Genetic history of the Iberian Peninsula

PCA plot of 17 contemporary Iberian populations[1]

The ancestry of modern Iberians (comprising the Spanish and Portuguese) is consistent with the geographical situation of the Iberian Peninsula in the South-west corner of Europe, showing characteristics that are largely typical in Southern and Western Europeans. As is the case for most of the rest of Southern Europe, the principal ancestral origin of modern Iberians are Early European Farmers who arrived during the Neolithic. The large predominance of Y-Chromosome Haplogroup R1b, common throughout Western Europe, is also testimony to a sizeable input from various waves of (predominantly male) Western Steppe Herders that originated in the Pontic-Caspian Steppe during the Bronze Age.[2][3]

Modern Iberians' genetic inheritance largely derives from the pre-Roman inhabitants of the Iberian Peninsula who were deeply Romanized after the conquest of the region by the ancient Romans:[4][5]

There are also some genetic influences from the Alans and Germanic tribes who arrived after the Roman period, including the Suebi, Hasdingi Vandals, and Visigoths.[10][11][12] Due to its position on the Mediterranean Sea, like other Southern European countries, there were also contacts with other Mediterranean peoples such as the Phoenicians, Ancient Greeks and Carthaginians who briefly settled along Iberia's eastern and southern coasts, the Sephardi Jewish community, and Berbers and Arabs arrived during Al-Andalus, all of them leaving some North African and Middle Eastern genetic influences, particularly in the south and west of the Iberian Peninsula.[13][14][5][15][16][17][4] Similar to Sardinia, Iberia was shielded from settlement from the Middle East and Caucasus region by its western geographic location, and thus has lower levels of Western Asian and Middle Eastern admixture than Italy and Greece, most of which probably arrived to Iberia during historic rather than prehistoric times, especially in the Roman period.[18][19]

Population Genetics: Methods and Limitations

The foremost pioneer of the study of population genetics was Luigi Luca Cavalli-Sforza. Cavalli-Sforza used classical genetic markers to analyse DNA by proxy. This method studies differences in the frequencies of particular allelic traits, namely polymorphisms from proteins found within human blood (such as the ABO blood groups, Rhesus blood antigens, HLA loci, immunoglobulins, G-6-P-D isoenzymes, among others). Subsequently, his team calculated genetic distances between populations, based on the principle that two populations that share similar frequencies of a trait are more closely related than populations that have more divergent frequencies of the trait.[20]

Since then, population genetics has progressed significantly and studies using direct DNA analysis are now abundant and may use mitochondrial DNA (mtDNA), the non-recombining portion of the Y chromosome (NRY) or autosomal DNA. MtDNA and NRY DNA share some similar features which have made them particularly useful in genetic anthropology. These properties include the direct, unaltered inheritance of mtDNA and NRY DNA from mother to offspring and father to son, respectively, without the 'scrambling' effects of genetic recombination. We also presume that these genetic loci are not affected by natural selection and that the major process responsible for changes in base pairs has been mutation (which can be calculated).[21]

Whereas Y-DNA and mtDNA haplogroups represent but a small component of a person's DNA pool, autosomal DNA has the advantage of containing hundreds and thousands of examinable genetic loci, thus giving a more complete picture of genetic composition. Descent relationships can only to be determined on a statistical basis, because autosomal DNA undergoes recombination. A single chromosome can record a history for each gene. Autosomal studies are much more reliable for showing the relationships between existing populations but do not offer the possibilities for unraveling their histories in the same way as mtDNA and NRY DNA studies promise, despite their many complications.[citation needed]

Analyses of nuclear and ancient DNA

(A) Geographic distribution of the inferred proportions in the map. Shadings for proportions are scaled according to the maximum and minimum proportions of each source. (B) Representation of the ADMIXTURE proportions in each target population based on the statistically significant models obtained with qpAdm.

Nuclear DNA analysis shows that Spanish and Portuguese populations are most closely related to other populations of western Europe.[22][23][24] There is an axis of significant genetic differentiation along the east–west direction, in contrast to remarkable genetic similarity in the north–south direction. North African admixture, associated with the Islamic conquest, can be dated to the period between c. AD 860–1120.[25]

A study published in 2019 using samples of 271 Iberians spanning prehistoric and historic times proposes the following inflexion points in Iberian genomic history:[26]

  1. Mesolithic: hunter-gatherers from the European Steppes of Western Russia, Georgia and Ukraine are the first humans to settle the northwest of the Iberian Peninsula.
  2. Neolithic: neolithic farmers settle the entire Iberian Peninsula from Anatolia.
  3. Chalcolithic: Inflow of Central European hunter-gatherers and some gene inflow from sporadic contact with North Africa.
  4. Bronze Age: Steppe inflow from Central Europe.
  5. Iron Age: Additional Steppe gene flow from Central Europe, - the genetic pool of the Basque people remains mostly intact from this point on.
  6. Roman period: genetic inflow from Central and Eastern Mediterranean. Some additional inflow of North African genes detected in Southern Iberia.
  7. Visigothic period: no detectable inflows.
  8. Muslim period: Inflow from Northern Africa. Following the Reconquista, there is further genetic convergence between North and South Iberia.

North African influence

Main article: African admixture in Europe
Distribution of North African Admixture in the Iberian Peninsula

A number of studies have focused on ascertaining the genetic impact of historical North African population movements into Iberia on the genetic composition of modern Spanish and Portuguese populations. Initial studies pointed to the Straits of Gibraltar acting more as a genetic barrier than a bridge during prehistorical times,[27][28][29] while other studies point to a higher level of recent North African admixture among Iberians than among other European populations,[30][31][32][33][34][35][36] albeit this is as a result of more recent migratory movements, particularly the Moorish invasion of Iberia in the 8th century.

In terms of autosomal DNA, the most recent study regarding African admixture in Iberian populations was conducted in April 2013 by Botigué et al. using genome-wide SNP data for over 2000 European, Maghreb, Qatar and Sub-Saharan individuals of which 119 were Spaniards and 117 Portuguese, concluding that Spain and Portugal hold significant levels of North African ancestry. Estimates of shared ancestry averaged from 4% in some places to 10% in the general population; the populations of the Canary Islands yielded from 0% to 96% of shared ancestry with north Africans, although the Canary islands are a Spanish exclave located in the African continent, and thus this output is not representative of the Iberian population; these same results did not exceed 2% in other western or southern European populations.[37][38][39][40] 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. Indeed, a prior 2011 autosomal study by Moorjani et al. found Sub-Saharan ancestry in many parts of southern Europe at ranges of between 1-3%, "the highest proportion of African ancestry in Europe is in Iberia (Portugal 4.2±0.3% and Spain 1.4±0.3%), 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 North Africans."[30][34][35]

Recent studies show minor relationships between some Iberian regions and North African populations as a result of the Al-Andalus historical period which in Portugal lasted between the 8th and 12th centuries AD, and in southern Spain continued until the late 15th century AD. Iberia is the European region that has a more prominent presence of haplogroup E3b of the human Y chromosome (E-M81),[41] of haplogroup U (U6) and Haplotype Va, and this may be the result of some original common western Mediterranean population. In Portugal, North African Y-chromosome haplogroups (especially those typically North-West African) are at a frequency of 7.1%.[42] Some studies of mitochondrial DNA also find evidence of the North African haplogroup U6, especially in northern Portugal.[43] Although the frequency of U6 is low (4–6%), it was estimated that approximately 27% of the population of northern Portugal had some North African ancestry, as U6 is also not a common lineage in North Africa.[44] According to some studies, the North African and Arab elements in the ancestry of today's Iberians are more than trivial when compared to the basis of pre-Islamic ancestry, and the Strait of Gibraltar seems to function more as a genetic bridge than a barrier.[45][46][47]

However, a study that has used different genetic markers has reached different conclusions. In an autosomal study by Spínola et al. (2005), which analyzed the human leukocyte antigen (HLA genes) (inherent in all ancestors in direct paternal and maternal lineages) in hundreds of individuals from Portugal, showed that the Portuguese population has been influenced by other Europeans and North Africans, via many ancient migrations. According to the authors, the North and the South of Portugal show a greater similarity towards North Africans as opposed to the people of the center of the country, who seem closer to other Europeans, since the North of Portugal seems to have concentrated, certainly due to the pressure of Arab expansion, an ancient genetic pole originating from many North Africans and other Europeans, influences through millennia, [clarification needed] while southern Portugal shows a North African genetic influence, probably the result of origins recent from the Amazigh people who accompanied the Arab expansion.[48]

Moura (literally "Moorish"), a small southern town near the Spanish border, known for its Moorish heritage

According to a study published in the American Journal of Human Genetics in December 2008, 30% of modern Portuguese (23.6% in the north and 36.3% in the south) have DNA that shows they have male Sephardic Jewish ancestry and 14% (11.8 in the North and 16.1% in the South) have Moorish ancestry.[49] Despite the possible alternative sources for lineages attributed to a Sephardic Jewish origin, these proportions were testimony to the importance of religious conversion (voluntary or forced), shown by historical episodes of social and religious intolerance.

Five component admixture plots for various contemporary Iberian populations against other European, West Asian, North African and West African populations[50]

In terms of paternal Y-Chromosome DNA, recent studies coincide in that Iberia has the greatest presence of the typically Northwest African Y-chromosome haplotype marker E-M81 in Europe, with an average of 3%.[31][32] as well as Haplotype Va.[51][33] Estimates of Y-Chromosome ancestry vary, with a 2008 study published in the American Journal of Human Genetics using 1140 samples from throughout the Iberian peninsula, giving a proportion of 10.6% North African ancestry[34][35][36] to the paternal composite of Iberians. A similar 2009 study of 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%).[31]

Other studies of the Iberian gene-pool have estimated significantly lower levels of North African Ancestry. According to Bosch et al. 2000 "NW African populations may have contributed 7% of Iberian Y chromosomes".[28] A wide-ranging study by Cruciani et al. 2007, using 6,501 unrelated Y-chromosome samples from 81 populations found that: "Considering both these E-M78 sub-haplogroups (E-V12, E-V22, E-V65) 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 percent, 3.6 percent and 6.6 percent, respectively".[52] A 2007 study estimated the contribution of northern African lineages to the entire male gene pool of Iberia as 5.6%."[53] In general aspects, according to (Bosch et al. 2007) "...the origins of the Iberian Y-chromosome pool may be summarized as follows: 5% recent NW African, 78% Upper Paleolithic and later local derivatives (group IX), and 10% Neolithic" (H58, H71).[54]

Mitochondrial DNA studies of 2003, coincide in that the Iberian Peninsula holds higher levels of typically North African Haplotype U6,[35][55] as well as higher frequencies of Sub-Saharan African Haplogroup L in Portugal.[56][57][58][59] High frequencies are largely concentrated in the south and southwest of the Iberian peninsula, therefore overall frequency is higher in Portugal (7.8%) than in Spain (1.9%) with a mean frequency for the entire peninsula of 3.8%. There is considerable geographic divergence across the peninsula with high frequencies observed for Western Andalusia (14.6%)[59] and Córdoba (8.3%).,[56] Southern Portugal (10.7%), South West Castile (8%). Adams et al. and other previous publications, propose that the Moorish occupation left a minor Jewish, Saqaliba[60] and some Arab-Berber genetic influence mainly in southern regions of Iberia.[61][34]

The most recent and comprehensive genomic studies establish that North African genetic ancestry can be identified throughout most of the Iberian Peninsula, ranging from 0% to 11%, but is highest in the south and west, while being absent or almost absent in the Basque Country and northeast.[62][18][19]

Current debates revolve around whether U6 presence is due to Islamic expansion into the Iberian peninsula or prior population movements[34][35][36] and whether Haplogroup L is linked to the slave trade or prior population movements linked to Islamic expansion. A majority of Haplogroup L lineages in Iberia being North African in origin points to the latter.[56][58][30][35][63] In 2015, Hernández et al. concluded that "the estimated entrance of the North African U6 lineages into Iberia at 10 ky correlates well with other L African clades, indicating that some U6 and L lineages moved together from Africa to Iberia in the Early Holocene while a majority were introduced during historic times."[64]

Haplogroups

Y-DNA haplogroup frequencies in the Iberian Peninsula[34]

Y-Chromosome haplogroups

Main article: Haplogroup R-DF27

Like other Western Europeans, among Spaniards and Portuguese the Y-DNA Haplogroup R1b is the most frequent, occurring at over 70% throughout most of Spain.[65] R1b is particularly dominant in the Basque Country and Catalonia, occurring at rate of over 80%. In Iberia, most men with R1b belong to the subclade R-P312 (R1b1a1a2a1a2; as of 2017). The distribution of haplogroups other than R1b varies widely from one region to another.

In Portugal as a whole the R1b haplogroups rate 70%, with some areas in the Northwest regions reaching over 90%.[66]

Although R1b prevails in much of Western Europe, a key difference is found in the prevalence in Iberia of R-DF27 (R1b1a1a2a1a2a). This subclade is found in over 60% of the male population in the Basque Country and 40-48% in Madrid, Alicante, Barcelona, Cantabria, Andalucia, Asturias and Galicia.[67] R-DF27 constitutes much more than the half of the total R1b in the Iberian Peninsula. Subsequent in-migration by members of other haplogroups and subclades of R1b did not affect its overall prevalence, although this falls to only two thirds of the total R1b in Valencia and the coast more generally.[65] R-DF27 is also a significant subclade of R1b in parts of France and Britain. R-S28/R-U152 (R1b1a1a2a1a2b) is the prevailing subclade of R1b in Northern Italy, Switzerland and parts of France, but it represents less than 5.0% of the male population in Iberia. Ancient samples from the central European Bell Beaker culture, Hallstatt culture and Tumulus culture belonged to this subclade.[68][69][70] R-S28/R-U152 is slightly significant in Seville, Barcelona, Portugal and Basque Country at 10-20% of the total population, but it is represented at frequencies of only 3.0% in Cantabria and Santander, 2.0% in Castille and Leon, 6% in Valencia, and under 1% in Andalusia.[65] Sephardic Jews I1 0% I2*/I2a 1% I2 0% Haplogroup R1a 5% R1b 13% G 15% Haplogroup J2 2 25% J*/J1 22% E-M2151b1b 9% T 6% Q 2% [71]

Haplogroup J, mostly subclades of Haplogroup J-M172 (J2), is found at levels of over 20% in some regions, while Haplogroup E has a general frequency of about 10% – albeit with peaks surpassing 30% in certain areas. Overall, E-M78 (E1b1b1a1 in 2017) and E-M81 (E1b1b1b1a in 2017) both constitute about 4.0% each, with a further 1.0% from Haplogroup E-M123 (E1b1b1b2a1) and 1.0% from unknown subclades of E-M96.[34] (E-M81 is widely considered to represent relatively historical migrations from North Africa).

Mitochondrial DNA

MtDNA haplogroup frequencies in the main Iberian regions[72]
Main articles: Haplogroup H (mtDNA) and Haplogroup U (mtDNA)

There have been a number of studies about the mitochondrial DNA haplogroups (mtDNA) in Europe. In contrast to Y DNA haplogroups, mtDNA haplogroups did not show as much geographical patterning, but were more evenly ubiquitous. Apart from the outlying Sami, all Europeans are characterized by the predominance of haplogroups H, U and T. The lack of observable geographic structuring of mtDNA may be due to socio-cultural factors, namely patrilocality and a lack of polyandry.[73]

The subhaplogroups H1 and H3 have been subject to a more detailed study and would be associated to the Magdalenian expansion from Iberia c. 13,000 years ago:[56]

  • H1 encompasses an important fraction of Western European mtDNA, reaching its local peak among contemporary Basques (27.8%) and appearing at a high frequency among other Iberians and North Africans. Its frequency is above 10% in many other parts of Europe (France, Sardinia, British Isles, Alps, large portions of Eastern Europe), and above 5% in nearly all the continent. Its subclade H1b is most common in eastern Europe and NW Siberia.[74] So far, the highest frequency of H1 - 61%- has been found among the Tuareg of the Fezzan region in Libya.[75]
  • H3 represents a smaller fraction of European genome than H1 but has a somewhat similar distribution with peak among Basques (13.9%), Galicians (8.3%) and Sardinians (8.5%). Its frequency decreases towards the northeast of the continent, though. Studies have suggested haplogroup H3 is highly protective against AIDS progression.[76]

A 2007 European-wide study including Spanish Basques and Valencian Spaniards found Iberian populations to cluster the furthest from other continental groups, implying that Iberia holds the most ancient European ancestry. In this study, the most prominent genetic stratification in Europe was found to run from the north to the south-east, while another important axis of differentiation runs east–west across the continent. It also found, despite the differences, that all Europeans are closely related.[77]

Subregions

Spain

Frequencies of Y-DNA haplogroups in Spanish regions[34][78]
Region Sample size C E G I J2 JxJ2 R1a R1b Notes
Aragon 34 6% 0% 18% 12% 0% 3% 56%
Andalusia East 95 4% 3% 6% 9% 3% 1% 72%
Andalusia West 73 15% 4% 5% 14% 1% 4% 54%
Asturias 20 15% 5% 10% 15% 0% 0% 50%
Basques 116 1% 0% 8% 3% 1% 0% 87%
Castilla La Mancha 63 4% 10% 2% 6% 2% 2% 72%
Castile North-East 31 9% 3% 3% 3% 0% 0% 77%
Castile North-West 100 19% 5% 3% 8% 1% 2% 60%
Catalonia 80 >0%[79] 3% 6% 3% 6% 0% 0% 81%
Extremadura 52 18% 4% 10% 12% 0% 0% 50%
Galicia 88 17% 6% 10% 7% 1% 0% 57%
Valencia 73 >0%[80] 10% 1% 10% 5% 3% 3% 64%
Majorca 62 9% 6% 8% 8% 2% 0% 66%
Menorca 37 19% 0% 3% 3% 0% 3% 73%
Ibiza 54 8% 13% 2% 4% 0% 0% 57%
Seville 155 7% 4% 12% 8% 3% 1% 60%
Huelva 22 14% 0% 9% 14% 0% 0% 59%
Cadiz 28 4% 0% 14% 14% 4% 0% 51%
Cordoba 27 11% 0% 15% 15% 0% 0% 56%
Málaga 26 31% 4% 0% 15% 0% 8% 43%
Leon 60 10% 7% 3% 5% 2% 7% 62%
Cantabria 70 13% 9% 6% 3% 3% 4% 58%

Portugal

Distribution of the R1b haplogroup in Europe

Excerpts from the Abstract of a study published[81] in 2015:

"[...] In the case of Portugal, previous population genetics studies have already revealed the general portrait of HVS-I and HVS-II mitochondrial diversity, becoming now important to update and expand the mitochondrial region analysed. Accordingly, a total of 292 complete control region sequences from continental Portugal were obtained, under a stringent experimental design to ensure the quality of data through double sequencing of each target region.* Furthermore, H-specific coding region SNPs were examined to detail haplogroup classification and complete mitogenomes were obtained for all sequences belonging to haplogroups U4 and U5. In general, a typical Western European haplogroup or Atlantic modal haplotype (AMH) composition was found in mainland Portugal, associated to high level of mitochondrial genetic diversity. Within the country, no signs of substructure were detected. The typing of extra coding region SNPs has provided the refinement or confirmation of the previous classification obtained with EMMA tool in 96% of the cases. Finally, it was also possible to enlarge haplogroup U phylogeny with 28 new U4 and U5 mitogenomes."

The AMH reaches the highest frequencies in the Iberian Peninsula, in Great Britain and Ireland. In the Iberian Peninsula it reaches 70% in Portugal as a whole, with more than 90% in NW Portugal and nearly 90% in Galicia (NW Spain), while the highest value is to be found among the Basques (NE Spain).

The Atlantic modal haplotype (AMH) or haplotype 15 is a Y chromosome haplotype of Y-STR microsatellite variations, associated with the Haplogroup R1b. It was discovered prior to many of the SNPs now used to identify subclades of R1b and references to it can be found in some of the older literature. It corresponds most closely with subclade R1b1a2a1a(1) [L11].

The AMH is the most frequently occurring haplotype amongst human males in Atlantic Europe. It is characterized by the following marker alleles:

  • DYS388 12
  • DYS390 24
  • DYS391 11
  • DYS392 13
  • DYS393 13
  • DYS394 14 (also known as DYS19)

See also

Wikimedia Commons has media related to Genetic studies on Iberian Peninsula.

References

  1. ^ Pimenta J, Lopes AM, Carracedo A, Arenas M, Amorim A, Comas D (2019). "Spatially explicit analysis reveals complex human genetic gradients in the Iberian Peninsula". Sci Rep. 9 (1): 7825. Bibcode:2019NatSR...9.7825P. doi:10.1038/s41598-019-44121-6. PMC 6534591. PMID 31127131.
  2. ^ Nelis, Mari; Esko, Tõnu; Mägi, Reedik; Zimprich, Fritz; Zimprich, Alexander; Toncheva, Draga; Karachanak, Sena; Piskáčková, Tereza; Balaščák, Ivan; Peltonen, Leena; Jakkula, Eveliina; Rehnström, Karola; Lathrop, Mark; Heath, Simon; Galan, Pilar; Schreiber, Stefan; Meitinger, Thomas; Pfeufer, Arne; Wichmann, H-Erich; Melegh, Béla; Polgár, Noémi; Toniolo, Daniela; Gasparini, Paolo; D'Adamo, Pio; Klovins, Janis; Nikitina-Zake, Liene; Kučinskas, Vaidutis; Kasnauskienė, Jūratė; Lubinski, Jan; Debniak, Tadeusz; Limborska, Svetlana; Khrunin, Andrey; Estivill, Xavier; Rabionet, Raquel; Marsal, Sara; Julià, Antonio; Antonarakis, Stylianos E.; Deutsch, Samuel; Borel, Christelle; Attar, Homa; Gagnebin, Maryline; Macek, Milan; Krawczak, Michael; Remm, Maido; Metspalu, Andres; Fleischer, Robert C. (8 May 2009). "Genetic Structure of Europeans: A View from the North–East". PLOS ONE. 4 (5): e5472. Bibcode:2009PLoSO...4.5472N. doi:10.1371/journal.pone.0005472. PMC 2675054. PMID 19424496.
  3. ^ Novembre, John; Johnson, Toby; Bryc, Katarzyna; Kutalik, Zoltán; Boyko, Adam R.; Auton, Adam; Indap, Amit; King, Karen S.; Bergmann, Sven; Nelson, Matthew R.; Stephens, Matthew; Bustamante, Carlos D. (31 August 2008). "Genes mirror geography within Europe". Nature. 456 (7218): 98–101. Bibcode:2008Natur.456...98N. doi:10.1038/nature07331. PMC 2735096. PMID 18758442.
  4. ^ a b Bycroft, Clare; Fernandez-Rozadilla, Ceres; Ruiz-Ponte, Clara; Quintela, Inés; Carracedo, Ángel; Donnelly, Peter; Myers, Simon (1 February 2019). "Patterns of genetic differentiation and the footprints of historical migrations in the Iberian Peninsula". Nature Communications. 10 (1): 551. Bibcode:2019NatCo..10..551B. doi:10.1038/s41467-018-08272-w. PMC 6358624. PMID 30710075.
  5. ^ a b Olalde, Iñigo; Mallick, Swapan; Patterson, Nick; Rohland, Nadin; Villalba-Mouco, Vanessa; Silva, Marina; Dulias, Katharina; Edwards, Ceiridwen J.; Gandini, Francesca; Pala, Maria; Soares, Pedro; Ferrando-Bernal, Manuel; Adamski, Nicole; Broomandkhoshbacht, Nasreen; Cheronet, Olivia; Culleton, Brendan J.; Fernandes, Daniel; Lawson, Ann Marie; Mah, Matthew; Oppenheimer, Jonas; Stewardson, Kristin; Zhang, Zhao; Arenas, Juan Manuel Jiménez; Moyano, Isidro Jorge Toro; Salazar-García, Domingo C.; Castanyer, Pere; Santos, Marta; Tremoleda, Joaquim; Lozano, Marina; et al. (15 March 2019). "The genomic history of the Iberian Peninsula over the past 8000 years". Science. 363 (6432): 1230–1234. Bibcode:2019Sci...363.1230O. doi:10.1126/science.aav4040. PMC 6436108. PMID 30872528.
  6. ^ "Iberians - MSN Encarta". Encarta.msn.com. Archived from the original on 30 October 2009. Retrieved 12 January 2022.
  7. ^ Álvarez-Sanchís, Jesús (28 February 2005). "Oppida and Celtic society in western Spain". E-Keltoi: Journal of Interdisciplinary Celtic Studies. 6 (1).
  8. ^ a b "Ethnographic Map of Pre-Roman Iberia (Circa 200 B.C.)". Arqueotavira.com. Archived from the original on 11 June 2004. Retrieved 12 January 2022.
  9. ^ "Spain - History". Britannica.com.
  10. ^ "Les Wisigoths dans le Portugal médiéval : état actuel de la question". L'Europe héritière de l'Espagne wisigothique. Collection de la Casa de Velázquez. Books.openedition.org. 23 January 2014. pp. 326–339. ISBN 9788490960981.
  11. ^ https://alpha.sib.uc.pt/?q=content/o-património-visigodo-da-l%C3%ADngua-portuguesa [dead link‍]
  12. ^ Quiroga, Jorge López (January 2017). "(PDF) IN TEMPORE SUEBORUM. The time of the Suevi in Gallaecia (411–585 AD)". Jorge López Quiroga-Artemio M. Martínez Tejera (Coord.): In Tempore Sueborum. The Time of the Sueves in Gallaecia (411–585 Ad). The First Medieval Kingdom of the West, Ourense. Academia.edu. Retrieved 21 January 2020.
  13. ^ James S. Amelang. "The Expulsion of the Moriscos: Still more Questions than Answers" (PDF). Intransitduke.org. Universidad Autónoma, Madrid. Retrieved 22 January 2022.
  14. ^ Jónsson 2007, p. 195.
  15. ^ Adams, Susan M.; Bosch, Elena; Balaresque, Patricia L.; Ballereau, Stéphane J.; Lee, Andrew C.; Arroyo, Eduardo; López-Parra, Ana M.; Aler, Mercedes; Grifo, Marina S. Gisbert; Brion, Maria; Carracedo, Angel; Lavinha, João; Martínez-Jarreta, Begoña; Quintana-Murci, Lluis; Picornell, Antònia; Ramon, Misericordia; Skorecki, Karl; Behar, Doron M.; Calafell, Francesc; Jobling, Mark A. (12 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): 725–736. doi:10.1016/j.ajhg.2008.11.007. PMC 2668061. PMID 19061982.
  16. ^ Torres, Gabriela (31 December 2008). "El español "puro" tiene de todo". BBC Mundo.
  17. ^ Cervantes virtual: La invasión árabe. Los árabes y el elemento árabe en español. 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". The American Journal of Human Genetics. 83 (6): 725–736. doi:10.1016/j.ajhg.2008.11.007. PMC 2668061. PMID 19061982.
  18. ^ a b Bycroft, Clare; et al. (2019). "Patterns of genetic differentiation and the footprints of historical migrations in the Iberian Peninsula". Nature Communications. 10 (1): 551. Bibcode:2019NatCo..10..551B. doi:10.1038/s41467-018-08272-w. PMC 6358624. PMID 30710075.
  19. ^ a b Olalde, Iñigo; et al. (2019). "The genomic history of the Iberian Peninsula over the past 8000 years". Science. 363 (6432): 1230–1234. Bibcode:2019Sci...363.1230O. doi:10.1126/science.aav4040. PMC 6436108. PMID 30872528.
  20. ^ Cavalli-Sforza LL, Menozzi P, Piazza A (1993). The History and Geography of Human Genes. Princeton University Press. p. 51. ISBN 978-0-691-08750-4. Retrieved 2009-07-22.
  21. ^ Milisauskas S (2002). European Prehistory: a survey. Birkhauser. p. 58. ISBN 978-0-306-46793-6.
  22. ^ Nelis, Mari; Esko, Tõnu; Mägi, Reedik; Zimprich, Fritz; Zimprich, Alexander; Toncheva, Draga; Karachanak, Sena; Piskácková, Tereza; Balascák, Ivan; Peltonen, L; Jakkula, E; Rehnström, K; Lathrop, M; Heath, S; Galan, P; Schreiber, S; Meitinger, T; Pfeufer, A; Wichmann, HE; Melegh, B; Polgár, N; Toniolo, D; Gasparini, P; d'Adamo, P; Klovins, J; Nikitina-Zake, L; Kucinskas, V; Kasnauskiene, J; Lubinski, J; Debniak, T (2009). Fleischer, Robert C. (ed.). "Genetic Structure of Europeans: A View from the North–East". PLOS ONE. 4 (5): e5472. Bibcode:2009PLoSO...4.5472N. doi:10.1371/journal.pone.0005472. PMC 2675054. PMID 19424496.
  23. ^ Wade, Nicholas (13 August 2008). "The Genetic Map of Europe". The New York Times. Retrieved 17 October 2009.
  24. ^ Novembre, John; Johnson, Toby; Bryc, Katarzyna; Kutalik, Zoltán; Boyko, Adam R.; Auton, Adam; Indap, Amit; King, Karen S.; Bergmann, Sven; Nelson, Matthew R.; Stephens, Matthew; Bustamante, Carlos D. (2008). "Genes mirror geography within Europe". Nature. 456 (7218): 98–101. Bibcode:2008Natur.456...98N. doi:10.1038/nature07331. PMC 2735096. PMID 18758442.
  25. ^ Bycroft, Clare; Fernandez-Rozadilla, Ceres; Ruiz-Ponte, Clara; Quintela, Inés; Carracedo, Ángel; Donnelly, Peter; Myers, Simon (1 February 2019). "Patterns of genetic differentiation and the footprints of historical migrations in the Iberian Peninsula". Nature Communications. 10 (1): 551. Bibcode:2019NatCo..10..551B. doi:10.1038/s41467-018-08272-w. PMC 6358624. PMID 30710075.
  26. ^ Olalde, Iñigo; Mallick, Swapan; Patterson, Nick; Rohland, Nadin; Villalba-Mouco, Vanessa; Silva, Marina; Dulias, Katharina; Edwards, Ceiridwen J.; Gandini, Francesca; Pala, Maria; Soares, Pedro (2019-03-15). "The genomic history of the Iberian Peninsula over the past 8000 years". Science. 363 (6432): 1230–1234. Bibcode:2019Sci...363.1230O. doi:10.1126/science.aav4040. ISSN 0036-8075. PMC 6436108. PMID 30872528.
  27. ^ Dupanloup, I.; Bertorelle, G; Chikhi, L; Barbujani, G (2004). "Estimating the Impact of Prehistoric Admixture on the Genome of Europeans". Molecular Biology and Evolution. 21 (7): 1361–72. doi:10.1093/molbev/msh135. PMID 15044595. S2CID 17665038.
  28. ^ a b Bosch, Elena; Calafell, Francesc; Comas, David; Oefner, Peter J.; Underhill, Peter A.; Bertranpetit, Jaume (2001). "High-Resolution Analysis of Human Y-Chromosome Variation Shows a Sharp Discontinuity and Limited Gene Flow between Northwestern Africa and the Iberian Peninsula". The American Journal of Human Genetics. 68 (4): 1019–29. doi:10.1086/319521. PMC 1275654. PMID 11254456.
  29. ^ Comas, David; Calafell, Francesc; Benchemsi, kiNoufissa; Helal, Ahmed; Lefranc, Gerard; Stoneking, Mark; Batzer, Mark A.; Bertranpetit, Jaume; Sajantila, Antti (2000). "Alu insertion polymorphisms in NW Africa and the Iberian Peninsula: Evidence for a strong genetic boundary through the Gibraltar Straits". Human Genetics. 107 (4): 312–9. doi:10.1007/s004390000370. PMID 11129330. S2CID 9618737.
  30. ^ a b c Moorjani P, Patterson N, Hirschhorn JN, Keinan A, Hao L, Atzmon G, et al. (2011). "The History of African Gene Flow into Southern Europeans, Levantines, and Jews". PLOS Genet. 7 (4): e1001373. doi:10.1371/journal.pgen.1001373. PMC 3080861. PMID 21533020.
  31. ^ a b c Capelli, Cristian; Onofri, Valerio; Brisighelli, Francesca; Boschi, Ilaria; Scarnicci, Francesca; Masullo, Mara; Ferri, Gianmarco; Tofanelli, Sergio; Tagliabracci, Adriano; Gusmao, Leonor; Amorim, Antonio; Gatto, Francesco; Kirin, Mirna; Merlitti, Davide; Brion, Maria; Verea, Alejandro Blanco; Romano, Valentino; Cali, Francesco; Pascali, Vincenzo (2009). "Moors and Saracens in Europe: estimating the medieval North African male legacy in southern Europe". European Journal of Human Genetics. 17 (6): 848–52. doi:10.1038/ejhg.2008.258. PMC 2947089. PMID 19156170.
  32. ^ a b Semino, Ornella; Magri, Chiara; Benuzzi, Giorgia; Lin, Alice A.; Al-Zahery, Nadia; Battaglia, Vincenza; MacCioni, Liliana; Triantaphyllidis, Costas; Shen, Peidong; Oefner, Peter J.; Zhivotovsky, Lev A.; King, Roy; Torroni, Antonio; Cavalli-Sforza, L. Luca; Underhill, Peter A.; Santachiara-Benerecetti, A. Silvana (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". The American Journal of Human Genetics. 74 (5): 1023–34. doi:10.1086/386295. PMC 1181965. PMID 15069642.
  33. ^ a b Gérard, Nathalie; Berriche, Sala; Aouizérate, Annie; Diéterlen, Florent; Lucotte, Gérard (2006). "North African Berber and Arab Influences in the Western Mediterranean Revealed by Y-Chromosome DNA Haplotypes". Human Biology. 78 (3): 307–16. doi:10.1353/hub.2006.0045. PMID 17216803. S2CID 13347549.
  34. ^ a b c d e f g h Adams, Susan M.; Bosch, Elena; Balaresque, Patricia L.; Ballereau, Stéphane J.; Lee, Andrew C.; Arroyo, Eduardo; López-Parra, Ana M.; Aler, Mercedes; Grifo, Marina S. Gisbert; Brion, Maria; Carracedo, Angel; Lavinha, João; Martínez-Jarreta, Begoña; Quintana-Murci, Lluis; Picornell, Antònia; Ramon, Misericordia; Skorecki, Karl; Behar, Doron M.; Calafell, Francesc; Jobling, Mark A. (2008). "The Genetic Legacy of Religious Diversity and Intolerance: Paternal Lineages of Christians, Jews, and Muslims in the Iberian Peninsula". The American Journal of Human Genetics. 83 (6): 725–36. doi:10.1016/j.ajhg.2008.11.007. PMC 2668061. PMID 19061982.
  35. ^ a b c d e f González, Ana M.; Brehm, Antonio; Pérez, José A.; Maca-Meyer, Nicole; Flores, Carlos; Cabrera, Vicente M. (April 2003). "Mitochondrial DNA affinities at the Atlantic fringe of Europe". American Journal of Physical Anthropology. 120 (4): 391–404. doi:10.1002/ajpa.10168. PMID 12627534. S2CID 6430969.
  36. ^ a b c Giacomo, F.; Luca, F.; Popa, L. O.; Akar, N.; Anagnou, N.; Banyko, J.; Brdicka, R.; Barbujani, G.; et al. (2004). "Y chromosomal haplogroup J as a signature of the post-neolithic colonization of Europe". Human Genetics. 115 (5): 357–371. doi:10.1007/s00439-004-1168-9. PMID 15322918. S2CID 18482536.
  37. ^ Botigué, L. R.; Henn, B. M.; Gravel, S; Maples, B. K.; Gignoux, C. R.; Corona, E; Atzmon, G; Burns, E; Ostrer, H; Flores, C; Bertranpetit, J; Comas, D; Bustamante, C. D. (2013). "Gene flow from North Africa contributes to differential human genetic diversity in southern Europe". Proceedings of the National Academy of Sciences. 110 (29): 11791–11796. Bibcode:2013PNAS..11011791B. doi:10.1073/pnas.1306223110. PMC 3718088. PMID 23733930.
  38. ^ Estimating gene flow from North Africa to southern Europe Archived 2015-04-30 at archive.today, David Comas, one of the authors of the study
  39. ^ "La cifra del 20% sólo se da en Canarias, para el resto del país oscila entre el 10% y 12%", explica Comas.", David Comas, one of the authors of the study, Los españoles somos los europeos con más genes magrebíes, Huffington post, June 2013
  40. ^ Flores, Carlos; Villar, Jesús; Guerra, Luisa; Hernández, Alexis; Basaldúa, Santiago; Corrales, Almudena; Pino-Yanes, María (2011-03-30). "North African Influences and Potential Bias in Case-Control Association Studies in the Spanish Population". PLOS ONE. 6 (3): e18389. Bibcode:2011PLoSO...618389P. doi:10.1371/journal.pone.0018389. ISSN 1932-6203. PMC 3068190. PMID 21479138.
  41. ^ Gérard, Nathalie; Berriche, Sala; Aouizérate, Annie; Diéterlen, Florent; Lucotte, Gérard (2006). "North African Berber and Arab Influences in the Western Mediterranean Revealed by Y-Chromosome DNA Haplotypes". Human Biology. 78 (3): 307–316. doi:10.1353/hub.2006.0045. PMID 17216803. S2CID 13347549.
  42. ^ Gonçalves, Rita; Freitas, Ana; Branco, Marta; Rosa, Alexandra; Fernandes, Ana T.; Zhivotovsky, Lev A.; Underhill, Peter A.; Kivisild, Toomas; Brehm, António (2005). "Y-chromosome Lineages from Portugal, Madeira and Açores Record Elements of Sephardim and Berber Ancestry". Annals of Human Genetics. 69 (4): 443–454. doi:10.1111/j.1529-8817.2005.00161.x. hdl:10400.13/3018. PMID 15996172. S2CID 3229760.
  43. ^ 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:10.1353/hub.2005.0041. hdl:10216/109268. PMID 16201138. S2CID 20901589.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  44. ^ 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. Vol. 120, no. 4. pp. 391–404. doi:10.1002/ajpa.10168. PMID 12627534.{{cite news}}: CS1 maint: multiple names: authors list (link)
  45. ^ Dupanloup I, Bertorelle G, Chikhi L, Barbujani G (July 2004). "Estimating the impact of prehistoric admixture on the genome of Europeans". Molecular Biology and Evolution. Vol. 21, no. 7. pp. 1361–72. doi:10.1093/molbev/msh135. PMID 15044595.{{cite news}}: CS1 maint: multiple names: authors list (link)
  46. ^ Bosch E, Calafell F, Comas D, Oefner PJ, Underhill PA, Bertranpetit J (April 2001). "High-resolution analysis of human Y-chromosome variation shows a sharp discontinuity and limited gene flow between northwestern Africa and the Iberian Peninsula". American Journal of Human Genetics. Vol. 68, no. 4. pp. 1019–29. doi:10.1086/319521. PMC 1275654. PMID 11254456.{{cite news}}: CS1 maint: multiple names: authors list (link)
  47. ^ Comas, David; Calafell, Francesc; Benchemsi, Noufissa; Helal, Ahmed; Lefranc, Gerard; Stoneking, Mark; Batzer, Mark A.; Bertranpetit, Jaume; Sajantila, Antti (2000). "Alu insertion polymorphisms in NW Africa and the Iberian Peninsula: evidence for a strong genetic boundary through the Gibraltar Straits". Human Genetics. 107 (4): 312–319. doi:10.1007/s004390000370. PMID 11129330. S2CID 9618737.
  48. ^ Spínola, H.; Middleton, D.; Brehm, A. (2005). "HLA genes in Portugal inferred from sequence-based typing: in the crossroad between Europe and Africa". Tissue Antigens. 66 (1): 26–36. doi:10.1111/j.1399-0039.2005.00430.x. PMID 15982254.
  49. ^ "The Genetic Legacy of Religious Diversity and Intolerance: Paternal Lineages of Christians, Jews, and Muslims in the Iberian Peninsula".
  50. ^ Hernández CL, Pita G, Cavadas B, López S, Sánchez-Martínez LJ, Dugoujon JM, et al. (2020). "Human Genomic Diversity Where the Mediterranean Joins the Atlantic". Mol Biol Evol. 37 (4): 1041–1055. doi:10.1093/molbev/msz288. PMC 7086172. PMID 31816048.
  51. ^ Lucotte, Gérard; Gérard, Nathalie; Mercier, Géraldine (2011-04-05). "North African Genes in Iberia Studied by Y-Chromosome DNA Haplotype 5". Human Biology. 73 (5).
  52. ^ Cruciani, F.; La Fratta, R.; Trombetta, B.; Santolamazza, P.; Sellitto, D.; Colomb, E. B.; Dugoujon, J. -M.; Crivellaro, F.; Benincasa, T. (2007). "Tracing Past Human Male Movements in Northern/Eastern Africa and Western Eurasia: New Clues from Y-Chromosomal Haplogroups E-M78 and J-M12". Molecular Biology and Evolution. 24 (6): 1300–1311. doi:10.1093/molbev/msm049. PMID 17351267.
  53. ^ Cruciani, F.; La Fratta, R.; Trombetta, B.; Santolamazza, P.; Sellitto, D.; Colomb, E. B.; Dugoujon, J.-M.; 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, A. I.; Efremov, G. D.; Coppa, A.; Novelletto, A.; Scozzari, R. (2007). "Tracing Past Human Male Movements in Northern/Eastern Africa and Western Eurasia: New Clues from Y-Chromosomal Haplogroups E-M78 and J-M12". Molecular Biology and Evolution. 24 (6): 1300–11. doi:10.1093/molbev/msm049. PMID 17351267.
  54. ^ Bosch, E; Calafell, F; Comas, D; Oefner, PJ; Underhill, PA; Bertranpetit, J (April 2001). "High-resolution analysis of human Y-chromosome variation shows a sharp discontinuity and limited gene flow between northwestern Africa and the Iberian Peninsula". Am. J. Hum. Genet. 68 (4): 1019–29. doi:10.1086/319521. PMC 1275654. PMID 11254456.
  55. ^ Plaza, S.; Calafell, F.; Helal, A.; Bouzerna, N.; Lefranc, G.; Bertranpetit, J.; Comas, D. (2003). "Joining the Pillars of Hercules: mtDNA Sequences Show Multidirectional Gene Flow in the Western Mediterranean". Annals of Human Genetics. 67 (Pt 4): 312–28. doi:10.1046/j.1469-1809.2003.00039.x. PMID 12914566. S2CID 11201992.
  56. ^ a b c d Pereira, Luisa; Cunha, Carla; Alves, Cintia; Amorim, Antonio (2005). "African Female Heritage in Iberia: A Reassessment of mtDNA Lineage Distribution in Present Times". Human Biology. 77 (2): 213–29. doi:10.1353/hub.2005.0041. hdl:10216/109268. PMID 16201138. S2CID 20901589.
  57. ^ Cerezo, M.; Achilli, A.; Olivieri, A.; Perego, U. A.; Gomez-Carballa, A.; Brisighelli, F.; Lancioni, H.; Woodward, S. R.; Lopez-Soto, M.; Carracedo, A.; Capelli, C.; Torroni, A.; Salas, A. (27 March 2012). "Reconstructing ancient mitochondrial DNA links between Africa and Europe". Genome Research. 22 (5): 821–826. doi:10.1101/gr.134452.111. PMC 3337428. PMID 22454235.
  58. ^ 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:10.1007/s00439-003-1024-3. hdl:10400.13/3046. PMID 14513360. S2CID 8870699.
  59. ^ a b Hernández, Candela L; Reales, Guillermo; Dugoujon, Jean-Michel; Novelletto, Andrea; Rodríguez, Juan; Cuesta, Pedro; Calderón, Rosario (2014). "Human maternal heritage in Andalusia (Spain): its composition reveals high internal complexity and distinctive influences of mtDNA haplogroups U6 and L in the western and eastern side of region". BMC Genetics. 15 (1): 11. doi:10.1186/1471-2156-15-11. PMC 3905667. PMID 24460736.
  60. ^ Gordon, Matthew; Hain, Kathryn A. (2017). Concubines and Courtesans: Women and Slavery in Islamic History. Oxford University Press. ISBN 9780190622183.
  61. ^ "Tracing Past Human Male Movements in Northern/Eastern Africa and Western Eurasia". Academic.oup.com. Retrieved 2020-01-21.
  62. ^ https://reich.hms.harvard.edu/sites/reich.hms.harvard.edu/files/inline-files/2019_Olalde_Science_IberiaTransect_2.pdf [bare URL PDF]
  63. ^ Alvarez, Luis; Santos, Cristina; Ramos, Amanda; Pratdesaba, Roser; Francalacci, Paolo; Aluja, María Pilar (1 February 2010). "Mitochondrial DNA patterns in the Iberian Northern plateau: Population dynamics and substructure of the Zamora province". American Journal of Physical Anthropology. 142 (4): 531–539. doi:10.1002/ajpa.21252. PMID 20127843.
  64. ^ Hernández, Candela L.; Soares, Pedro; Dugoujon, Jean M.; Novelletto, Andrea; Rodríguez, Juan N.; Rito, Teresa; Oliveira, Marisa; Melhaoui, Mohammed; Baali, Abdellatif; Pereira, Luisa; Calderón, Rosario; Achilli, Alessandro (28 October 2015). "Early Holocenic and Historic mtDNA African Signatures in the Iberian Peninsula: The Andalusian Region as a Paradigm". PLOS ONE. 10 (10): e0139784. Bibcode:2015PLoSO..1039784H. doi:10.1371/journal.pone.0139784. PMC 4624789. PMID 26509580.
  65. ^ a b c Myres, Natalie M; Rootsi, Siiri; Lin, Alice A; Järve, Mari; King, Roy J; Kutuev, Ildus; Cabrera, Vicente M; Khusnutdinova, Elza K; Pshenichnov, Andrey; Yunusbayev, Bayazit; Balanovsky, Oleg; Balanovska, Elena; Rudan, Pavao; Baldovic, Marian; Herrera, Rene J; Chiaroni, Jacques; Di Cristofaro, Julie; Villems, Richard; Kivisild, Toomas; Underhill, Peter A (25 August 2010). "A major Y-chromosome haplogroup R1b Holocene era founder effect in Central and Western Europe". European Journal of Human Genetics. 19 (1): 95–101. doi:10.1038/ejhg.2010.146. PMC 3039512. PMID 20736979.
  66. ^ Marques, Sofia L.; Goios, Ana; Rocha, Ana M.; Prata, Maria João; Amorim, António; Gusmão, Leonor; Alves, Cíntia; Alvarez, Luis (March 2015). "Portuguese mitochondrial DNA genetic diversity—An update and a phylogenetic revision". Forensic Science International: Genetics. 15: 27–32. doi:10.1016/j.fsigen.2014.10.004. PMID 25457629.
  67. ^ Valverde, Laura; Illescas, Maria José; Villaescusa, Patricia; Gotor, Amparo M.; García, Ainara; Cardoso, Sergio; Algorta, Jaime; Catarino, Susana; Rouault, Karen; Férec, Claude; Hardiman, Orla; Zarrabeitia, Maite; Jiménez, Susana; Pinheiro, Maria Fátima; Jarreta, Begoña M.; Olofsson, Jill; Morling, Niels; de Pancorbo, Marian M. (March 2016). "New clues to the evolutionary history of the main European paternal lineage M269: dissection of the Y-SNP S116 in Atlantic Europe and Iberia". European Journal of Human Genetics. 24 (3): 437–441. doi:10.1038/ejhg.2015.114. PMC 4755366. PMID 26081640.
  68. ^ Olalde, Iñigo; Brace, Selina; Allentoft, Morten E.; Armit, Ian; Kristiansen, Kristian; Booth, Thomas; Rohland, Nadin; Mallick, Swapan; Szécsényi-Nagy, Anna; Mittnik, Alissa; Altena, Eveline; Lipson, Mark; Lazaridis, Iosif; Harper, Thomas K.; Patterson, Nick; Broomandkhoshbacht, Nasreen; Diekmann, Yoan; Faltyskova, Zuzana; Fernandes, Daniel; Ferry, Matthew; Harney, Eadaoin; de Knijff, Peter; Michel, Megan; Oppenheimer, Jonas; Stewardson, Kristin; Barclay, Alistair; Alt, Kurt Werner; Liesau, Corina; Ríos, Patricia; et al. (March 2018). "The Beaker phenomenon and the genomic transformation of northwest Europe". Nature. 555 (7695): 190–196. Bibcode:2018Natur.555..190O. doi:10.1038/nature25738. PMC 5973796. PMID 29466337.
  69. ^ Allentoft, Morten E.; Sikora, Martin; Sjögren, Karl-Göran; Rasmussen, Simon; Rasmussen, Morten; Stenderup, Jesper; Damgaard, Peter B.; Schroeder, Hannes; Ahlström, Torbjörn; Vinner, Lasse; Malaspinas, Anna-Sapfo; Margaryan, Ashot; Higham, Tom; Chivall, David; Lynnerup, Niels; Harvig, Lise; Baron, Justyna; Casa, Philippe Della; Dąbrowski, Paweł; Duffy, Paul R.; Ebel, Alexander V.; Epimakhov, Andrey; Frei, Karin; Furmanek, Mirosław; Gralak, Tomasz; Gromov, Andrey; Gronkiewicz, Stanisław; Grupe, Gisela; Hajdu, Tamás; Jarysz, Radosław; Khartanovich, Valeri; Khokhlov, Alexandr; Kiss, Viktória; Kolář, Jan; Kriiska, Aivar; Lasak, Irena; Longhi, Cristina; McGlynn, George; Merkevicius, Algimantas; Merkyte, Inga; Metspalu, Mait; Mkrtchyan, Ruzan; Moiseyev, Vyacheslav; Paja, László; Pálfi, György; Pokutta, Dalia; Pospieszny, Łukasz; Price, T. Douglas; Saag, Lehti; Sablin, Mikhail; Shishlina, Natalia; Smrčka, Václav; Soenov, Vasilii I.; Szeverényi, Vajk; Tóth, Gusztáv; Trifanova, Synaru V.; Varul, Liivi; Vicze, Magdolna; Yepiskoposyan, Levon; Zhitenev, Vladislav; Orlando, Ludovic; Sicheritz-Pontén, Thomas; Brunak, Søren; Nielsen, Rasmus; Kristiansen, Kristian; Willerslev, Eske (June 2015). "Population genomics of Bronze Age Eurasia". Nature. 522 (7555): 167–172. Bibcode:2015Natur.522..167A. doi:10.1038/nature14507. PMID 26062507. S2CID 4399103.
  70. ^ Peter de Barros, Damgaard (9 May 2018). "137 ancient human genomes from across the Eurasian steppes". Nature. 557 (7705): 369–374. Bibcode:2018Natur.557..369D. doi:10.1038/s41586-018-0094-2. hdl:1887/3202709. PMID 29743675. S2CID 13670282.
  71. ^ "Eupedia".
  72. ^ Barral-Arca R, Pischedda S, Gómez-Carballa A, Pastoriza A, Mosquera-Miguel A, López-Soto M, et al. (2016). "Meta-Analysis of Mitochondrial DNA Variation in the Iberian Peninsula". PLOS ONE. 11 (7): e0159735. Bibcode:2016PLoSO..1159735B. doi:10.1371/journal.pone.0159735. PMC 4956223. PMID 27441366.
  73. ^ Rosser et al. (2000)
  74. ^ Loogväli EL, Roostalu U, Malyarchuk BA, et al. (November 2004). "Disuniting uniformity: a pied cladistic canvas of mtDNA haplogroup H in Eurasia". Molecular Biology and Evolution. 21 (11): 2012–21. doi:10.1093/molbev/msh209. PMID 15254257.
  75. ^ Ottoni, Claudio; Primativo, Giuseppina; Hooshiar Kashani, Baharak; Achilli, Alessandro; Martínez-Labarga, Cristina; Biondi, Gianfranco; Torroni, Antonio; Rickards, Olga; Kayser, Manfred (21 October 2010). "Mitochondrial Haplogroup H1 in North Africa: An Early Holocene Arrival from Iberia". PLOS ONE. 5 (10): e13378. Bibcode:2010PLoSO...513378O. doi:10.1371/journal.pone.0013378. PMC 2958834. PMID 20975840.
  76. ^ Hendrickson SL, Hutcheson HB, Ruiz-Pesini E, et al. (November 2008). "Mitochondrial DNA Haplogroups influence AIDS Progression". AIDS. 22 (18): 2429–39. doi:10.1097/QAD.0b013e32831940bb. PMC 2699618. PMID 19005266.
  77. ^ Bauchet, M; McEvoy, B; Pearson, LN; Quillen, EE; Sarkisian, T; Hovhannesyan, K; Deka, R; Bradley, DG; Shriver, MD (2007). "Measuring European Population Stratification with Microarray Genotype Data". The American Journal of Human Genetics. 80 (5): 948–56. doi:10.1086/513477. PMC 1852743. PMID 17436249.
  78. ^ Flores, Carlos; Maca-Meyer, Nicole; González, Ana M.; Oefner, Peter J.; Shen, Peidong; Pérez, Jose A.; Rojas, Antonio; Larruga, Jose M.; Underhill, Peter A. (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): 855–863. doi:10.1038/sj.ejhg.5201225. PMID 15280900.
  79. ^ Haplogroup C* (C-M130) has been found among males with the surname Llach and originating from Garrotxa, Catalonia, Spain. It was not found among males with the same surname from other areas, or males with other surnames of Catalan origin (Cognoms Catalans, n.d., Resultat; access 15 September 2015). The Cognoms Catalans project, which researches "genetic surnames" in Catalonia, Valencia and the Balearic Islands, is based at Universitat Pompeu Fabra, Barcelona.
  80. ^ C Haplogroup – Y-DNA Classic Chart (21 January 2017).
  81. ^ Marques, Sofia L; Goios, Ana; Rocha, Ana M; Prata, Maria J; Amorim, António; Gusmão, Leonor; Alves, Cíntia; Alvarez, Luis (March 2015). "Portuguese mitochondrial DNA genetic diversity- an update and a phylogenetic revision". FSI Genetics. 15: 27–32. doi:10.1016/j.fsigen.2014.10.004. PMID 25457629.

Works cited