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Revision as of 16:30, 1 July 2014 editJeezuuz1934 (talk | contribs)73 edits Barracuda, you have still deleted a lot of studies. Its ok to shorten but not to censor what you dislike, see talk page.Tag: Incorrectly formatted external link or image← Previous edit Revision as of 16:44, 1 July 2014 edit undo83.55.46.126 (talk) →Haplogroup L lineages frequencies: Deleting section on Eastern Europe as per BarracudaNext edit →
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In ], Haplogroup L lineages are present at lower frequencies than ], and only detected in certain regions, such as between 2 and 3% in ], ],<ref>{{cite paper | pmc = 1852723 | title = Mitochondrial DNA Variation of Modern Tuscans Supports the Near Eastern Origin of Etruscans | author = Achilli et al. | year = 2008 }}</ref> ] and ].<ref>{{cite paper | pmid = 19852679 | title = Human mitochondrial DNA variation in Southern Italy | author = Ottoni et al. | year = 2009 }}</ref> In ], Haplogroup L lineages are present at lower frequencies than ], and only detected in certain regions, such as between 2 and 3% in ], ],<ref>{{cite paper | pmc = 1852723 | title = Mitochondrial DNA Variation of Modern Tuscans Supports the Near Eastern Origin of Etruscans | author = Achilli et al. | year = 2008 }}</ref> ] and ].<ref>{{cite paper | pmid = 19852679 | title = Human mitochondrial DNA variation in Southern Italy | author = Ottoni et al. | year = 2009 }}</ref>

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 ] clusters. {{Harvcoltxt|Malyarchuk et al.|2005}} detected only two monophyletic clusters, L1b and L3b in Russians with an estimated age no greater than 6,500 years. {{Harvcoltxt|Malyarchuk et al.|2008}} identified African L1b, L2a, L3b, L3d and M1 clades in Slavic populations at low frequencies. L1b, L3b and L3d had matches with West African populations, indicating that these lineages probably entered Europe through Iberia. One lineage L2a1a, appeared to be much older, indicating that it may have entered Europe in prehistoric times. This clade was possibly related to L2a1 clades identified in ten individuals of ] heritage from France, Germany, Poland, Romania and Russia. L2a lineages are widespread throughout Africa, as a result the origins of this lineage are uncertain.<ref name="malyarchuk2008">{{Harvcoltxt|Malyarchuk et al.|2008}}</ref><ref>{{cite journal |author=Behar DM |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 |ref=harv |author-separator=, |author2=Metspalu E |author3=Kivisild T |display-authors=3 |last4=Achilli |first4=A |last5=Hadid |first5=Y |last6=Tzur |first6=S |last7=Pereira |first7=L |last8=Amorim |first8=A |last9=Quintanamurci |first9=L}}</ref>


] is also found in Europe at low frequencies. In a study by Gonzalez et al. 2007, haplogroup M1 had a frequency of 0.3%. The origins of haplogroup M1 have yet to be conclusively established. One clade of haplgroup M1, M1a is widely accepted to be of East African origin. About 40% of all clades of M1 found in Europe are M1a and consequently of recent East African origin.{{citation needed|date=May 2013}} ] is also found in Europe at low frequencies. In a study by Gonzalez et al. 2007, haplogroup M1 had a frequency of 0.3%. The origins of haplogroup M1 have yet to be conclusively established. One clade of haplgroup M1, M1a is widely accepted to be of East African origin. About 40% of all clades of M1 found in Europe are M1a and consequently of recent East African origin.{{citation needed|date=May 2013}}

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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 both North African and Sub Saharan 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.

Distribution of R1a (purple) and R1b (red). (Black represents all other haplogroups)

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.

Main genetic compositions

File:Haplogroups europe.png
Y-haplogroups in Europe. Iberia is together with France, England, North Italy and Switzerland mainly R1b.

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

A number of studies have tried to find out the genetic impact of North African and Middle Eastern population movements on the modern Spanish and Portuguese ancestry, through comparison of genetic markers in Spain and Portugal with North Africa and the Near East. The most recent and thorough study about Moorish influence in the Iberian Peninsula was conducted April 2013 by Pompeu Fabra University using genome-wide SNP data for over 2000 individuals. This study concluded that the Iberian Peninsula holds significantly higher levels of both North African and Sub-Saharan ancestry than the rest of the European continent with a sharp difference between Iberia and France. Estimates of shared ancestry with North African populations were found to be much higher than previously reported, accounting for up to 20% of individual genomes, whereas these do not exceed 2% in Southern European populations outside Spain and Portugal and are practically inexistent north of the Pyrenees.

Iberia has the greatest presence of the typically Northwest African Y-chromosome haplotype marker E-M81 in Europe. and Haplotype Va.

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 Near Eastern and 10.6 percent having DNA reflecting North African ancestors. 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, including Levantine Arabs, earlier Phoenician colonization or even population movements during the Neolithic. Chris Tyler-Smith, a collaborator with the team that carried the study, argues that the individual differences in Y-chromosome markers suggests a common ancestry more recent than several millennia.

Priorly, Capelli et al. 2009 reported that North African male haplogroups, especially E1b1b1b (E-M81), E1b1b1a-b (M78 derived chromosomes showing the rare DYS439 allele 10, or E-V65) and a subset of J1 (M267 derived), represented, on average, 7-8% of the current Iberian male lineages.

Historically introduced NW African types in Iberia (Capelli et al. (2009))
Sample N E1b1b1b (M81) E1b1b1a-b (V65) J1 (subset) Total %
Spain 717 5.2 1 1.5 7.7
Portugal 659 5 0.3 1.8 7.1
Iberia 1376 5.1 0.7 1.7 7.4

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.

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.

Country Region Number tested Study %
Italy Countrywide 583 Brisighelli et al. (2012) 0.8%
Italy Countrywide 411 Plaza et al. (2003) 0.1%
Italy Sicily 169 Plaza et al. (2003) 0.6%
Italy Sicily 106 Maca-Meyer et al. (2003) 0.94%
Spain Central Spain 50 Plaza et al. (2003) 2.0%
Spain Galicia 103 Plaza et al. (2003) 1.9%
Spain Galicia 135 Maca-Meyer et al. (2003) 2.2%
Spain Catalonia 118 Maca-Meyer et al. (2003) 1.6%
Spain Huelva 135 Hernandez et al. (2014) 8.8%
Spain Maragatos 49 Maca-Meyer et al. (2003) 8.1%
Spain Canary Islands 300 Brehm et al. (2003) 14.0%
Portugal Countrywide 54 Plaza et al. (2003) 5.6%
Portugal North Portugal 184 Maca-Meyer et al. (2003) 4.3%
Portugal Central Portugal 161 Brehm et al. (2003) 1.9%
Portugal Madeira 155 Brehm et al. (2003) 3.9%
Iberia Spain & Portugal 887 Plaza et al. (2003) 1.8%

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

Haplogroup L lineages frequencies

Haplogroup L is the dominant haplogroup of Sub-Saharan Africa (96-100%) with lower levels occurring also in adjacent areas of North Africa, the Arabian Peninsula and parts of the Middle East.

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

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

In Iberia the mean frequency of Haplogroup L lineages reaches 3.83% and 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 with increase frequencies observed for Madeira (Portugal), Southern Portugal, Cordoba (Southern Spain) and North Western Castile (Spain). In the Autonomous regions of Portugal, L haplogroups constituted about 13% of the lineages in Madeira, significantly more than in the Azores. In the Spanish archipelago of 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. 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. Gonzalez et al. (2003) harvcoltxt error: no target: CITEREFGonzalez_et_al.2003 (help) 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 in prehistoric times rather than during the slave trade. According to Pereira et al. (2005) harvcoltxt error: no target: CITEREFPereira_et_al.2005 (help), the sub-Saharan lineages found in Iberia matched lineages from diverse regions in Africa. They suggest this pattern is more compatible with recent arrival of these lineages after slave trading began in the 15th century. According to the study, alternative scenarios that invoke much older and demographically more significant introductions (Gonzalez et al. (2003) harvcoltxt error: no target: CITEREFGonzalez_et_al.2003 (help)) or that claim a substantial role of the Roman and/or Islamic periods on the introduction of sub-Saharan lineages seem unlikely. Casas et al. (2006) harvcoltxt error: no target: CITEREFCasas_et_al.2006 (help) extracted DNA from human remains that were exhumed from old burial sites in Al-Andalus, Spain. The remains date to between 12th-13th century. The frequency of Sub-Saharan lineages detected in the medieval samples was 14.6% and 8.3% in the present population of Priego de Cordoba. The authors suggest the Muslim occupation and prehistoric migrations before the Muslim occupation would have been the source of these lineages. The highest frequencies of Sub-Saharan lineages found so far in Europe were observed by Alvarez et al. 2010 in the comarca of Sayago (18.2%) which is according to the authors "comparable to that described for the South of Portugal" and by Pereira et al. 2010 in Alcacer do Sal (22%).

In Italy, Haplogroup L lineages are present at lower frequencies than Iberia, and only detected in certain regions, such as between 2 and 3% in Latium, Tuscany, Basilicata and Sicily.

Haplogroup M1 is also found in Europe at low frequencies. In a study by Gonzalez et al. 2007, haplogroup M1 had a frequency of 0.3%. The origins of haplogroup M1 have yet to be conclusively established. One clade of haplgroup M1, M1a is widely accepted to be of East African origin. About 40% of all clades of M1 found in Europe are M1a and consequently of recent East African origin.


Country Region Number tested Study %
Albania 42 Belledi et al. (2000) 2.38%
Britain 76 Piercy et al. (1993) 1.00%
Finland 50 Sajantila et al. (1995) 2.00%
Germany 333 Pliss et al. (2005) 1.20%
Norway Central Norway 74 Passarino et al. (2002) 1.40%
Norway - Rosenberg et al. (2002) 1.00%
Norway - Wilson et al. (2001) 1.00%
Italy Countrywide 583 Brisighelli et al. (2012) 1.20%
Italy Tuscany 322 Achilli et al. (2007) 1.90%
Italy Latium 138 Achilli et al. (2007) 2.90%
Italy Marche 813 Achilli et al. (2007) 0.98%
Italy Volterra 114 Achilli et al. (2007) 2.60%
Italy Central Italy 83 Plaza et al. (2003) 1.20%
Italy Lombardy 177 Achilli et al. (2007) 0.00%
Italy Piedmont 169 Achilli et al. (2007) 0.00%
Italy Sardinia 84 Pereira et al. (2005) 2.90%
Italy Sardinia 73 Plaza et al. (2003) 2.80%
Italy Southern Italy 37 Plaza et al. (2002) 8.10%
Italy Campania (South Italy) 313 Achilli et al. (2007) 0.32%
Italy Basilicata (South Italy) 92 Ottoni et al. (2009) 2.20%
Italy Apulia & Calabria (South Italy) 226 Achilli et al. (2007) 0.00%
Italy Southern Italy 115 Sarno et al. (2014) 0.00%
Italy Sicily 49 Semino et al. (1989) 4.40%
Italy Sicily 106 Cali et al. (2003) 0.94%
Italy Sicily 105 Achilli et al. (2007) 1.90%
Italy Sicily 169 Plaza et al. (2003) 0.60%
Italy Sicily 198 Sarno et al. (2014) 1.01%
Iberia Spain & Portugal (South) 310 Casas et al. (2006) 7.40%
Spain Countrywide 312 Alvarez et al. (2007) 2.90%
Spain Central Spain 50 Plaza et al. (2003) 4.00%
Spain North-West Spain 216 Achilli et al. (2007) 3.70%
Spain Galicia 92 Pereira et al. (2005) 3.30%
Spain Zamora 214 Alvarez et al. (2010) 4.70%
Spain Sayago 33 Alvarez et al. (2010) 18.18%
Spain Cordoba 108 Casas et al. (2006) 8.30%
Spain Huelva 135 Hernandez et al. (2014) 5.70%
Spain Valencia 30 Plaza et al. (2003) 3.40%
Spain Catalonia 101 Alvarez-Iglesias et al. (2009) 2.97%
Spain Balearic Islands 231 Picornell et al. (2005) 2.20%
Spain Canary Islands 300 Brehm et al. (2003) 6.60%
Portugal Countrywide 549 Pereira et al. (2005) 5.83%
Portugal North 100 Pereira et al. (2010) 5.00%
Portugal Center 82 Pereira et al. (2010) 9.70%
Portugal South 195 Brehm et al. (2003) 11.30%
Portugal South 303 Achilli et al. (2007) 10.80%
Portugal Coruche 160 Pereira et al. (2010) 8.70%
Portugal Pias 75 Pereira et al. (2010) 3.90%
Portugal Alcacer do Sal 50 Pereira et al. (2010) 22.00%
Portugal Azores 179 Brehm et al. (2003) 3.40%
Portugal Madeira 155 Brehm et al. (2003) 12.90%
Greece Crete 202 Achilli et al. (2007) 0.99%
Cyprus Cyprus 91 Irwin et al. (2008) 3.30%

A similar study by Auton et al. 2009, which also contains an admixture analysis chart but no cluster membership coefficients, shows little to no sub-Saharan African influence in a wide array of European samples, i.e. Albanians, Austrians, Belgians, Bosnians, Bulgarians, Croatians, Cypriots, Czechs, Danes, Finns, Frenchmen, Germans, Greeks, Hungarians, Irish, Italians, Kosovars, 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

References

  1. ^ Balaresque, Patricia; Bowden, Georgina R.; Adams, Susan M.; Leung, Ho-Yee; King, Turi E.; Rosser, Zoë H.; Goodwin, Jane; Moisan, Jean-Paul; Richard, Christelle; Millward, Ann; Demaine, Andrew G.; Barbujani, Guido; Previderè, Carlo; Wilson, Ian J.; Tyler-Smith, Chris; Jobling, Mark A. (2010). Penny, David (ed.). "A Predominantly Neolithic Origin for European Paternal Lineages". PLoS Biology. 8 (1): e1000285. doi:10.1371/journal.pbio.1000285. PMC 2799514. PMID 20087410.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  2. Alonso, Santos; Flores, Carlos; Cabrera, Vicente; Alonso, Antonio; Martín, Pablo; Albarrán, Cristina; Izagirre, Neskuts; De La Rúa, Concepción; García, Oscar (2005). "The place of the Basques in the European Y-chromosome diversity landscape". European Journal of Human Genetics. 13 (12): 1293–302. doi:10.1038/sj.ejhg.5201482. PMID 16094307. {{cite journal}}: Invalid |display-authors=9 (help)
  3. B. Arredi, E. S. Poloni and C. Tyler-Smith (2007). "The peopling of Europe". In Crawford, Michael H. (ed.). Anthropological genetics: theory, methods and applications. Cambridge, UK: Cambridge University Press. p. 394. ISBN 0-521-54697-4.
  4. Swarthy, blue-eyed caveman revealed using DNA from ancient tooth
  5. Blue-Eyed Hunter-Gatherers Roamed Prehistoric Europe, Gene Map Reveals
  6. Stone Age Spaniard had blue eyes, dark skin
  7. Stone Age Europeans had dark skin and blue eyes, Spanish researchers say
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  12. 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. {{cite journal}}: Invalid |display-authors=9 (help)
  13. ^ Most British men are descended from ancient farmers, The Guardian, 20 January 2010
  14. ^ Most European males 'descended from farmers', BBC News, 20 January 2010
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  16. Flores, Carlos; Maca-Meyer, Nicole; González, Ana M; Oefner, Peter J; Shen, Peidong; Pérez, Jose A; Rojas, Antonio; Larruga, Jose M; Underhill, Peter A (2004). "Reduced genetic structure of the Iberian peninsula revealed by Y-chromosome analysis: implications for population demography". European Journal of Human Genetics. 12 (10): 855–63. doi:10.1038/sj.ejhg.5201225. PMID 15280900. {{cite journal}}: Invalid |display-authors=9 (help)
  17. Morelli, Laura; Contu, Daniela; Santoni, Federico; Whalen, Michael B.; Francalacci, Paolo; Cucca, Francesco (2010). Lalueza-Fox, Carles (ed.). "A Comparison of Y-Chromosome Variation in Sardinia and Anatolia is More Consistent with Cultural Rather than Demic Diffusion of Agriculture". PLoS ONE. 5 (4): e10419. doi:10.1371/journal.pone.0010419. PMC 2861676. PMID 20454687.{{cite journal}}: CS1 maint: unflagged free DOI (link)
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  21. 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.
  22. 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.
  23. 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.
  24. ^ 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)
  25. ^ González, Ana M.; Brehm, Antonio; Pérez, José A.; Maca-Meyer, Nicole; Flores, Carlos; Cabrera, Vicente M. (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.
  26. 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): 357–71. doi:10.1007/s00439-004-1168-9. PMID 15322918.
  27. Sutton, Wesley K.; Knight, Alec; Underhill, Peter A.; Neulander, Judith S.; Disotell, Todd R.; Mountain, Joanna L. (2006). "Toward resolution of the debate regarding purported crypto-Jews in a Spanish-American population: Evidence from the Y chromosome". Annals of Human Biology. 33 (1): 100–11. doi:10.1080/03014460500475870. PMID 16500815.
  28. Zalloua, P; Platt, D; El Sibai, M; Khalife, J; Makhoul, N; Haber, M; Xue, Y; Izaabel, H; et al. (2008). "Identifying Genetic Traces of Historical Expansions: Phoenician Footprints in the Mediterranean". The American Journal of Human Genetics. 83 (5): 633–42. doi:10.1016/j.ajhg.2008.10.012. PMC 2668035. PMID 18976729.
  29. "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). "
  30. "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 other population movements such as Arabs, Phoenicians, or even the first Neolithic settlers thousands of years ago" http://www.elmundo.es/elmundo/2008/12/04/ciencia/1228409780.html
  32. Spanish Inquisition left genetic legacy in Iberia, New Scientist, December 4, 2008
  33. Capelli, Cristian; Onofri, Valerio; Brisighelli, Francesca; Boschi, Ilaria; Scarnicci, Francesca; Masullo, Mara; Ferri, Gianmarco; Tofanelli, Sergio; et al. (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. See table
  34. 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.
  35. 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): 312–9. doi:10.1007/s004390000370. PMID 11129330. {{cite journal}}: Invalid |display-authors=9 (help)
  36. 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.
  37. 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.
  38. Goncalves, Rita; Freitas, Ana; Branco, Marta; Rosa, Alexandra; Fernandes, Ana T.; Zhivotovsky, Lev A.; Underhill, Peter A.; Kivisild, Toomas; Brehm, Antonio (2005). "Y-chromosome Lineages from Portugal, Madeira and Acores Record Elements of Sephardim and Berber Ancestry". Annals of Human Genetics. 69 (4): 443–54. doi:10.1111/j.1529-8817.2005.00161.x. PMID 15996172. {{cite journal}}: Invalid |display-authors=9 (help)
  39. 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.
  40. 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)
  41. ^ http://genome.cshlp.org/content/22/5/821/F1.large.jpg
  42. http://genome.cshlp.org/content/22/5/821.full#ref-12
  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. PMID 16201138. {{cite journal}}: Invalid |ref=harv (help)CS1 maint: multiple names: authors list (link)
  44. ^ 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. PMID 14513360. {{cite journal}}: Invalid |ref=harv (help)CS1 maint: multiple names: authors list (link)
  45. Alvarez; et al. (2010). "Mitochondrial DNA Patterns in the Iberian Northern Plateau: Population Dynamics and Substructure of the Zamora Province". PMID 20127843. {{cite journal}}: Cite journal requires |journal= (help); Explicit use of et al. in: |author= (help)
  46. 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, Alvarez et al. (2010)
  47. Attention: This template ({{cite doi}}) is deprecated. To cite the publication identified by doi:10.1002/ajhb.21049, please use {{cite journal}} (if it was published in a bona fide academic journal, otherwise {{cite report}} with |doi=10.1002/ajhb.21049 instead.
  48. Achilli; et al. (2008). "Mitochondrial DNA Variation of Modern Tuscans Supports the Near Eastern Origin of Etruscans". PMC 1852723. {{cite journal}}: Cite journal requires |journal= (help); Explicit use of et al. in: |author= (help)
  49. Ottoni; et al. (2009). "Human mitochondrial DNA variation in Southern Italy". PMID 19852679. {{cite journal}}: Cite journal requires |journal= (help); Explicit use of et al. in: |author= (help)
  50. Different genetic components in the Norwegian population revealed by the analysis of mtDNA and Y chromosome polymorphisms
  51. PLOS ONE: Uniparental Markers of Contemporary Italian Population Reveals Details on Its Pre-Roman Heritage
  52. 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)
Human genetics
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