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The ancestry of modern Iberians (Spanish and Portuguese) is consistent with the geographical situation of the Iberian Peninsula in the south-west of Europe:There is a strong connexion particularly with Atlantic Europe but also Mediterranean Europe and the Near East, albeit the latter is lesser than other Mediterranean countries (notably Greece) due to Spain being the furthest away from the Bosphorous region considered the main bridge of Neolithic expansions into Europe. Nevertheless, Iberia has the strongest link in Europe to North Africa, purportedly as a result of northward population movements during the seven centuries of Muslim rule over the peninsula. Iberia's Mediterranean and Near Eastern connexion is also likely to be largely of historical rather prehistorical origin, with the region's prehistorical ancestral basis being fundamentally western European.

Caveats

Archaeogenetics is a new science and most of its findings are recent and remain controversial as the understanding of the issues faced expands rapidly. For instance, the issue concerning the dating of the original R1 haplogroup mutation and its subclades has varied widely, originally dated at over 30 000 years BP (Before Present) to the later studies that show a much younger date of 18 600 BP. Such differences in dating affect conclusions about whether or not a population is largely palaeolithic or neolithic in origin.

Other problems relating to starting assumptions and the propensity of haplogroups to suffer genetic drift, population bottlenecks and founder effects point to the many difficulties faced by researchers and will require much time and effort to resolve with the help of a better understanding of the haplogroup clades, complex statistical analysis of extensive autosomally based studies correlated with archaeological research.

Analysis of prehistoric populations

2014 Study

A 2014 genetic study revealed that a Stone age man from the Mesolithic period, who lived in the Iberian Peninsula about 7,000 years ago, had blue eyes, black or brown hair, and dark skin. His DNA proved to be different from the genetic make up of most today's european populations.

The basques

The distinctiveness already noted by studies of 'classical markers', the subsequent discovery of frequency maxima of R1b in Basque populations, and the apparently "pre-Indo-European" nature of the Basque language resulted in a popular and long-held view that Basques are "living fossils" of the earliest modern humans who colonized Europe. Specifically, Y haplogroup R1b was deemed to be a Palaeolithic marker, highest in Basques, from when western Europe was repopulated after the Last Glacial Maximum, maybe 25,000 years ago.

The distinctiveness already noted by studies of 'classical markers', the subsequent discovery of frequency maxima of R1b in Basque populations, and the apparently "pre-Indo-European" nature of the Basque language resulted in a popular and long-held view that Basques are "living fossils" of the earliest modern humans who colonized Europe. Specifically, Y haplogroup R1b was deemed to be a Palaeolithic marker, highest in Basques, from when western Europe was repopulated after the Last Glacial Maximum, maybe 25,000 years ago.

Finally, several ancient DNA samples have been recovered and amplified from Palaeolithic sites in the Basque region. The collection of mtDNA haplogroups sampled there differed significantly compared to their modern frequencies. The authors concluded that there is "discontinuity" between ancient and modern Basques.

Main genetic compositions

General position among European populations

According to Dupanloup et al. (2004) the main components in the European genomes appear to derive from ancestors whose features were similar to those of modern Basques and Near Easterners, with average values greater than 35% for both these parental populations, regardless of whether or not molecular information is taken into account. The lowest degree of both Basque and Near Eastern admixture is found in Finland, whereas the highest values are, respectively, 70% ("Basque") in Spain and roughly 60% ("Near Eastern") in the Balkans.

DNA analysis

DNA analysis shows that the Spanish and Portuguese are most closely related to other populations of western Europe: the French, the Andorrans, the Italians, the Irish, the British, the Germans, and the Swiss.

2007 study

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.

2010 study

A controversial study, published in January 2010, claimed that Europeans (including Spaniards and Portuguese) are of a nearly equal combination of the European hunter gatherers of the earlier Paleolithic period and more recent Neolithic period ancestry from the Near East, that likely came from Anatolia. After initially focusing upon E1b1b as a Neolithic marker, the study looked at the male Y haplogroup R1b1b, which is much more common in Western Europe. Mark Jobling said: "We focused on the commonest Y-chromosome lineage in Europe, carried by about 110 million men, it follows a gradient from south-east to north-west, reaching almost 100% frequency in Ireland. We looked at how the lineage is distributed, how diverse it is in different parts of Europe, and how old it is."

The results suggested that the lineage R1b1a2 (R-M269) (from 2003 to 2005 what is now R1b1a2 was designated R1b3; from 2005 to 2008 it was R1b1c and from 2008 to 2011 it was R1b1b2), like E1b1b or J lineages, spread together with farming from the Near East. Dr Patricia Balaresque added: "In total, this means that more than 80% of European Y chromosomes descend from incoming farmers. In contrast, most maternal genetic lineages seem to descend from hunter-gatherers. To us, this suggests a reproductive advantage for farming males over indigenous hunter-gatherer males during the switch from hunting and gathering, to farming".

Previous Y-chromosome and mtDNA analysis had suggested Paleolithic ancestry among populations in the Iberian Peninsula and that Iberia may have played a role in the re-population of western Europe after the last glaciation. This shows an ancestral bond between Iberia and the rest of western Europe, and in particular with Atlantic Europe, which share high frequencies of these haplogroups. R1b1a2, the most common western European haplogroup, arose 4,000 to 8,000 years ago in southwest Asia and later spread to Europe.

However, a study by Laura Morelli et al. found the assumptions and conclusions flawed and demonstrated that cultural, not demic, diffusion was the main means by which agriculture was spread in Europe in the Neolithic period.

Autosomal studies

Haplogroup composition of the ancient Iberians was very similar to that found in the modern Iberian Peninsula populations, suggesting a long-term genetic continuity since pre-Roman times.

Links with African populations

North-African admixture

An earlier European-wide study had pointed to a small North African and Arab element in modern day Iberian ancestry when compared to the pre-Islamic ancestral basis, and the Gibraltar Strait seems to have functioned much more as a genetic barrier than a bridge.

A 2007 study (published 2007) had estimated the contribution of northern African lineages to the entire male gene pool of Iberia as 5.6%."

A number of studies focus on the genetic impact of the centuries of Muslim rule in the Iberian peninsula on the genetic make up of the modern Iberian population. Iberia has the greatest presence of the typically Northwest African Y-chromosome haplotype marker E-M81 in Europe. and Haplotype Va. However it should be stated the other clades of the Haplogroup E are also present in other European populations, Italy and the balkans have the greatest presence of E-M78 and E-V12 respectively, which is overall higher than E-M81 admixture in Iberia

In January 2009, a study by Capelli et al. Spanish individuals found the total contribution of specific North African male haplotypes in Spain as 7.7%.

As for Mtdna analysis (Mitochondrial DNA), although present at only low levels, Iberia has much higher frequencies of typically North African Haplogroup U6 than those generally observed in Europe. It is difficult to ascertain whether U6's presence is the consequence of Islam's expansion into Europe during the Middle Ages, or it is rather the result of ancient demic processes that predate the Islamic presence.

According to a widely publicized recent study (December 2008) published in the American Journal of Human Genetics, 19.8 percent of modern Iberians (Spain + Portugal) have DNA reflecting some minor Levantine (Sephardic Jewish) ancestry and 10.6 percent having DNA reflecting minor Moorish ancestry. The study's speculation on Sephardic origin of the Levantine markers has been widely contested since it is likely that such markers also reflect other migrations from the Near East such as what medieval Andalusians called "Syrians", earlier Phoenician colonization or even population movements during the Neolithic.

On the other hand, Chris Tyler-Smith, a collaborator with the team that carried the study, argues that the individual differences in Y-chromosome markers suggest that Iberians and Sephardic Jews must share ancestry more recent than several millennia, even though in also a recent study (October 2008) they attributed those same lineages in Iberia and the Balearic Islands as of Phoenician origin.

Haplogroup L lineages frequencies

Haplogroup L is the dominant haplogroup of Sub-Saharan Africa (96-100%). Presence of the L Haplogroup (mainly found all over Africa, specially Sub-Saharan; the Arabic Peninsula and the Middle East) in Iberia is minor but slightly more abundant than in the rest of European populations; it matches with the geographic position of the peninsula and the historic migrations that have taken place in it. However most of the L lineages in Iberia match Northwest African L lineages rather than contemporary Sub-Saharan L lineages, so it's category as Sub-Saharan is debatable.

• A 2011 study by Moorjani et al. found that almost all Southern Europeans and some central and nordic Europeans have inherited 1%–3% Sub-Saharan ancestry (3.2% in Portugal, 2.9% in Sardinia, 2.7% in Southern Italy, 2.4% in South-western Spain, 0.8% in North-eastern Spain, 1% in England, 1.4% in Norway and 1.1% in Northern Italy) with an average mixture date of around 55 generations ago, "consistent with North African gene flow at the end of the Roman Empire and subsequent Arab migrations".

In Iberia the mean frequency of Haplogroup L lineages reaches 3.83% and the frequency is higher in Portugal (5.83%) than in Spain (2.4%). Furthermore, in western Iberia, increasing frequencies are observed for Galicia (3.26%) and northern Portugal (3.21%), through the center (5.02%) and to the south of Portugal (11.38%).

According to a study in 2012 by Cerezo et al., about 65% of the European L lineages most likely arrived in rather recent historical times (Romanization period, Arab conquest of the Iberian Peninsula and Sicily, Atlantic slave trade) and about 35% of L mtDNAs form European-specific subclades, revealing that there was gene flow from sub-Saharan Africa toward Europe as early as 11,000 yr ago.

In Iberia the mean frequency of Haplogroup L lineages reaches 3.83% and the frequency is higher in Portugal (5.83%) than in Spain (2.4% average) slightly above the european average. In both countries frequencies vary widely between regions with increase frequencies observed for , Southern Portugal and Cordoba (Southern Spain). Regarding Iberia, current debates are concerned with whether these lineages are associated with prehistoric migrations, the Islamic occupation of Iberia or the slave trade. Pereira et al. (2000) harvcoltxt error: no target: CITEREFPereira_et_al.2000 (help) suggested that African lineages in Iberia were predominantly the result of Atlantic Slave Trade, especially in Portugal, where slave trade was considered one of the most important economic activies of the country. Gonzalez et al. (2003) harvcoltxt error: no target: CITEREFGonzalez_et_al.2003 (help) however revealed that most of the L lineages in Iberia matched Northwest African L lineages rather than contemporary Sub-Saharan L lineages. The authors suggest this pattern indicates that most of the Sub-Saharan L lineages entered Iberia (and the rest of Europe) in prehistoric times rather than during the slave trade.

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

See also

• Genetic history of Europe

References

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40. "La cifra de los sefardíes puede estar sobreestimada, ya que en estos genes hay mucha diversidad y quizá absorbieron otros genes de Oriente Medio" ("The Sephardic result may be overestimated, since there is much diversity in those genes and maybe absorbed other genes from the Middle East"). ¿Pone en duda Calafell la validez de los tests de ancestros? “Están bien para los americanos, nosotros ya sabemos de dónde venimos” (Puts Calafell in doubt the validity of ancestry tests? "They can be good for the Americans, we already know from where we come from)"
41. "El doctor Calafell matiza que (...) los marcadores genéticos usados para distinguir a la población con ancestros sefardíes pueden producir distorsiones". "ese 20% de españoles que el estudio señala como descendientes de sefardíes podrían haber heredado ese rasgo de movimiento más antiguos, como el de los fenicios o, incluso, primeros pobladores neolíticos hace miles de años." "Dr. Calafell clarifies that (...) the genetic markers used to distinguish the population with Sephardim ancestry may produce distortions. The 20% of Spaniards that are identified as having Sephardim ancestry in the study could have inherited that same marker from older movements like the Phoenicians, or even the first Neolithic settlers thousands of years ago" http://www.elmundo.es/elmundo/2008/12/04/ciencia/1228409780.html
43. 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. {{cite journal}}: Unknown parameter |laydate= ignored (help); Unknown parameter |laysource= ignored (help); Unknown parameter |laysummary= ignored (help)
45. http://forwhattheywereweare.blogspot.com/2012/03/claim-that-many-african-matrilineages.html
46. http://www.plosgenetics.org/article/info%3Adoi%2F10.1371%2Fjournal.pgen.1001373
47. Gonzalez et al. (2003) harvcoltxt error: no target: CITEREFGonzalez_et_al.2003 (help)
48. PLOS ONE: Uniparental Markers of Contemporary Italian Population Reveals Details on Its Pre-Roman Heritage
49. Different genetic components in the Norwegian population revealed by the analysis of mtDNA and Y chromosome polymorphisms
50. Moorjani P; Patterson N; Hirschhorn JN; Keinan A; Hao L; et al. (2011). McVean, Gil (ed.). "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. {{cite journal}}: Unknown parameter |author-separator= ignored (help)CS1 maint: unflagged free DOI (link)
51. 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. PMID 16201138. {{cite journal}}: Invalid |ref=harv (help)
52. Cerezo M; Achilli A; Olivieri A; et al. (May 2012). "Reconstructing ancient mitochondrial DNA links between Africa and Europe". Genome Res. 22 (5): 821–6. doi:10.1101/gr.134452.111. PMC 3337428. PMID 22454235. {{cite journal}}: Unknown parameter |author-separator= ignored (help)
53. 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. PMID 16201138. {{cite journal}}: Invalid |ref=harv (help)CS1 maint: multiple names: authors list (link)
54. Auton A; Bryc K; Boyko AR; et al. (May 2009). "Global distribution of genomic diversity underscores rich complex history of continental human populations". Genome Research. 19 (5): 795–803. doi:10.1101/gr.088898.108. PMC 2675968. PMID 19218534. {{cite journal}}: Invalid |ref=harv (help); Unknown parameter |author-separator= ignored (help)

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