Estimating the Celtic / Indo-European input in Spaniards

The Celtic original population most probably descends from the preceeding Central European Late Neolithic/Bronze Age cultures, such as the Bell Beakers or the Unetice.

The Iberian Peninsula before the Bronze Age was genetically middle-neolithic-like, that is, like the Early Neolithic farmers but with an increased level of WHG. But no steppe ancestry present.

But here I don't want to estimate raw steppe ancestry, but the whole LN/BA package. I've estimated this input to be around ~50% in Spaniards.

Using the nMonte stats, spaniards can be modeled as being roughly 40-50% Central Euro BA, and 40-50% Neolithic farmer, and also, depending on the regions of Spain, some minor north-african and/or west-asian might be added to make it a better fit :

      Spanish_Valencia

"Bell_Beaker_LN_Average"   39.75

"Stuttgart"     27.25

"Spain_MN__average"     15.7

"Unetice_BA"   14.95

"Mozabite_Berber"       2.35

Total of :  54.7%  Central-Euro LN/BA + 43.0% Neolithic + 2.35% north-african

Other regions might get more Unetice  than Bell Beaker :

 Spanish_Aragon
"Spain_MN__average"     33.25
"Unetice__BA"   28.9
"Bell_Beaker_LN_Average"        19.45
"Stuttgart"     14.8
"Mozabite_Berber"       3.6

Total :  48.4%  Central-Euro LN/BA + 48.1% Neolithic + 3.6 north-african

El mito de la mezcla de razas en España

En el estudio de Günther et al. 2015 se ha analizado una muestra de un individuo de la cueva de El Portañón (en Burgos) de la Edad de Bronze, que resulta ser genéticamente muy parecido a los españoles actuales.

Esto se puede visualizar fácilmente mediante un mapa genético de poblaciones (PCA plot), donde el indivudo "español" de la Edad de Bronze se solapa con los españoles actuales :

Españoles en los mapas genéticos

En estos mapas genéticos se puede visualizar la similud genétcia de entre varias poblaciones  (cuanto más cerca aparecen dos poblaciones en el mapa, más cercanas genéticamente).

Los españoles (del sur o del norte) aparecemos entre Franceses e Italianos del Norte (de Lombardia), algo que se asemeja a nuestra posición geográfica, sin embargo los norte-africanos quedan muy lejanos a nosotros, más emparentados estos últimos con poblaciones del medio-oriente.

Otras poblaciones del mediterráneo, como Italianos del Sur, Sicilianos y algunos Griegos, quedan algo lejanos a nosotros, de hecho hay aproximadamente la misma distancia genética entre ellos y nosotros, que la que hay entre poblaciones del Noroeste de Europa con nosotros.

Hacer click en la imágenes para agrandar :

Ancient DNA from Hunter-Gatherer and Farmer Groups from Northern Spain Supports a Random Dispersion Model for the Neolithic Expansion into Europe

Hervella et al. 2012

Samples : 

Haplogroups :

Haplogroup H is the major one, showing a frequency of 45% in the ancient samples analysed. This figure is similar to that observed in some present-day European populations including the North of the Iberian Peninsula

Two of the Palolithic hunter-gatherer from Cantabria belong to mtDNA H .

The three groups of hunter-gatherers considered in this analysis (from Scandinavia, Central Europe and the Cantabrian fringe on the Iberian Peninsula) did not show statistically significant differences between one another, but they are significantly different from any population compared, because to the high frequency of haplotypes within the haplogroup U (50%–80%) .

Abstract

Background/Principal Findings

The phenomenon of Neolithisation refers to the transition of prehistoric populations from a hunter-gatherer to an agro-pastoralist lifestyle. Traditionally, the spread of an agro-pastoralist economy into Europe has been framed within a dichotomy based either on an acculturation phenomenon or on a demic diffusion. However, the nature and speed of this transition is a matter of continuing scientific debate in archaeology, anthropology, and human population genetics. In the present study, we have analyzed the mitochondrial DNA diversity in hunter-gatherers and first farmers from Northern Spain, in relation to the debate surrounding the phenomenon of Neolithisation in Europe.

Methodology/Significance

Analysis of mitochondrial DNA was carried out on 54 individuals from Upper Paleolithic and Early Neolithic, which were recovered from nine archaeological sites from Northern Spain (Basque Country, Navarre and Cantabria). In addition, to take all necessary precautions to avoid contamination, different authentication criteria were applied in this study, including: DNA quantification, cloning, duplication (51% of the samples) and replication of the results (43% of the samples) by two independent laboratories. Statistical and multivariate analyses of the mitochondrial variability suggest that the genetic influence of Neolithisation did not spread uniformly throughout Europe, producing heterogeneous genetic consequences in different geographical regions, rejecting the traditional models that explain the Neolithisation in Europe.

Conclusion

The differences detected in the mitochondrial DNA lineages of Neolithic groups studied so far (including these ones of this study) suggest different genetic impact of Neolithic in Central Europe, Mediterranean Europe and the Cantabrian fringe. The genetic data obtained in this study provide support for a random dispersion model for Neolithic farmers. This random dispersion had a different impact on the various geographic regions, and thus contradicts the more simplistic total acculturation and replacement models proposed so far to explain Neolithisation.

R-L21 in Northern Spain

The study of Martínez-Cruz et al. 2012 shows a distribution of Y-DNA haplgroups from Basques and surrounding regions, from Spain and France. My focus is on the sublcade of R1b , R-L21, which is most common in the British Isles and Brittany, and until now it has not been much tested for in Iberia.
In some regions of Spain tested in this study it surpasses 20% of frequency.  Here is the table for all haplogorups and samples. The regions corresponding to Spain and their R-L21 frequency are :

Southwestern Gipuzkoa, n = 13/57 = 22.81%
Roncal Valley, Navarra: n = 11/53 = 20.8 %
Alaba, n = 11/51 = 21.57%
Guipuzkoa, n = 9/47 = 19.15%
Central/Western Navarra, n = 9/60 = 15.0%
Bizkaia, Basque Country, n = 7/57 = 12.28%
La Rioja, n = 6/54 = 11.11 %
Western Bizkaia , n = 3/19 = 10.53%
Northwestern Navarra, n = 5/51 = 9.80%
Northern Aragón : n = 1/27 = 3.7%

Total : n = 75/476 = 15.76%

mtDNA L in Europe and Spain

Here is a compilation from many different studies, for the frequencies of the maternal Sub-Saharan haplogroup mtDNA L,  for countries from all around  Europe. At the bottom, there is a compilation for the total of Spain. We'll see that spanish percentages are similar to other countries and/or regions of Europe. 

For some obscure reasons, on the Wikipedia articles  "mtDNA L in Europe", and  "African admixture in Europe" there is only  Iberia and Italy as european countries having mtDNA L, we will see here that this is not the case, there is in all corners of the continent. And I advise you if you dare to edit those articles,  you'll probably get blocked.


Slovaks              n = 2/207    0.97%      Malyarchuk et al.2006

Czechs              n = 1/279     0.36%      Malyarchuk et al., 2008

German-Danish  n = 1/161     0.62%      Richards et al. 1996
Germany            n = 1/335     0.30%      Achilli et al 2007
Germany            n = 4/333    1.20%      Pliss et al. 2005

Albania             n =  1/42      2.38%     Belledi et al. 2000
Bosnia             n=1/144        0.70%    Achilli et al 2007
Bulgaria           n=1/141        0.71%    Achilli et al 2007
Balkans          n= 1/556        0.18%    Regueiro et al. 2012 + King et al. 2011


Poland              n = 1/542     0.20%      Achilli et al 2007
Poland              n = 1/436     0.22%      Malyarchuk et al., 2002
Poles                n = 1/849     0.12%      Malyarchuk et al., 2008


British               n=1/100      1.00%      Piercy et al. 1993
England            n = 2/335     0.60%      Achilli et al 2007
England            n  = 1/142    0.70%     Helgason et al., 2001
Great Britain     n = 1/114     0.90 %     García et al. 2011
Scotland           n = 1/891     0.10 %     García et al. 2011


Finnish             n = 1/50       2.00%     Sajantila et al. 1995
Finland            n = 1/121     0.83%     Achilli et al 2007
Finns               n = 3/580     0.52%     Pliss et al. 2005


Norway          n = 1/74       1.40%     Passarino et al 2002
Norwegians    n = 2/397     0.50%      Pliss et al. 2005


Russia            n =  2/683     0.29%     Malyarchuk et al. 2008


France            n = 1/332      0.30%    Achilli et al 2007
France            n = 5/692      0.72%     Richards et al. 2007
France            n =  3/433     0.69%     García et al. 2011
France            n = 2/320      0.63%     Pliss et al. 2005
France                                 1.40%     Gónzalez et al. 2003


Switzerland      n=1/228        0.44%    Achilli et al 2007


Crete, Greece  n= 2/202       0.99%    Achilli et al 2007


Italy                n = 8/411      1.94%     Plaza et al. 2003,  Romano et al. 2003

NW Italy         n = 100        2.00%    Brisighelli et al. 2012

Sardinian         n = 2/69       2.90%     Di Rienzo and Wilson 1991
Sardinia          n =2/370       0.54%    Achilli et al 2007

Sicily               n= 1/106       0.94%     Cali et al. 2001
Sicily               n = 2/105      1.90%     Achilli et al 2007
Sicily              n = 3/465       0.65%    Plaza et al. 2003,  Romano et al. 2003
South Italy     n = 2/313        0.64%    Boattini et al. 2013 + Stefania Sarno et al. 2014

Tuscany          n= 6/322        1.86%    Achilli et al 2007
Marche (Italy) n=8/813         0.98%    Achilli et al 2007
Latium(Italy)    n=4/138        2.90%    Achilli et al 2007
Murlo (Italy)    n = 1/86       1.16%     Achilli et al 2007
Volterra (Italy) n =  3/114     2.63%    Achilli et al 2007
Casentino (Italy) n =  2/122  1.64%   Achilli et al 2007
Campania        n = 3/313      0.32%   Achilli et al 2007



Spain Total   n = 51 / 4.329     1.18%    From here :

Spain   n = 15/813   1.84%       Achilli et al 2007
Spain,  n =  0/686    0.00%       Rhouda 2006
Spain   n =  0/233    0.00%,      Lopez-Parra 2009
Spain   n =  1/251    0.40%       Plaza et al. 2003
Spain   n =  9/312    2.90%       Álvarez et al. 2007
Spain   n =  1/108    0.90%       Casas et al. 2006 *
Spain   n =  2/226    0.88%       Maca-Meyer et al. 2003
Spain   n = 15/742   2.02%      García et al. 2011
Spain (Basques)  n = 0/462      García et al. 2011
Spain, Zamora Province**  n =10/214    4.67%      Álvarez  et al. 2010
Spain, n = 8/496     1.61%       Pereira et al. 2005

Note that in the García et al. 2011 there is overlapping with other studies, that's why the total shown here  is not the same as the total you might get from the study.

- * Note that Casas et al. 2006 only considered L1-L2 sequences to be of sub-Saharan origin.
-** This study inlcudes only a sample from a small town of the province of Zamora, thus it is not very representative of the population of Spain, I prefer wider regions, anyways I have included them.
- Note about the subclade L3, some scholars don't consider it as Sub-Saharan (Maca-Meyer 2003 et al.) in her words :
""L3 is more related to Eurasian haplogroups than to the most divergent African clusters L1 and L2". L3 is the haplogroup from which all modern humans outside of Africa derive"

Total E-M81 in Spain

UPDATE (13/03/2012) : Added the samples from Martínez-Cruz et al. 2012, which includes Navarrese, Burgos (Castilla),  La Rioja and Cantabria.

Spanish Basques     n=2/55    3.6%    Cruciani 2004.
Basques                  n=1/45    2.2%    Underhill 2000. 
Basque Country      n=1/116  0.9%    Adams et al.
Basques                 n=1/168   0.6%    Capelli 2009 (Alonso et al. + Garcia et al)
Basques                  n=1/43    2.3%    Capelli 2009 from Bosch et al.
------------
Total Basques  6/427 = 1.4%


Navarra, Cinco Villas       n=0/42    0.0%       López-Parra 2009
Lapurdi, Navarra              n =0/66   0.0%       Martínez-Cruz et al. 2012
Central/Western Navarra  n = 1/60  1.7%       Martínez-Cruz et al. 2012
Northwestern Navarra     n = 0/51   0.0%       Martínez-Cruz et al. 2012
Valle Roncar, Navarra     n = 2/53   3.77%     Martínez-Cruz et al. 2012
 ------
Total Navarra :   3/272 = 1.10%


Pasiegos*               n=15/64     24%        Maca-Meyer 2003.
Lebaniegos             n=0/37       0.0%       Maca-Meyer 2003.
Other Cantabrians   n=6/36      16.7%      Maca-Meyer 2003.
Cantabria               n=17/161   13.0%      Brión et al. 2004
Cantabria               n=1/18        5.56%     Martínez-Cruz et al. 2012
--------
Total Cantabria      n=39/316 = 12.23%


La Rioja,   n = 0/54      0.0%                 Martínez-Cruz et al. 2012
 

Asturians             n=2/90       2.2%       Cruciani 2004.
Asturias              n=0/20       0.0%       Adams et al.   
-----------
Total Asturias      n=2/110 = 1.82%


Aragón                   n=3/34    8.8%     Adams et al.
Aragón, Jacetania   n=0/31    0.0%     López-Parra 2009
Northern Aragón    n =0/27   0.0%     Martínez-Cruz et al. 2012 
------------------
Aragón :  3/92  3.26 %
  

Galicia             n=2/19      10.5%    Flores et al.2004
Galicia             n=9/88      10.2%    Adams et al. 2008
Galicia             n=12/292    4.1%    Brión et al.2004
--------
Total Galicia    23/399= 5.76%


Valencia            n=2/31     6.5%       Flores et al.2004
Valencia            n=4/73     5.48%    Adams et al.
----------
Total Valencia   6/104 = 5.77%


Castille                  n=1/21     4.8%     Flores et al.2004
Burgos                  n =1/20    5.0%     Martínez-Cruz et al. 2012
Castilla Mancha     n=2/63     3.2%    Adams et al.
Castile NE            n=3/31      9.7%    Adams et al.
Castille NW          n=10/100 10.0%   Adams et al.
----------
Total Castilla   17/235 = 7.23%


Catalans            n=0/16     0.0%     Bosch et al.2001
Catalonia           n=1/80    1.25%   Adams et al.
Vall d'Arán         n=1/25    4.00%   López-Parra 2009
Cerdeña             n=0/37    0.00%   López-Parra 2009
Alt Urgell          n=0/34    0.00%   López-Parra 2009
-----------
Total Catalonia  2/192 = 1.04%


Extremadura         n=4/52   8.0%   Adams et al.
-----------

Majorca             n=2/62    3.22%    Adams et al.
Minorca             n=3/37    8.1%      Adams et al.
Ibiza                 n=0/54     0.0%      Adams et al.
----------
Balearic Islands : 5/153 = 3.27%


Southern Spaniards   n=1/62      1.6%     Scozzari 2001.
Huelva, Andalusia     n=5/167   2.99%    Ambrosio 2010.
Huelva, Andalusia     n=1/22      4.5%     Flores et al.2004
Seville, Andalusia     n=7/155     4.5%     Flores et al.2004
Cadiz, Andalusia      n=0/28       0.0%     Flores et al.2004
Cordoba, Andalusia  n=2/27      7.4%     Flores et al.2004
Malaga, Andalusia   n=3/26     11.5%     Flores et al.2004
Andalusia East         n=2/95      2.1%     Adams et al.
Andalusia West       n=10/73    13.7%    Adams et al.
Andalusians           n=2/37        5.4%     (Capelli 2009)  Bosch et al.
------------------
Total Andalusia :  33/692 = 4.77%

TOTAL SPAIN =   143/3.098 =  4.62 %

*without Pasiegos = 128/3034 = 4.22 %

Canarians were not included for obvious reasons.


Observations :

- We obtain a result of between 4 and 5%, slightly lower than that observed for Portugal (5.73%).

- The closest spanish region to the national average would be Andalusia (4.77%).

- As mentioned before, I reiterate the lack of correlation between islamic history in Iberia and these subclade frequencies, such that Andalusia (the place were Islamic religion lasted longer, specifically Western Andalusia) has less E-M81 than some northern regions such as Galicia, Cantabria, Valencia, Castille, which essentially means the presence of this subclade in Iberia has very little to do with the islamic period, in other words, the moors (in the ethnic sense)  had a very minoritary presence in Spain.

- Another obersvation I make, is the difference between Galicia and it's close neighbour Asturias, which has amongst the lowest frequency of the spanish communities.

- The lowest presence is in Catalonia, with aprox. 1%,  followed by Basques (1.4%) and the highest in Cantabria (12.75% with Pasiegos, 9.8% without Pasiegos).

- Obviously the samples of the various communities are still very small and unrepresentitative, so this percentages are only orientative.

- Compared to other nations, the closest country besides Portugal in terms of similar percentages, would be France, with 4.42 % (see here).

Total E-M81 in Portugal

Southern Portuguese  n=6/49   12.2%   Cruciani 2004.
Southern Portugal      n=6/78  8.00%   Adams et al.
South Portugal          n=6/100   6.0%   Goncalves 2005
Total South :   20/257 = 7.8%

Central Portugal    n=5/102   4.9%   Goncalves 2005

Northern Pprtuguese   n=2/50    4.0%    Cruciani 2004.
Northern Portuguese    n=6/109   5.5%   Flores et al.
Northern Portugal        n=2/60    3.0%    Adams et al.
North Portugal            n=6/101   5.9%   Goncalves 2005
Total North :  17/320 = 5.3%

Portugal               n= 33/659  5.0%   Capelli et al. 2009 (includes Beleza et al.)

Total Portugal (all samples)  n = 70 / 1.308 = 5.35%

Very similar to the percentage of Galicia (5.76%) and below 6% again when considering many studies. This percentage is similar to some areas of France ( like Auvergne ( 5.62%,  Cruciani 2004), and the North of Portugal has higher than spanish southern region of Andalusia (4.7% see my previous entry). I will publish another entry with the total for Spain.

E-M81 in Galicia

Total of E-M81 in Galicia (NW Spain), considering many studies :

Galicia             n=2/19    10.5%    Flores et al.2004
Galicia             n=9/88    10.2%    Adams et al.
Galicia             n=12/292   4.1%    Brion et al.2004
--------
Total Galicia    23/399= 5.76%

It's actually below 6% when combining different studies, and similar to those found in some areas of France, like Auvergne ( 5.62%,  Cruciani 2004), and higher than those found in Andalusia (4.7% , see here). This presence is obviously due to pre-historic migration, because muslim history in Iberia doesn't correlate with genetics, as seen here, when Andalusia has less E-M81 than some northern spanish communities.

North-African marker E-M81 in Europe

Here is a list obtained from different studies, for the typical North-African y-dna subclade E-M81 (part of the marco haplogroup E) , for populations of Europe (outside of Iberia) :


West-Flanders : 1/110 = 0.9%   Larmuseau et al. 2013
Vlaams-Waals Brabant :  1/124 =  0.8%  Larmuseau et al. 2013
Belgium Total (all the study) :  2/773 = 0.3%  Larmuseau et al. 2013
------------------

Germany   n= 1/345    0.29%    Rebala et al. 2012
---------------

Haskovo, Bulgaria n = 1/41  2.4%  Karachanak 2013.
Total Bulgaria    n = 1/808  0.12%   Karachanak 2013.
----------------

Greeks               n = 3/92    3.3%   Battaglia et al. 2008
Macedonian-Greeks  n=1/57   1.8%   Battaglia et al. 2008

----------------
Croats               n = 1/89   1.1%   Battaglia et al. 2008
----------------

North-Italy
North-Italians      n=1/67  1.5%   Cruciani 2004.
Lombard             n=1/18  5.6%   Scozzari 2001.
Venetian              n=20     0.0%   Scozzari 2001.
Ligurian               n=17     0.0%   Scozzari 2001.
Bologna               n =29    3.4%  Boattini et al. 2013           

Central-Italy
Central Italians    n=2/89     2.2%   Cruciani 2004.
Latium                n=1/66     1.5%   Scozzari 2001.
Toscana             n=123       0.8%    Boattini et al. 2013

South Italy/Sicily
Southern Italians   n=0/87    0.0%   Cruciani 2004.
Sicilians                 n=1/136   0.7%   Cruciani 2004.
Sicily                    n= 141     2.8%   Boattini et al. 2013
Cosenza              n=1/45      2.22%

Sardinia
Sardinians             n=1/367   0.3%   Cruciani 2004.
Sardinia                n=82        1.2%   Boattini et al. 2013
-------------------------- 

French               n=3/85    3.5%   Cruciani 2004.
French              n=3/73  4.1%   Scozzari 2001.
Auvergne           n=5/89  5.62   Cruciani 2004.
Île-de-France     (5/91)   5.49   Cruciani 2004.
Gascony             n=0/24  0.0%    Adams et al.
Béarn, Gascony   n=1/56  1.78%  Martínez-Cruz et al. 2012,
Bigorre, Gacony  n=1/44  2.72%  Martínez-Cruz et al. 2012

Provence-Alpes-Côte d'Azur	n=1/45 2.22%	Ramos-Luis et al. 2013.
Nord-Pas-de-Calais	n=3/70 4.29%	Ramos-Luis et al. 2013
Midi-Pyrénées	n=1/67 1.49%	Ramos-Luis et al. 2013.

North-African marker in Andalusia

We find the following percentages for the typcial north-african (Berber) paternal marker, the Y-DNA haplogroup E-M81, obtained from different sources :

Southern Spaniards  n=1/62    1.6%    Scozzari 2001.
Huelva, Andalusia   n=5/167   2.99%  Ambrosio 2010.
Huelva, Andalusia   n=1/22      4.5%   Flores et al.2004
Seville, Andalusia  n=7/155     4.5%    Flores et al.2004
Cadiz, Andalusia    n=0/28      0.0%    Flores et al.2004
Cordoba, Andalusia  n=2/27   7.4%    Flores et al.2004
Malaga, Andalusia   n=3/26    11.5%   Flores et al.2004
Andalusia East      n=2/95       2.1%    Adams et al.
Andalusia West      n=10/73   13.7%   Adams et al.
-------------------------
Total South Spain :  31/655 = 4.73%

It is a similar result to percentages found in  French  (3/73 = 4.1% , Scozzari et al. 2001) or in regions of France ( 5.6% in Auvergne (5/89) and Île-de-France 5/91 = 5.5% ). This suggests that the presence of Islamic rule, which was longer in the South of Spain than anywhere else in Iberia, had very limited or no impact in the genetic pool of the population. We can conclude that the moorish population in Andalusia was extremely minoritary, the place were Islamic rule was stronger and longer. We can't also rule out the possibily that the presence of this marker, or part of it, could have entered in much earlier times (Neolithic or earlier), which would make the presence of moors in Al-Andalus even more insignificant. So, in other words, we can safely say the Islamic rule was just that, a military and religious rule, with no impact on the demographics and genetics of the authoctonous populations.


Ambrosio et al. 


The distribution of E-M81 haplogroup, a Berber marker, was found at a frequency of 3% in our sample. The distribution of M81 frequencies in Iberia seems to be not concordant with the regions where Islamic rule was most intense and long-lasting. The study also showed that most of M78 derived allele (6.6%) led to the V13* subhaplogroup. We also found the most basal and rare paragroup M78* and others with V12 and V65 mutations. The lineage defined by M34 mutation, which is quite frequent in Jews, was detected as well. Conclusions: The haplogroup E among Western Andalusians revealed a complex admixture of genetic markers from the Mediterranean space, with interesting signatures of populations from the Middle East and the Balkan Peninsula and a surprisingly low influence by Berber populations compared to other areas of the Iberian Peninsula.

Sub-Saharan admixture in the Mediterranean basin

González-Pérez et al. (2010) have analyzed populations from the northern and southern shores of the Mediterranean, with Central Europeans and West Africans as external references. In the Discussion section, they admit that the inflated "Alu/STR estimate might be artefactual" and favor the estimate based on the Alu  loci set alone because it's consistent with previous mtDNA, Y-chromosome and 500,000-SNP structure data.
According to the more accurate latter method, Sub-Saharan African admixture is ~13% in North Africa and "imperceptible" or noise (~ 0.01%) in Southern Europe:

Pyrenean Y-DNA

In the pie charts, dark blue refers
to R1b1b2d, dark light to R1b1b2c, white to other R, orange
to I2a2, salmon to other I and pink to lineages that entered
Iberia from the Neolithic onwards. The light gray slice refers to
R plus I lineages.

From the study of  López-Parra et al. 2009 which analyses 5 pyrenean populations (25 males from Vall d'Arán (Lleida province), 34 from Alt Urgell (Lleida province), 37 from Cerdanya (Girona province), 31 from Jacetania (Huesca province) and 42 from Cinco Villas (Navarre province).


As expected, the vast majority by far of these populations belong to haplogroup R1b1b2-M269 (total 131/169 = 77.5%), especially the subhaplogroup R1b1b2*, and the second most common haplogroup among the total Pyrenean samples was haplogroup I which accounted for 12.4% (21/169) of paternal lineages, being I2a2 the most common clade. The rest of haplogroups are quite minoritary. Here are the distributions for each of the pyrenean populations sampled :

Cerdeña (n=37)
R1b: 75.7%  I: 16.2%  G : 2.7%  J: 2.7%  E: 0% 

Alt Urgell (n=34)
R1b: 76.5%  I: 8.8%  J: 8.8%  E: 2.9% 

Vall d'Arán (n=25)
R1b : 84%  I: 12%  E: 4%  J: 0%

Jacetania (N=31) :
R1b : 64.%  I: 16.1%   J: 9.7%  R1: 6.45%    G: 3.2%   E: 0%

Cinco Villas (n=42)
R1b: 85.7%  I:  14.3%

TOTAL (n=169) :  R1b (131/169) : 77.5%  I (21/169) : 12.4%  J (7/169) = 4.14%, G (3/169) = 1.78% E (3/169) = 1.78%,    R1 (2/169) = 1.18%  and  Other 2/169 = 1.18%.

Summary

The male-mediated genetic legacy of the Pyrenean population was assessed through the analysis of 12 Y-STR and 27 Y-SNP loci in a sample of 169 males from 5 main geographical areas in the Spanish Pyrenees: Cinco Villas (Western Pyrenees), Jacetania and Valle de Arán (Central Pyrenees) and Alto Urgel and Cerdaña (Eastern Pyrenees). In the Iberian context, the Pyrenean samples present some specificities, being characterizeded by a high proportion of chromosomes R1b1b2-M269 (including the usually uncommon R1b1b2d-SRY2627 and R1b1b2c-M153 types) or I2a2-M26 and low proportions of other haplogroups. Our results indicate that an old pre-Neolithic substrate is preponderant in populations of the whole Pyrenean fringe. However, AMOVA revealed a high level of substructure within Pyrenean populations, partially explained by drift effects as well as by the signature of an ancient genetic differentiation between Western and Eastern Pyrenees.

Other interesting extracts from the study : 

I2a2 is virtually absent east of the Italian Apennines and shows the highest incidences in north-eastern Iberia/southern France, with the exception of the isolated and dramatic peak of frequency (40.9%) in Sardinia (Rootsi et al. 2004).
This was well illustrated in the phylogram of I2a2 in Figure 1 from Rootsi et al. (2004) based upon the population data available at the moment, within which Basques (Spanish and French mixed) and Bearnais (in the French Atlantic Pyrenees) showed the highest continental frequencies (6% and 7.7% respectively). New data from Spanish Basques (Alonso et al. 2005) did not reproduce such elevated values; in Biscay I2a2 was not detected and its frequency was 1.3% in Gipuzkoa and 4.5% in Alava plus Navarra. From the new data presented here it seems that the Pyrenees might indeed have been the region where I2a2 arose and from which it initiated the spreading process after the LGM.

One of the signs comes from the concentration of high frequencies of I2a2 among populations from the entire Pyrenean range. Our data strongly reinforce previous evidence that I2a2 arose during Mesolithic times in a region close to or within the Pyrenees. The dispersal of I2a2 from its place of origin throughout the Pyrenees and beyond, implied not only gene exchange but also considerable movement of people. Very likely, the demographic event associated with the expansion of I2a2 was the Ice-age repopulation of
Europe from the Franco-Cantabrian refuge. A number of studies on human mtDNA diversity have already indicated that the Franco-Cantabrian glacial refuge was a major source for the European gene pool (Achilli et al. 2004), and our data on I2a2 seemingly lend support to the role of the region as a Mesolithic diffusion center of male lineages.

Celtiberia population estimate

From the book  ENTRE Celtas e Íberos. Las poblaciones protohistóricas de las Galias e Hispania/ editado por Luis Berrocal-Rangel y Philippe Gardes. — Madrid : Real Academia de la Historia : Casa de Velázquez, 2001. — 248 p. : il. ; 30 cm. —(Bibliotheca Archaeologica Hispana ; 8).

Note, the Celtiberia here refers only to the proper Celtiberian tribes, and not all the Celts of Iberia.

Abstract

The demographic analysis of the Celtiberian is a hard question. To solve it we have analysed the ethnoarchaeological data and we have contrasted it with archaeological data, as the surface of the known Celtiberian civitates and oppida. We have contrasted the results with the few informations transmitted by classical historians and geographers, as the numbers of warriors in Celtiberian armies and the extension of the populi and civitates of Celtiberia after Plinius and Ptolomeus. As conclusion of the analysis, we can consider that Celtiberia, extended about 45.000 km2 had a theoretical demographic density of about 5/6 h./km2 in mountain areas, but could reach 8/10 h./km2 in the best areas, as river valleys and peripheral plains. These results allow us a calculation of the total Celtiberian population between 250.000 h. and about 350.000/450.000 h., a bigger and more logical number. The first demographic anlysis of the Celtiberia offers a interesanting information and very useful for further studies on Archaeology and Ancient History and Geography, and also to better understand the territorial and urban planning, the demography and the sociology of the Celtiberians.

ADMIXTURE up to K=20 (Rasmussen et al. 2011)

The recently published paper by Rasmussen et al. is focused on the Aboriginal Australians, and inlcudes a massive ADMIXTURE analysis up to 20 ancestral components (K=20). The Spaniard sample (taken from Behar et al.) in this run is similar-looking to other Western-European populations, showing the dark-blue component (peaks in Lithuanians) at similar levels as the French, and the whiteish-blue component (peaks in Sardinians).

The original size can be found in the supplementary PDF here (page 159) :

Sub-Saharan haplogroup mtDNA L in Spain and Europe

In the study of Rhouda et al. (2006) in a sample of 686 spaniards, not a single individual with haplogroup mtDNA L was found. In the rest of Western Europe, the recent study of García et al. 2011 finds L up to Northern Germany and Denmark . We observe the following percentages  :

Hérault (Languedoc) 2.4 %
Rhône (Lyonnais) 4.4 %
Vendée and Vienne (Poitou) 0.8 %
Calvados and Seine-Maritime (Normandy) 1.8 %
Somme (Picardie) 1.3 %
France Miscellanea 0.6 %
Scotland 0.1 %
England 0.7 %
Great Britain 0.9 %
North-Germany and Denmark 0.7 %

Skin reflectance of selected world populations

From the study of Nina Jablonski and George Chaplin (Academy of Sciences of California).
We can see that Spaniards, from León (NW Spain) and Basques (N Spain), are as light or even lighter than other Western Europeans. Higher values indicate lighter skintone.

Country and population or area	Observed reflectance at 685 nm
EUROPE
Netherlands	67•37
Germany (Mainz)	66•90
United Kingdom (Northern)	66•10
Spain (Basques)	65•70
United Kingdom (Wales)	65•00
Ireland (Rossmore)	64•75
Spain (Leon)	64•66
Belgium	63•14
United Kingdom (London)	62•30
WEST ASIA
Iraq/Syria (Kurds)	61•12
Turkey	59•15
Israel	58•20
Lebanon	58•20
Jordan	53•00
Saudi Arabia	52•50
NORTH AFRICA
Algeria (Aures)	58•05
Tunisia	56•30
Morrocco	54•85
Libya (Tripoli)	54•40
Libya (Fezzan)	44•00
SOUTH ASIA
India (Northern)	53•26
Pakistan	52•30
India (Southern)	46•70
EAST ASIA
China (Southern)	59•17
Vietnam	55•90
Japan (Northern)	54•90
Philippines (Manila)	54•10
Cambodia	54•00
Japan (Southwest)	53•55
Nepal (Eastern)	50•42
AUSTRO-MELANESIA
Papua New Guinea	35•30
Australia (Darwin)	19•30
AMERICAS
Greenland (Southern)	55•70
Peru (Nunoa)	47•70
Peru (Maranon)	43•05
SUB-SAHARAN AFRICA
South Africa (Hottentot)	46•80
Botswana (San)	42•40
Zaire	33•20
Kenya	32•40
Ethiopia	31•70
Tanzania (Sandewe)	28•90
Namibia	25•55
Cameroon (Fali)	21•50
Mozambique (Chopi)	19•45

Source : "The Evolution of Human Skin Coloration". J Hum Evol, 2000; pp. 74-75

R1b frequencies in Spain and Europe

The paper from Belareseque et al. 2010 gives us new percentages of R1b1b2 from different countries of Europe, as we can see in the table below. There is also a map of frequencies. As we can see again, Spain has among the highest percentages of Europe, especially the Spanish Basques, with 87.1 %, second most in Europe after Wales with 92.3 %.

Here are the levels for the regions of Spain :

• Spain Basques........................87.1 %
• Catalonia, Spain .....................81.3 %
• Andalucia East, Spain ............72.0 %
• Castilla La Mancha, Spain..... 72.0 %
• Galicia, Spain.........................58.0 %
• Andalucia West, Spain...........55.0 %

 Complete table with all countries :

Map of frequencies : 

Spaniards genetically similar to Western and Northern Europeans

Study from Gayán et al. 2010 , some extracts :
 
In this study we have sampled over 800 unrelated individuals from the population of Spain, and have genotyped them with a genome-wide coverage. We have carried out linkage disequilibrium, haplotype, population structure and copy-number variation (CNV) analyses, and have compared these estimates of the Spanish population with existing data from similar efforts. Conclusions: In general, the Spanish population is similar to the Western and Northern Europeans but has a more diverse haplotypic structure. These results suggest that the general Spanish population, as characterized in the present study by sampling from eight different cities widely-spaced across Spain, is generally similar to other European populations, although more genetically diverse than Western and Northern Europeans. Moreover, the Spanish population is remarkably homogeneous within itself in terms of global genetic structure. In view of these results, the population of Spain is sufficiently genetically similar to the CEU sample (White-Americans from Utah) so that the CEU HapMap dataset could be used to infer genotypes for the Spanish population.

North-African mtDNA in Spain...and Europe

The minor presence of the typical north-african mtDNA haplgroup U6  in Spain, even if at low levels, was tought to be higher than anywherelse in Europe. Interestingly we will see that this is not the case. The truth is that U6 is found everywhere in Europe, but it has been found at higher frequencies in France. Obviously these studies make sure the testees are natives, having the 4 grandparents from the same area, and their surnames native. We see now the studies :

In this study of Dubut et al. 2004 there is a total of 4.5% of U6 in Finistère (Brittany, France) and 1.4% in Périgord-Limousin.

In this very recent study of García et al. 2011 we find these frequencies for France :

• U6a1/a2/a3 Perigord-Limousin (Limousin) 1.4 %
                     Finistère (Brittany) 0.7 %
• U6a  France Miscellanea 0.6 %

Spain : 

Andalusians n=3/158    1.9%   Plaza 2003.

Asturias   n=1/89     1.1%    Garcia 2011.

Aragón     n=0/119    0.0%    Garcia 2011.

Catalonia-Aragón  n=2/164  1.2%  Garcia 2011.
Catalonia n=3/80    3.9%      Garcia 2011.
Catalonia n=78     0.0 %    Plaza 2003.

Central Spain   n=1/50  2.0%   Plaza 2003.

Basques    n=45  ??   0.6%    Maca-Meyer 2003.
Basques    n=0/377    0.0%    Garcia 2011.
Basques    n=1/85     1.2%    Garcia 2011.
Basques    n=173      0.0%    Plaza 2003.

Galicia      ??       2.3%    Maca-Meyer 2003.
Galicia    n=3/185    1.5%    Garcia 2011.
Galicia    n=2/103      1.9%    Plaza 2003.

Cantabria  n=1/105    1.0%    García 2011.
Potes      n=0/72     0.0%    Maca-Meyer 2003.

Lebaniegos n=0/72   0.0%      Maca-Meyer 2003.
Pasiegos   n=0/82   0.0%      Maca-Meyer 2003.

Valencia   n=0/30     0.0 %     Plaza 2003.

% M1 and U6a1a = 2/214 = 0.93%   Álvarez 2010


TOtal SPain = 18/2.022 = 0.89%