | pmid = 19341322 | doi = 10.3378/1534-6617-80.5.535 | s2cid = 25142338}}

According to an ancient DNA analyses on Natufian skeletal remains from present-day northern Israel, the Natufians in fact shared no evident genetic affinity to sub-Saharan Africans. It was not possible 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.Lazaridis 2016 Quote: "However, no affinity of Natufians to sub-Saharan Africans is evident in our genome-wide analysis, as present-day sub-Saharan Africans do not share more alleles with Natufians than with other ancient Eurasians (Extended Data Table 1)." 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%).{{cite bioRxiv | vauthors = Lazaridis I, Nadel D, Rollefson G, Merrett DC, Rohland N, Mallick S, Fernandes D, Novak M, Gamarra B, Sirak K, Connell S |display-authors= 6 |date=17 June 2016|title=The genetic structure of the world's first farmers|biorxiv=10.1101/059311}} -- [http://biorxiv.org/content/biorxiv/suppl/2016/06/16/059311.DC1/059311-1.pdf Table S6.1 - Y-chromosome haplogroups]{{cite journal | vauthors = Lazaridis I, Nadel D, Rollefson G, Merrett DC, Rohland N, Mallick S, Fernandes D, Novak M, Gamarra B, Sirak K, Connell S, Stewardson K, Harney E, Fu Q, Gonzalez-Fortes G, Jones ER, Roodenberg SA, Lengyel G, Bocquentin F, Gasparian B, Monge JM, Gregg M, Eshed V, Mizrahi AS, Meiklejohn C, Gerritsen F, Bejenaru L, Blüher M, Campbell A, Cavalleri G, Comas D, Froguel P, Gilbert E, Kerr SM, Kovacs P, Krause J, McGettigan D, Merrigan M, Merriwether DA, O'Reilly S, Richards MB, Semino O, Shamoon-Pour M, Stefanescu G, Stumvoll M, Tönjes A, Torroni A, Wilson JF, Yengo L, Hovhannisyan NA, Patterson N, Pinhasi R, Reich D | display-authors = 6 | title = Genomic insights into the origin of farming in the ancient Near East | journal = Nature | volume = 536 | issue = 7617 | pages = 419–424 | date = August 2016 | pmid = 27459054 | pmc = 5003663 | doi = 10.1038/nature19310 | bibcode = 2016Natur.536..419L }}, Supplementary Table 1. 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{{cite journal | vauthors = Lazaridis I, Nadel D, Rollefson G, Merrett DC, Rohland N, Mallick S, Fernandes D, Novak M, Gamarra B, Sirak K, Connell S, Stewardson K, Harney E, Fu Q, Gonzalez-Fortes G, Jones ER, Roodenberg SA, Lengyel G, Bocquentin F, Gasparian B, Monge JM, Gregg M, Eshed V, Mizrahi AS, Meiklejohn C, Gerritsen F, Bejenaru L, Blüher M, Campbell A, Cavalleri G, Comas D, Froguel P, Gilbert E, Kerr SM, Kovacs P, Krause J, McGettigan D, Merrigan M, Merriwether DA, O'Reilly S, Richards MB, Semino O, Shamoon-Pour M, Stefanescu G, Stumvoll M, Tönjes A, Torroni A, Wilson JF, Yengo L, Hovhannisyan NA, Patterson N, Pinhasi R, Reich D | display-authors = 6 | title = Genomic insights into the origin of farming in the ancient Near East | journal = Nature | volume = 536 | issue = 7617 | pages = 419–424 | date = August 2016 | pmid = 27459054 | pmc = 5003663 | doi = 10.1038/nature19310 | bibcode = 2016Natur.536..419L }} from Neolithic Iranian farmers from the Zagros Mountains, caring up to 62% of the Basal Eurasians and Ancient North Eurasians (ANE). This might suggest that different strains of Basal Eurasians contributed to Natufians and Zagros farmers,{{cite journal | vauthors = Broushaki F, Thomas MG, Link V, López S, van Dorp L, Kirsanow K, Hofmanová Z, Diekmann Y, Cassidy LM, Díez-Del-Molino D, Kousathanas A, Sell C, Robson HK, Martiniano R, Blöcher J, Scheu A, Kreutzer S, Bollongino R, Bobo D, Davudi H, Munoz O, Currat M, Abdi K, Biglari F, Craig OE, Bradley DG, Shennan S, Veeramah K, Mashkour M, Wegmann D, Hellenthal G, Burger J | display-authors = 6 | title = Early Neolithic genomes from the eastern Fertile Crescent | journal = Science | volume = 353 | issue = 6298 | pages = 499–503 | date = July 2016 | pmid = 27417496 | pmc = 5113750 | doi = 10.1126/science.aaf7943 | bibcode = 2016Sci...353..499B }}{{cite journal | vauthors = Gallego-Llorente M, Connell S, Jones ER, Merrett DC, Jeon Y, Eriksson A, Siska V, Gamba C, Meiklejohn C, Beyer R, Jeon S, Cho YS, Hofreiter M, Bhak J, Manica A, Pinhasi R | display-authors = 6 | title = The genetics of an early Neolithic pastoralist from the Zagros, Iran | journal = Scientific Reports | volume = 6 | pages = 31326 | date = August 2016 | pmid = 27502179 | pmc = 4977546 | doi = 10.1038/srep31326 | bibcode = 2016NatSR...631326G }}{{cite journal | vauthors = Fernández E, Pérez-Pérez A, Gamba C, Prats E, Cuesta P, Anfruns J, Molist M, Arroyo-Pardo E, Turbón D | display-authors = 6 | title = Ancient DNA analysis of 8000 B.C. near eastern farmers supports an early neolithic pioneer maritime colonization of Mainland Europe through Cyprus and the Aegean Islands | journal = PLOS Genetics | volume = 10 | issue = 6 | pages = e1004401 | date = June 2014 | pmid = 24901650 | pmc = 4046922 | doi = 10.1371/journal.pgen.1004401 | doi-access = free }} 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.{{cite book | vauthors = Goring-Morris AN, Hovers E, Belfer-Cohen A | date = 2009 | chapter = The Dynamics of Pleistocene and Early Holocene Settlement Patterns in the Levant: An Overview. | title = Transitions in Prehistory: Essays in Honor of Ofer Bar-Yosef | veditors = Shea JJ, Lieberman DE | publisher = Oxbow Books | isbn = 978-1-84217-340-4 }} 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.{{cite journal | vauthors = Olszewski DI | year = 2006 | title = Issues in the Levantine Epipaleolithic: The Madamaghan, Nebekian and Qalkhan (Levant Epipaleolithic) | journal = Paléorient | volume = 32 | issue = 1| pages = 19–26 | doi=10.3406/paleo.2006.5168}} 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.{{cite journal | vauthors = Richter T, Garrard AN, Allock S, Maher LA | year = 2011 | title = Interaction before Agriculture: Exchanging Material and Sharing Knowledge in the Final Pleistocene Levant | journal = Cambridge Archaeological Journal | volume = 21 | issue = 1| pages = 95–114 | doi = 10.1017/S0959774311000060 | s2cid = 162887983 | url = http://discovery.ucl.ac.uk/1343637/1/download20.pdf }}

{{Portal|Evolutionary biology|Europe}}

• African immigration to Europe (contemporary history)
• Genetic history of the Middle East
• Genetic history of North Africa
• Genetic history of Europe
• Barbary slave trade
• Abkhazians of African descent

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• {{cite journal | vauthors = Cavalli-Sforza LL | title = Genes, peoples, and languages | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 94 | issue = 15 | pages = 7719–24 | date = July 1997 | pmid = 9223254 | pmc = 33682 | doi = 10.1073/pnas.94.15.7719| bibcode = 1997PNAS...94.7719C | doi-access = free }}
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{{human genetics}}

Europe, African Admixture In