# Population Genetics > mtDNA Haplogroups >  African MtDna Signatures in the Iberian Peninsula

## Angela

This is a long awaited look at the issue, and particularly because they looked at whole mtDna sequences.

Candela L. Hernandez et al:
*Early Holocenic and Historic mtDNA African Signatures in the Iberian Peninsula: The Andalusian Region as a Paradigm*http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0139784

Abstract
_"Determining the timing, identity and direction of migrations in the Mediterranean Basin, the role of “migratory routes” in and among regions of Africa, Europe and Asia, and the effects of sex-specific behaviors of population movements have important implications for our understanding of the present human genetic diversity. A crucial component of the Mediterranean world is its westernmost region. Clear features of transcontinental ancient contacts between North African and Iberian populations surrounding the maritime region of Gibraltar Strait have been identified from archeological data. The attempt to discern origin and dates of migration between close geographically related regions has been a challenge in the field of uniparental-based population genetics. Mitochondrial DNA (mtDNA) studies have been focused on surveying the H1, H3 and V lineages when trying to ascertain north-south migrations, and U6 and L in the opposite direction, assuming that those lineages are good proxies for the ancestry of each side of the Mediterranean. To this end, in the present work we have screened entire mtDNA sequences belonging to U6, M1 and L haplogroups in Andalusians—from Huelva and Granada provinces—and Moroccan Berbers. We present here pioneer data and interpretations on the role of NW Africa and the Iberian Peninsula regarding the time of origin, number of founders and expansion directions of these specific markers. The estimated entrance of the North African U6 lineages into Iberia at 10 ky correlates well with other L African clades, indicating that U6 and some L lineages moved together from Africa to Iberia in the Early Holocene. Still, founder analysis highlights that the high sharing of lineages between North Africa and Iberia results from a complex process continued through time, impairing simplistic interpretations. In particular, our work supports the existence of an ancient, frequently denied, bridge connecting the Maghreb and Andalusia."
_

They're much clearer in the actual body of the paper, where they posit two peaks of migration, one about 10,000 ybp, and one in the historic era.
_Thus, the most parsimonious model for the oldest demographic events and migrations across the Mediterranean Basin, based on mtDNA evidence, is the following: after 20 ky, U6 lineages had an extensive population expansion in northwest Africa, associated with the emergence of the Iberomaurusian industry in the Maghreb; this pool was further enriched by sub-Saharan L lineages, especially L1b, which began to arrive in North Africa in the beginning of the African Humid Period (AHP, ~11–5.5 ky BP) [57]; U6 and L lineages were introduced from northwest Africa into Iberia in the post-glacial period, most probably by the time of the Younger Dryas/beginning of the Holocene. The opening of the trans-Saharan communications with the African Humid Period, also allowed the southern migration of U6 sequences, and its local expansion as detected in the BSP for lineages observed in sub-Saharan Africa.
_
_"The recent migration peaks identified in the FA of U6 and L lineages could be associated with the Islamic rule of Iberia and the slave trade period, respectively. The HVS-I FA attributes a comparatively lower proportion of these recently introduced sequences into Iberia when compared with the post-glacial one: 1/3 vs 2/3, respectively. The analysis performed by [26] on the phylogeography of L sequences observed in Iberia and remaining Europe pointed to 65% of its introduction in recent times (Romanization period, Islamic expansion and Atlantic slave trade), and 35% at older times, as earlier as 11ky. The complete sequence based FA for U6 agrees more with these results, attributing a half-half proportion of sequences in both periods, showing the higher resolution of complete mitogenomes.  The historical-based expectation of a higher proportion of newly introduced U6 lineages in Andalusia was confirmed: 70% against 30% in the earlier migration."
_Put more simply,based on the complete sequences and phylogeography, the split would be 50/50 ancient and modern, except for Andalusia, where it's a 30/70 ancient/historical era mix. 


What I don't think is totally explained is the split between Western and Eastern Andalusia, unless there was a differing migration during the relocations after the Reconquista: 

*"Western Andalusians from Huelva show a distinctive African influence (11.8% of the total mtDNA variability) compared to those from the eastern part of the region (Granada province) where the proportion of African maternal haplogroups is much less pronounced (3.6%)".

*They also used a very extensive collection of mtDna from all around the Mediterranean Basin to do some contour maps not only of U6a and L1b, which are in the body of the paper, but of all the African mtDna.

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## LeBrok

I always thought that there was Mesolithic migration from Africa to Europe through Gibraltar. IIRC there was some West African or SSA signal in WHG or even SHG. I'll try to find a moment tomorrow to read this in full.

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## bicicleur

it looks like remnants of mtDNA indicate pré-Berber migrations, while all Y DNA involved has been whiped out

Berbers are E-M81 and YFull estimates their TMRCA to be a mere 2100 years

apart from E-M81 Berbers and J1 Arab Muslims there is not much other Y DNA left in nortwestern Africa

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## Hauteville

J1 is not exclusively arab, only J1c3 is. Sarno calculated that J1 in south Italy and Sicily is: 



> However, the estimated age for Sicilian and Southern-Italian J1 haplotypes refers to the end of the Bronze Age (32611345 YBP), thus suggesting more ancient contributions from the East.

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## Angela

They're proposing a 50/50 split based on the whole mtDna sequences for a Muslim Era/early migration for Iberia as a whole, and 70/30 for Andalucia, which makes sense given how long it was under Muslim rule and the frequency in part of it.

As for the y that was involved I don't know. The mtDna for U6 as a whole matches the one for E-M81 very well, although I don't know how that can be if E-M81 is that young. 

MtDna U6

Hernandez et al spatial distribution of mtDna U6.jpg

This is the one for U6a:
Hernandez et al spatial distribution of mtDna U6a.jpg

This is L1b:
Hernandez et al spatial distribution of mtDna L1b.jpg

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## Angela

I think the M1 map is very interesting as well.

Attachment 7475

It looks as if it's from a different part of North West Africa. It also looks as if it originally came from the east. I've always thought the whole Phoenician/Carthaginian thing was a bit overblown given the Phoenicians were traders, not colonists, and therefore probably a very male only mediated flow if any, but maybe the Carthaginians were different? That is around the area of the largest Carthaginian settlements isn't it?

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## Hauteville

Weren't carthaginians of phoenician origins?

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## Angela

> Weren't carthaginians of phoenician origins?


Yes, they were, but there's some indication they were a mixed Phoenician/Berber empire. Once the Levant was conquered I think the dynamic changed. They were on their own, as it were, trying to establish a new homeland. 

https://en.wikipedia.org/wiki/Carthage

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## bicicleur

> They're proposing a 50/50 split based on the whole mtDna sequences for a Muslim Era/early migration for Iberia as a whole, and 70/30 for Andalucia, which makes sense given how long it was under Muslim rule and the frequency in part of it.
> 
> As for the y that was involved I don't know. The mtDna for U6 as a whole matches the one for E-M81 very well, although I don't know how that can be if E-M81 is that young. 
> 
> MtDna U6
> 
> Hernandez et al spatial distribution of mtDna U6.jpg
> 
> This is the one for U6a:
> ...


afaik the Muslims that conquered first southern Spain were from the Omajade dynasty, who used to rule in Baghdad, but were expelled by fundamentalistic Muslims
those Omajade fugitives were mainly military men and had little women with them
they took the daughters of the beaten Visigoth rulers as hostages but had children with them
it has been told many Morish rulers dyed their hair to show they were genuine Morish and to conceal their blondness

in the 11 th century the Omajades in Spain were expelled by the fundamentalistic Almohavides from Morroco

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## bicicleur

> I think the M1 map is very interesting as well.
> 
> Attachment 7475
> 
> It looks as if it's from a different part of North West Africa. It also looks as if it originally came from the east. I've always thought the whole Phoenician/Carthaginian thing was a bit overblown given the Phoenicians were traders, not colonists, and therefore probably a very male only mediated flow if any, but maybe the Carthaginians were different? That is around the area of the largest Carthaginian settlements isn't it?


the Phoenicians were traders, but when they were expelled from Lebanon by the Assyrians, they founded Carthago

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## Hauteville

I can not see M1 map.

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## Hauteville

> the Phoenicians were traders, but when they were expelled from Lebanon by the Assyrians, they founded Carthago


Not only Carthage. They established emporiums through the Mediterranean basin.

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## Angela

> I can not see M1 map.


Sorry, let me try it again.

Hernandez et al spatial distribution of mtDna M.jpg

It's probably just gene flow from a slightly different part of northwest Africa.

I was just musing based on the fact that the M1 shows up where it does in Spain and you 
see it in Tunis.

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## Maleth

> the Phoenicians were traders, but when they were expelled from Lebanon by the Assyrians, they founded Carthago


Legend has it that it was internal feud that prompted queen Dido to leave Tyre and search for a new homeland and not foreign occupation

https://en.wikipedia.org/wiki/Dido

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## Maleth

> Yes, they were, but there's some indication they were a mixed Phoenician/Berber empire. Once the Levant was conquered I think the dynamic changed. They were on their own, as it were, trying to establish a new homeland. 
> 
> https://en.wikipedia.org/wiki/Carthage


I presume this is correct also the Berber population in the new Phonetician home (later to be named Carthage) would overwhelm the new phonetician arrivals which at some point must have mixed. i believe that the J1 commonly found in these areas today are not just an addition with the rise of Islam but can also be through very ancient migrations

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## Angela

> Legend has it that it was internal feud that prompted queen Dido to leave Tyre and search for a new homeland and not foreign occupation
> 
> https://en.wikipedia.org/wiki/Dido


I certainly like that version better. :)

Poor Dido, no happy ending for her...

This is my favorite painting of her and Aeneas.

http://harvardmagazine.com/sites/def...F14_09_001.jpg

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## Sile

> J1 is not exclusively arab, only J1c3 is. Sarno calculated that J1 in south Italy and Sicily is:


where did you see that J1c3 was exclusively arab?

There where no arabs in north-africa until after the fall of the Roman empire, so my guess for J1c3 , if arab was still in the arabian peninsula

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## Angela

I also think this map of the distribution of mtDna L3f is interesting.
Hernandez et al spatial distribution of mtDna L3f.jpg



That hotspot in France is pretty close to a hot spot of E-M81 in France. Perhaps this one makes sense as a recent Muslim Era trail that somehow wound up in France. I don't know if it's from the earliest foray up from Spain directly in the early days or the later one that came directly up the coast from the Mediterranean.

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## Angela

> where did you see that J1c3 was exclusively arab?
> 
> There where no arabs in north-africa until after the fall of the Roman empire, so my guess for J1c3 , if arab was still in the arabian peninsula


The Phoenicians were probably descendants of the Canaanites. We don't have ancient dna from the Canaanites, so we can't be certain what they carried. J2a is one good bet, but I wouldn't rule out certain clades of J1. I don't think all of it in the Levant is solely from the period after the "Arab" conquests, just as I don't think all of it in East Africa is that late.

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## Hauteville

> where did you see that J1c3 was exclusively arab?
> 
> There where no arabs in north-africa until after the fall of the Roman empire, so my guess for J1c3 , if arab was still in the arabian peninsula


J1c3 is the dominant Y-DNA of semite speaking peoples.

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## Maleth

> I certainly like that version better. :)
> 
> Poor Dido, no happy ending for her...
> 
> This is my favorite painting of her and Aeneas.
> 
> 
> http://harvardmagazine.com/sites/def...F14_09_001.jpg


Lovely painting that is. :) Always heard the Dido story as a child. Legend (local) also has it here that she visited Malta and contemplated in building her new city here, but it was far too small for her aspirations. Maybe its not a far fetched story considering its believed that it was the Phoenicians who founded the old town of Maleth (todays Mdina) some 700BC and two temples existed already in the entrance of the two largest harbours, Astarte and Melqart. Punic tombs are found all over the Islands.

http://www.mellieha.com/places_inter...punictombs.htm

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## Maleth

> I certainly like that version better. :)
> 
> Poor Dido, no happy ending for her...
> 
> This is my favorite painting of her and Aeneas.
> 
> http://harvardmagazine.com/sites/def...F14_09_001.jpg


That is a lovely painting indeed :). I always heard of the Dido version as a child. Legend (local) also has it that she visited Malta and contemplated in building her new city here, but it was eventually too small for her aspirations. Maybe its not a far fetched story considering that Malta was already an important Phoenician out post with two temples one dedicated to Astarte (todays fort St Angelo no remains left just documentation) and other to Melqart (today just the foundations remain). Also it was the phonecians who founded the first town Maleth (Maleth/Melita/Malta) which gave Malta its name. Also Punic style burial are found all over the Islands.

http://www.mellieha.com/places_inter...punictombs.htm

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## bicicleur

> Legend has it that it was internal feud that prompted queen Dido to leave Tyre and search for a new homeland and not foreign occupation
> 
> https://en.wikipedia.org/wiki/Dido


what's the catch ?

I'm sure the Roman writers must have given a twist to history to demonise Carthago and justifie their own wars

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## bicicleur

> The Phoenicians were probably descendants of the Canaanites. We don't have ancient dna from the Canaanites, so we can't be certain what they carried. J2a is one good bet, but I wouldn't rule out certain clades of J1. I don't think all of it in the Levant is solely from the period after the "Arab" conquests, just as I don't think all of it in East Africa is that late.


The Canaanites were Semites, so E-M123.
I guess J2a was in the Levant before arrival of the Semites, they were ousted by the Semites and then some of them fled to the Lebanon shores.
Ugarit was a Semititc city and the most important harbor for trade between Mesopotamia and the Mediterranean, it was destroyed by the Sea Peoples.
Many Semitic traders from Ugarit fled south toward the villages at the Lebanon shores and mixed with the local J2a.
This mixture of E-M123 and J2a were the Phoenicians. There may have been some J1 with that too.
Anyway if there is still some Phoenician DNA left it should be mainly E-M123 and probably as a second J2a with maybe a small minority of J1.

But I think most of J1 in northern Africa except Egypt is from Muslim Arabs.
The reason is the TMRCA of only 2100 years for E-M81.
At a given point, not so long ago the Berbers must have erased virtualy all other male DNA in the area.
It is known that the conquerer Muslim Arabs came in great numbers to live in or near the cities, while the Berbers remained the majority in the more rural areas.
It would help if we had a breakdown in subclades of the J1 people living in northern Africa.

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## bicicleur

> I also think this map of the distribution of mtDna L3f is interesting.
> Hernandez et al spatial distribution of mtDna L3f.jpg
> 
> 
> 
> That hotspot in France is pretty close to a hot spot of E-M81 in France. Perhaps this one makes sense as a recent Muslim Era trail that somehow wound up in France. I don't know if it's from the earliest foray up from Spain directly in the early days or the later one that came directly up the coast from the Mediterranean.


Before they were beaten by Karel Martel, many Muslim looters roamed through southern France.
They may have raped some women in the process, but they wouldn't have left any mtDNA.

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## bicicleur

> what's the catch ?
> 
> I'm sure the Roman writers must have given a twist to history to demonise Carthago and justifie their own wars


ah, I got it

When Dido sees Aeneas' fleet leaving she curses him and his Trojans and proclaims endless hate between Carthage and the descendants of Troy, foreshadowing the Punic Wars. 

not much this time, the Romans could come up with much more complicated stories to demonise their enemies

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## Maleth

> what's the catch ?
> 
> I'm sure the Roman writers must have given a twist to history to demonise Carthago and justifie their own wars


Oh...I never thought of a catch. Just had history in mind. Internal fueds = to demonaization? If not mistaken the Romans wrote very often about their own.

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## Maleth

> The Canaanites were Semites, so E-M123.
> I guess J2a was in the Levant before arrival of the Semites, they were ousted by the Semites and then some of them fled to the Lebanon shores.
> Ugarit was a Semititc city and the most important harbor for trade between Mesopotamia and the Mediterranean, it was destroyed by the Sea Peoples.
> Many Semitic traders from Ugarit fled south toward the villages at the Lebanon shores and mixed with the local J2a.
> This mixture of E-M123 and J2a were the Phoenicians. There may have been some J1 with that too.
> Anyway if there is still some Phoenician DNA left it should be mainly E-M123 and probably as a second J2a with maybe a small minority of J1.
> 
> But I think most of J1 in northern Africa except Egypt is from Muslim Arabs.
> The reason is the TMRCA of only 2100 years for E-M81.
> ...


I doubt if in this period if each group of people in certain geographical areas were so homogenized with one haplogroup eliminating the other. J1 or its precursor has been around long enough to penetrate neighboring regions from were it was born and vice versa

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## Sile

> ah, I got it
> 
> When Dido sees Aeneas' fleet leaving she curses him and his Trojans and proclaims endless hate between Carthage and the descendants of Troy, foreshadowing the Punic Wars. 
> 
> not much this time, the Romans could come up with much more complicated stories to demonise their enemies


Aeneas was a dardanian and not a trojan...........dardanus was a separate town/area from Ilios( Troy ) , but both where in the huge Troad "region"

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## Angela

If this is correct, the Saracens were established in Narbonne for more than forty years. It seems long enough to send for a few favorite wives.
*Expansion to Provence and Charles Martel*

 


_ Muslim troops leaving Narbonne to Pépin le Bref, in 759, after 40 years of occupation.

__Still, in 734, Umayyad forces (called "Saracens" at the time) under Abd el-Malik el Fihri, Abd al-Rahman's successor, received without a fight the submission of the cities of Avignon, Arles, and probably Marseille, ruled by count Maurontus. The patrician of Provence had called Andalusi forces in to protect his strongholds from the Carolingian thrust, maybe estimating his own garrisons too weak to fend off Charles Martel's well-organised, strong army made up of vassi enriched with Church lands._
_
Charles faced the opposition of various regional actors. To begin with the Gothic and Gallo-Roman nobility of the region, who feared his aggressive and overbearing policy.[6] Charles decided to ally with the Lombard King Liutprand in order to repel the Umayyads and the regional nobility of Gothic and Gallo-Roman stock. He also underwent the hostility of the dukes of Aquitaine, who jeopardized Charles' and his successor Pepin's rearguard (737, 752) during their military operations in Septimania and Provence. The dukes of Aquitaine in turn largely relied on the strength of the Basque troops, acting on a strategic alliance with the Aquitanians since mid-7th century._
_In 737, Charles captured and reduced Avignon to rubble, besides destroying the Umayyad fleet. The brother of Charles, Childebrand failed however in the siege of Narbonne. Charles attacked several other cities which had collaborated with the Umayyads, and destroyed their fortifications: Beziers, Agde, Maguelone, Montpellier, Nimes. Before his return to the northern Francia, Charles had managed to crush all opposition in Provence and Lower Rhone. Count Maurontus of Marseille fled to the Alps.
_
https://en.wikipedia.org/wiki/Umayya...Charles_Martel

See also:
http://www.uq.edu.au/events/event_vi...?event_id=9087

_"The Muslims of tenth-century Provence are a permanent fixture in the realm of folklore and legend. With their largely descriptive accounts, contemporary (9th and 10th century) medieval annals, cartularies, and chronicles form a rather one-dimensional portrait of Muslim settlement in southern France during the tenth century, which effectively thrived with little Christian opposition between the 890s and 970s. While these records provide a general framework of historical events, the long-term consequences of their history are far less familiar. In seeking to understand the Muslims’ impact on southern ‘French’ society, both during and after this period of settlement, this article examines their historical and cultural legacies in modern-French historiography; more specifically, it asks how the ‘Saracen legend’ was first created, why it has remained so prevalent in the historiography on tenth-century Provence, and finally, how it has shaped and perpetuated our understanding and view of the region."

_There is also this:

_ In her paper, "Medieval Cosmopolitanism and the Saracen-Christian Ethos," Marla Segolargues states that in_ _Floire et Blancheflor and Aucassin et Nicolette, two medieval Occitanian romances,the writers work actively to incorporate Islamic culture and its accomplishments into a hybrid communal identity. The hybrid elements of this identity are demonstrated in two ways: first,through the portrayal of mixed couples and second, through depiction of a biculturally constituted landscape and culture. Intercultural relations between the characters are dramatized through the structures of religious conversion. Each romance features a mixed couple, with one and the other, formerly Muslim but converted at some point in the narrative. In each work, the validity of Muslim lover's conversion is probed through an interrogation and a problematization of the process. As these conversions are probed, the hybrid identities of both the characters and their adoptive societies are revealed and accepted. In so doing, these works express cosmopolitan ideals that serve to distinguish Occitanians from their northern and western Christian neighbors. Much of the conflict in each work centers on negotiating an assimilable knowledge of the other within, and as such, the hybrid self."
__
_https://www.academia.edu/2488723/Medieval_Cosmopolitanism_and_the_Saracen-Christian_Ethos

One of the stories is about Aucassin, a young nobleman of _Provence_, who wishes to marry the adopted, converted former slave girl Nicolette.

This is a slightly later period, of course. I just remember things like this from, believe it or not, a university class on the poetry of the troubadours and the whole development of chivalric love, a development that grew out of Arabic poetry and social developments. How things have changed.

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## Drac II

> They're proposing a 50/50 split based on the whole mtDna sequences for a Muslim Era/early migration for Iberia as a whole, and 70/30 for Andalucia, which makes sense given how long it was under Muslim rule and the frequency in part of it.


The authors of this paper consider "recent" or "historic times" anything from Roman times on, so it is not necessarily "Muslim Era".

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## bicicleur

interesting
are the J1 in southern France J1-P58 or not?

the Merovingian kings didn't have an army to oppose intruders
it is Karel (Charles) Martel, servant of the Merovingian court who organised and trained a new army and beat the Muslims in Poitiers in 732
Karel Martel had the support of the pope and in 751 the pope crowned Pepijn (Pépin), son of Karel Martel as king of France
it was the end of the Merovingian dynasty and the beginning of the Karolingian line

I knew there were still many Saracene raids but I didn't know the Saracens did have some strongholds on French soil, eventualy with the support of some local nobility

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## Drac II

The authors mention Moorjani et al. as if it gives support to the idea that the "recent" mtDNA markers may be from Islamic times, but as they themselves seem aware, that study dated what they considered to be "sub-Saharan African" DNA in southern Europe to an average of about 55 generations ago (so around 1600 years), which is in fact pre-Islamic. Not even Islam itself existed back then.

They also tried to use Hellenthal et al. 2014 as if it supports a date range from 890 to 1754 CE for "recent" admixture in Southern Europe, but the "recent" admixture range for Spain in that study is from 1026 to 1670 CE (so 11th to 17th century AD), there is no admixture event that could have happened in the 700s, 800s, 900s or 1700s for Spain in that study. Italy in that study does have "recent" admixture events that they considered could have happened in the 700s-900s and later, though. However, they neglect to point out that Hellenthal et al.'s "recent" admixture ranges for Italy (except West Sicily) also extend back to Roman times, so they are not necessarily saying that it has to be due to "the Islamic expansion and slave trade" for Italy's case.

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## Hauteville

> If this is correct, the Saracens were established in Narbonne for more than forty years. It seems long enough to send for a few favorite wives.
> *Expansion to Provence and Charles Martel*
> 
>  
> 
> 
> _ Muslim troops leaving Narbonne to Pépin le Bref, in 759, after 40 years of occupation.
> 
> __Still, in 734, Umayyad forces (called "Saracens" at the time) under Abd el-Malik el Fihri, Abd al-Rahman's successor, received without a fight the submission of the cities of Avignon, Arles, and probably Marseille, ruled by count Maurontus. The patrician of Provence had called Andalusi forces in to protect his strongholds from the Carolingian thrust, maybe estimating his own garrisons too weak to fend off Charles Martel's well-organised, strong army made up of vassi enriched with Church lands._
> ...


There was also the Emirate of Fraxinet.

https://en.wikipedia.org/wiki/Fraxinet

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## bicicleur

> There was also the Emirate of Fraxinet.
> 
> https://en.wikipedia.org/wiki/Fraxinet


nice
2 sides of the same story
but still a lot of mystery

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## Angela

The authors _don't at all_ rely on the Moorjani et al or Hellenthal et al papers. They _rely_ on the phylogeography of the complete mitochrondrial sequences, a Founder Analysis (FA) methodology, and a Bayesian analysis. It's obvious in a close reading of the discussion and results sections as well as the methods section and the supplementary info section. That's what the paper is about.

What they do is _contrast_ the results of Moorjani et al and Hellenthal et al, which propose an almost entirely late date for the arrival of SSA autosomal input, with their results from uniparental marker analysis, and make a case for the fact that there is still a role of this kind of analysis, and that it may indeed be more accurate as to timing, and can serve to correct the failures of current methods of autosomal analysis with regard to problems with the dating of admixture. I happen to agree with them for what it's worth.

This is from the paragraph where they mention the Moorjani and Hellenthal papers.
"Date estimates for migrations across the Mediterranean Basin and the Sahara desert based on mtDNA data do not generally match the ones inferred by genome-wide information, these last tending to be considerably younger.

The current limitation on date estimates for admixture based on genome wide diversity, due to the phenomenon of recombination, shows the continued usefulness of uniparental markers to shed light on the human evolution history, spanning 200,000 years. In particular, our work supports the existence of an ancient, frequently denied, bridge connecting the Maghreb and Andalusia, located just across the Strait of Gibraltar but also possibly in a more eastern position with a pier on the small island of Alboran."


As for the precise causes of this "modern" admixture, this is what they have to say, and it_ is_ broader than just the Muslim era. However, the connection to Roman domination is not explored, beyond a reference to the joint administration of Northwest Africa and Iberia during the Roman period. Rather, the focus, other than the Islamic Era migration, is on the Atlantic slave trade.

"The recent migration peaks identified in the *FA of U6 and L lineages could be associated with the Islamic rule of Iberia and the slave trade period, respectively.* The HVS-I FA attributes a comparatively lower proportion of these recently introduced sequences into Iberia when compared with the post-glacial one: 1/3 vs 2/3, respectively. The analysis performed by [26] on the *phylogeography* of *L sequences* observed in* Iberia* and* remaining Europe* pointed to *65%* of its introduction in recent times (*Romanization period, Islamic expansion and Atlantic slave trade*), and 35% at older times, as earlier as 11ky. The* complete sequence based FA for U6 agrees more with these results, attributing a half-half proportion of sequences in both periods*, showing the higher resolution of complete mitogenomes. The *historical-based expectation of a higher proportion of newly introduced U6 lineages in Andalusia was confirmed: 70% against 30% in the earlier migration."
*
As to this Spanish slave trade admixture they also have this to say:


"In contrast,* only 24.2% (f1) and 20.3% (f2) of the L-lineages that entered Europe in the historic period match the ones that arrived in North Africa*, suggesting mainly independent introduction routes. As our data suggest, *an entrance of these lineages into Europe directly from sub-Saharan Africa in the historic period is likely related with the slave-trade in Europe."

*They seem to be proposing that while L1 is a mixed bag in terms of time of arrival, the great majority of the specific L2 and L3 clades that entered Iberia is the result of the trans-Atlantic slave trade.

"The position of the Iberian samples within *L2 and L3* phylogenies is of considerable complexity. In most cases, Iberian sequences do not cluster with the North Africans sequences but rather with those from the Middle East or Europe. Interestingly, the Spanish samples showed a tendency to group together in the trees although they originate from very distant peninsular regions. *It is likely that some of these Spanish mitochondrial sequences may correspond to descendants of emigrants to Hispanoamerica who returned to the Peninsula after mixing with Afro-American people.*"

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## Drac II

The authors don't try to rely only on autosomal analyses, but they do try to find support for their opinions in them:

"Date estimates for migrations across the Mediterranean Basin and the Sahara desert based on mtDNA data do not generally match the ones inferred by genome-wide information, these last tending to be considerably younger. A recent study [58] estimated that the migration from sub-Saharan Africa into North Africa occurred 40 generations ago (≈1.2 ky), matching trans-Saharan slave trade, while their estimate for the back-to-Africa settlement of North Africa (comparison with Near Eastern/European pools) was before 12 ky. *On the other hand, [**59**] showed that the 1–3% African ancestry observed in southern Europeans had an average mixture date of 55 generations ago,* matching North African flow at the end of the Roman Empire and subsequent Arab migrations. And even improved methods, allowing to disentangle between several migration events, such as the one developed in [49], c*onduct to dates of sub-Saharan African admixture in the range 890 to 1754 CE, again pointing to the Islamic expansion and slave trade.*"

As pointed out earlier, the results of those two studies they referred to actually do not exactly support what they say. Moorjani et al. estimated African DNA in Southern Europe to be older than the one in the Near East, dating to several centuries before Islam arrived anywhere in Europe. And Hellenthal et al. 2014 does not propose any such admixture dates as 890 or 1754 for Spain anywhere, so the authors of the mtDNA paper must have other parts of the Mediterranean in mind as well when they wrote that. For Italy such dates as the 800s were given as possible in Hellenthal et al., but they also can extend back to Roman times, so it is hardly clear-cut that it has to be because of "Islamic expansion and slave trade".

As for this quotation:

"The results for the introduction of L lineages *into Iberia or Europe* are similar to those, that is, a prehistoric episode would be the main contributor to the sub-Saharan presence in Mediterranean Europe and Iberia (Fig 4B and 4C). A deep inspection of the L-haplogroup founders introduced in North Africa showed that most of them (including basal L1b) reached the area at the older period of migration (Fig L in S1 File). Compared with those L lineages that entered into Mediterranean Europe, the L1b root haplotype clearly dominates the pool of L lineages. More importantly, 79.5% (_f1_) and 81.5% (_f2_) of the lineages that suggest an early Holocene (the Holocene represents the last 11,700 calendar yr before AD 2000 [47]) entrance in Europe also shows the same period of arrival into North Africa, indicating that the migration sequence was likely sub-Saharan Africa—North Africa—Mediterranean North. In fact, we detected genetic closeness between North African and European L-mitogenomes, especially lineages shared with Iberians. In contrast, only 24.2% (_f1_) and 20.3% (_f2_) of the L-lineages that *entered Europe* in the historic period match the ones that arrived in North Africa, suggesting mainly independent introduction routes. As our data suggest, an entrance of these lineages *into Europe* directly from sub-Saharan Africa in the historic period is likely related with *the slave-trade in Europe.*"

It refers to European slave trade in general, not just the Iberian ones, and its possible role in introducing some of these lineages.

As for the preciseness of their mtDNA analysis methods, the fact that they can't more exactly pinpoint whether the "recent" mtDNA lineages were introduced during Roman, medieval or more modern times (including descendants of immigrants from the Americas) pretty much casts doubts that they are better or can shed more light than the autosomal methods.

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