No Iberian in Iberomaurusian

After almost a century of controversy on the matter, it seems that archaeogenetics solved the riddle. Not in the sense I thought it would but it did anyhow.

Marieke van de Loosdrecht et al., Pleistocene North African genomes link Near Eastern and sub-Saharan African human populations. Science 2018 (pay per view). DOI:10.1126/science.aar8380 -- See also supplementary materials (free but hard to find). 

Ancient DNA samples from Taforalt (Iberomaurusian or Oranian culture, Upper Paleolithic of North Africa) show no trace of Paleoeuropean ancestry (WHG), however they show strong affinity to West Asians of Palestinian type, showing also some significant amount of African Aboriginal ancestry, probably closer to East African Hadza and Sandawe and ancient Mota than to West African types. The result is something roughly similar to Afars but not quite the same in any case. 

Fig. S8 - Taforalt individuals on the top PCs of present-day African, Near Eastern and South European populations.

Fig. S11 - ADMIXTURE results for a few informative K values.

So the conclusion must thus be that the Eurasian influence in North African Upper Paleolithic (call it Iberomaurusian, Oranian or my personal unorthodox preference: Taforaltian) arrived from West Asia with whichever intermediate stage in Egypt and Cyrenaica, where that influence is quite apparently much older in the archaeological record. This seems contradictory to the chronology of Taforaltian, with Western sites producing older radiocarbon dates but the genetic data seems overwhelming. 

I must say I wish they would have contrasted with older (and available) Paleoeuropean samples than WHG (Epipaleolithic) but I guess that some WHG influence would have shown up if there was some older European influx because the various Paleoeuropean layers are not disconnected. But it is still something someone should test, just in case. 

Haploid DNA

The Taforalt sample was rich in mtDNA U6a, with also one instance of M1b:

All six male samples carried Y-DNA E1b1b, with most of them being well defined as E1b1b1a1-M78 (see table S16 for details).


Related: Luxmanda: a 3,000 years-old proto-Horner in Tanzania.

North African Neolithic was influenced by Europe... and European Chalcolithic by Iberian Neolithic

Or so it seems considering the data of Fregel et al., a study I have in my to-do list for some time and that I don't see cited often or ever at all.

Rosa Fregel et al., Neolithization of North Africa involved the migration of people from both the Levant and Europe. BioRxiv 2017 (pre-pub). DOI:10.1101/191569

The critical piece is probably this selection from Admixture results but which repeats over and over through the study with many more analyzed populations from all West Eurasia and North Africa:

We see how KEB (Morocco Neolithic) is a mix of European Neolithic intermediate between Iberia (purple) and Sardinian (blue) on one side and, on the other, something like Mozabites (not shown in this detail, cream). TOR is a new Neolithic sample from Andalusia.

Another ancient Moroccan sample IAM (pre-Neolithic, not shown here either) is fully cream-colored like mostly are modern Mozabites. 

Interestingly we see for the first time the emergency of a purple-colored component that differentiates Iberian Early Neolithic from the rest (although this does not happen at lower K-values, so they are still related), a component that, in the MNChL (Middle Neolithic and Chalcolithic) period, somehow appears as dominant in Italy (no data for earlier times) and becomes quite dominant in Central Europe. 

This is intriguing to say the least. It must be said that modern Sardinians and Basques (these probably, not labeled) are low in the purple component, although less than other populations, and that somehow the Early Neolithic (blue) component made a comeback:

I do not want to over-interpret all this (autosomal genetics are not an exact science) but, judging on KEB, the purple component is not just a generic southern branch (Cardium Pottery) distinction but something specifically Iberian or Italo-Iberian. The matter needs more research but it is in any case very intriguing that the purple component seems to expand from Iberia or somewhere nearby (France?, Italy?) in the period leading to the Chalcolithic, a most critical one in the formation of the genetics of Europe.

There is a also a little hoard of DNAmt and Y-DNA, with G2a-M201 (in Europe), E1b-L19* (in pre-Neolithic North Africa) and T-M184 (in Neolithic North Africa) in the patrilineal side and quite a bit of varied K1a in the matrilineal one, as well as JT (also in both shores) and U6 and M1 in North Africa.

Worth reading and keeping in mind, no doubt.

Oldest evidence for cooking in pots comes from Libya

Quickies

Kambiz Karami at Anthropology.net (a most interesting blog to follow) mentioned this week that Libyan pottery remains, maybe as old as 10,000 years ago, have provided the oldest evidence of vegetable cooking of we know of, at least in pots (see below for the disclaimer). The remains indicate cooking of vegetables and meat and are associated to pictures of people gathering plants, as well as grinding stones (hand mills) with remains of such provisions.

Ref. Nature Plants.

Oddly enough, just two days later, he contradicted himself, mentioning that the charred remains of nuts and seeds from Palestine, dated from 780,000 years ago, actually provide the oldest evidence of vegetable cooking, even if it's clearly a more primitive way of cuisine and not yet at all the "refined" stew of ancient Libyans, which won them three stars in the Paleo-Michelin Guide for Nomads and five pitchforks in Popular Mesolithic Cuisine. 

Ref. PNAS.



Update (Jan 4): origin of Libyan and West Asian pottery could be Sudan

Jm8 mentions it in the comments, referencing to Anthromadness. I would need more data to judge (the ref. paper is behind paywall) but  it does sounds as probably correct to me, being Sudan/Nubia one of the earliest Mesolithic areas in the Western half of the Old World, one that is way too often and unfairly neglected but that definitely influenced the Levant prior to the development of Neolithic proper (what should explain a lot of things both linguistic and genetic).

Ancient aboriginal DNA from El Hierro (Canary Islands)

The island of El Hierro (lit. The Iron) is one of the westernmost of the Canary Islands. I have never visited but it seems to be very beautiful, far enough from the Sahara to enjoy a warm yet humid climate. It wasn't far enough to remain uninhabited however and now we get to know something more about its original dwellers, generally known as Guanches (although technically this name only applied to the inhabitants of La Palma originally).

Alejandra C. Ordóñez et al., Genetic studies on the prehispanic population buried in Punta Azul cave (El Hierro, Canary Islands). Journal of Archaeological Science 2016. Pay per view → LINK [doi:10.1016/j.jas.2016.11.004]

Abstract

The aim of this study was to establish the genetic studies of the population from one of the most important known aboriginal funerary spaces of the island of El Hierro (Canary Islands), the Punta Azul cave, which harbors remains of 127 individuals. Sixty-one adult tibiae were examined, 32 left and 29 right. Radiocarbon dating yields an antiquity of 1015–1210 AD. We have obtained an overall success rate of 88.5% for the molecular sexing, and of 90.16% for the uniparental markers. Short tandem repeats (STR) profiles were also possible for 45.9% of the samples. This performance is a consequence of the good conservation of the bones in their archaeological context. The mtDNA composition of the sample is characterized by the complete fixation of the H1-16260 lineage. These results can be explained by a mixture of consecutive founding events, a bottleneck episode at the beginning of the colonization and/or as a consequence of genetic drift. Paternal lineages were also affected by these processes but in a less acute way. These differences lead us to propose social behaviors as an explanation for this difference. The maternal transmission of the lineages, mentioned in ethnohistorical sources of the Archipelago, could be an explanation. These results could be in agreement with endogamous practices, but the autosomal STR results indicate a relative high diversity. These results have allowed us to characterize the Punta Azul cave population and see the way in which geographical isolation, the process of adaptation and specific social behaviors affected the aboriginal population of the Island.

And now the interesting stuff, the findings:

As should be expected, the remote and rather small western island, which is the most direct threat to the security of the USA and the Western World in general because of the danger its volcano may slide into the Ocean and cause a huge tsunami of devastating consequences, something that no nuclear arsenal can do anything against, shows clear indication of very strong genetic drift relative to its relatives of the larger islands, caused either by founder effects and/or endogamous drift. Otherwise it is within the general patterns for the pre-colonial islanders. 

We see a lot of likely mtDNA H and certain Y-DNA R1b1a2. The Canary Islands were settled c. 1000 BCE (11,000 HE) by people arrived from what is now Morocco and the Western Sahara. They seem to retain a somewhat archaic genetic pool, relatively rich in Europe-related genetic elements, not so abundant at all today in Northwest Africa anymore, and by this I mean of course very especially Y-DNA haplogroups I and R1b. While we can still find some R1b in NW Africa, haplogroup I is almost impossible to find nowadays, yet it was present in the Canary Islands prior to European conquest and nearly disappeared afterwards (so it's definitely not a colonial input, accidentally misidentified, not at all). 

See also: Leherensuge: Ancient Guanche Y-DNA.

Iberomaurusian ancient mtDNA

This is an issue that has lingered for a long time in the online anthropology circles. Once upon a time (2005) there was online a presentation in which a good deal of mitochondrial DNA (HVS-I) sequences from the key Iberomaurusian (or Oranian) culture site of Taforalt (North Morocco) were (not too formally) published. Eventually this presentation became almost impossible to find... however now Rym Kefi (the original author if I remember correctly) and colleagues have got back to resurrect that important data set from oblivion, plus sequences from Afalou cave as well.

Rym Kefi et al., On the origin of Iberomaurusians: new data based on ancient mitochondrial DNA and phylogenetic analysis of Afalou and Taforalt populations. Forsenic Sciences Research 2016. Freely available at the time & space of writing this → LINK [doi:10.1080/24701394.2016.1258406]

Abstract

The Western North African population was characterized by the presence of Iberomaurusian civilization at the Epiplaeolithic period (around 20,000 years before present (YBP) to 10,000 YBP). The origin of this population is still not clear: they may come from Europe, Near East, sub-Saharan Africa or they could have evolved in situ in North Africa. With the aim to contribute to a better knowledge of the settlement of North Africa we analysed the mitochondrial DNA extracted from Iberomaurusian skeletons exhumed from the archaeological site of Afalou (AFA) (15,000–11,000 YBP) in Algeria and from the archaeological site of Taforalt (TAF) (23,000–10,800 YBP) in Morocco. Then, we carried out a phylogenetic analysis relating these Iberomaurusians to 61 current Mediterranean populations.

The genetic structure of TAF and AFA specimens contains only North African and Eurasian maternal lineages. These finding demonstrate the presence of these haplotypes in North Africa from at least 20,000 YBP. The very low contribution of a Sub-Saharan African haplotype in the Iberomaurusian samples is confirmed. We also highlighted the existence of genetic flows between Southern and Northern coast of the Mediterranean.

The bulk of the data is this (tables 1 and 2):

Notice that all sequences are "old style", i.e. HVS-I only, and that's why haplogroup assessment is not always 100% certain, with particular emphasis on the "CRS" sequences, which have been proven in some ancient cases to belong not to the modern normal (H1, rarely H2 o other H) but to a very rare modern haplotype within U*, AFAIK only preserved in Asturias but somewhat common in the Magdalenian of South Germany. 

In any case, that they produce 100% safe H1, H6, H14, etc. and, even more surprisingly, J and T2 variants, should make people think about the hidden Paleolithic mtDNA diversity in the wider Mediterranean area. And by this I do not just mean North Africa but also all Southern Europe, including many areas that can't be considered part of the Mediterranean basin, such as most of the crucial Franco-Cantabrian Region, the most densely populated province, by far, of post-LGM Upper Paleolithic Europe, source of several cultural waves (including the Solutrean one, almost certainly at the origin of Iberomaurusian), yet outrageously neglected by research.

Table 5 and other materials in the paper also deal with which modern populations appear closest to the Iberomaurusian ancient mtDNA pool and these are:

1. Tuscans (0.00090)
2. Catalans (0.00134)
3. Galicians (0.00223)
4. Sicilians (0.00377)
5. El Alia (0.00699)
6. Valencian (0.00787)
7. Matmata (0.00788)
8. Slougia (0.00831)
9. Jerba Berbers (0.00934)

Figures in brackets are FST distances, the smaller the closer the match, statistically speaking. Populations in cursive are from Northwest Africa.

That is I'd say quite surprising because we tend to think of those near matches as quite impacted by Neolithic and post-Neolithic inflows, at least judging on other recent research. It seems that the debate on the origin of modern Europeans and North Africans refuses to come to a close, as different aspects of the evidence available may be somewhat contradictory. What do you make up of all this?

See also:

• Leherensuge: 2009 review of mtDNA H studies (some re. North Africa).
• I'd emphasize Sykes et al 2005 on Portuguese Epipaleolithic and Neolithic mtDNA also rich on "CRS" and other H sequences, because it provides another anchor point for the presence of mtDNA H in pre-Neolithic Europe.
• But for North African mtDNA studies Enafaa 2009 is pretty much seminal, showing that SW Europe related haplogroups H1, H3, H4 and H7 are absolutely hegemonic in the region's H.
• MtDNA H in Magdalenian Cantabria
• MtDNA U6 in Aurignacian from Romania
• On South Iberian Solutrean (surely at the origin of Iberomaurusian)
• And in general the Oranian label (alternative name of Iberomaurusian)

UPDATE: single marker inferred lineages (not always the haplogroup is reported correctly):

Down in the comments' section, Capra questions with good sense the certainty of inferred haplogroups. The most clear one is TafV27, claimed to be H6a1a8, which cannot actually be that (would need another HVS-I marker defining H6, which is missing), however this means that it must be HV0 and possibly V, a lineage that appears (per the aDNA literature) in the European Neolithic, already quite towards the West, out of nowhere, and that "nowhere" should not be West Asia, where it has never been sequenced, unlike K. 

I'm therefore reviewing here all the single-marker inferred haplogroups as carefully as possible, please double-check them and report if I seem to be committing any error:

• AF22B - 16126C (reported as JT or H14b1) - JT(xJ,xT) or H14b1 indeed, notice that JT* was probably also one of the lineages described (also by the HVS-I method) for Nerja cave (Solutrean, same time frame), just across Alboran Sea (cf. Fernández-Domínguez 2005). Whether is one or the other or even something else, it's almost certain that the lineage was shared across the sea between Europe and Africa, what just makes total sense for this culture.
• Taf55-IB - 16239T (reported as H1) - I actually get specific H1 variants (H1bf'bg'bh'ch specifically) or H17c. H sublineages in any case, unless it is an extinct or unreported R* lineage, most unlikely.
• TafV27 - 16298C (reported as H6a1a8) - must be HV0, possibly even V, a distributed Euro-African lineage with three hotspots: Kabylia, Catalonia and Lappland. Per Caramelli 2003 and successor studies, this lineage was already present in Italy (Pagilicci cave) in the Gravettian era.
• TafVIII - 16223T (reported as U4a2b) - It cannot be U4 at all because it lacks the HVS-I mutation 16356C. In fact the reverse 16223C mutation defines macro-haplogroup R, and I can't find any other matches within R, so it must be L3(xR), maybe N(xR) but maybe M or other L3. The possibilities are many, for example: N1, N2, N9, N11, etc. within N, a lot of possibilities within M, and also several within L3e'i'k'x(xL3x). The only thing we should be certain here is that it is not R and also not upstream of L3 (that's how good as it gets with HVS-I methodology, really). 
• TafXXV3 - 16126C (reported as H14b1) - as with AF22B, it can be JT(xJ,xT) or H14b1 and again I must emphasize that there is an extremely good chance that this matrilineage had relatives in Iberia (Nerja) within the Solutrean culture.

In case you want to re-check, the best resource is of course PhyloTree.

IMPORTANT CAVEAT: the above update was done assuming they had fully sequenced the HVS-I, which is not that long, but it seems that in many cases (second column of the tables above) they have only sequenced up to the 16317 locus, what makes prediction of haplogroup even harder. That would explain their H6a1a8 prediction, although there's no way they can exclude the much more common HV0 (H6 is unheard of in North Africa). Thanks again to Capra for pointing that out.

Paleolithic European mtDNA lineage U5b2c1 in Carthaginian man

Quickies

Elizabeth A. Matisoo-Smith et al. A European Mitochondrial Haplotype Identified in Ancient Phoenician Remains from Carthage, North Africa. PLoS ONE 2016. Open access → LINK [doi:10.1371/journal.pone.0155046]

Abstract

While Phoenician culture and trade networks had a significant impact on Western civilizations, we know little about the Phoenicians themselves. In 1994, a Punic burial crypt was discovered on Byrsa Hill, near the entry to the National Museum of Carthage in Tunisia. Inside this crypt were the remains of a young man along with a range of burial goods, all dating to the late 6th century BCE. Here we describe the complete mitochondrial genome recovered from the Young Man of Byrsa and identify that he carried a rare European haplogroup, likely linking his maternal ancestry to Phoenician influenced locations somewhere on the North Mediterranean coast, the islands of the Mediterranean or the Iberian Peninsula. This result not only provides the first direct ancient DNA evidence of a Phoenician individual but the earliest evidence of a European mitochondrial haplogroup, U5b2c1, in North Africa.

The lineage is the same one as La Braña 1, an Epipaleolithic man buried in a cave at the mountains NE of León. Its presence on a Carthaginian from the 6th century BCE almost certainly indicates that he had native Iberian maternal ancestry, that his family had arrived to Carthage from Gadir (modern Cádiz) or some of the other Phoenician colonies of Andalusia. The location of his burial at the acropolis and the wealth of the burial goods indicate that he belonged to the highest social elite of the still incipient Carthaginian empire. He has been nicknamed "Ariche" (the loved one) and his face reconstructed as you can see in this blog.

Thanks to Jamel of Lapurdi for the reference an a nice related discussion.

Algerian complex genetics

This is a rather interesting study that deals with the genetics of the Republic of Algeria, with several new samples.

Asmahan Bekada, Lara R. Arauna et al. Genetic Heterogeneity in Algerian Human Populations. PLoS ONE 2015. Open access → LINK [doi:10.1371/journal.pone.0138453]

Abstract

The demographic history of human populations in North Africa has been characterized by complex processes of admixture and isolation that have modeled its current gene pool. Diverse genetic ancestral components with different origins (autochthonous, European, Middle Eastern, and sub-Saharan) and genetic heterogeneity in the region have been described. In this complex genetic landscape, Algeria, the largest country in Africa, has been poorly covered, with most of the studies using a single Algerian sample. In order to evaluate the genetic heterogeneity of Algeria, Y-chromosome, mtDNA and autosomal genome-wide makers have been analyzed in several Berber- and Arab-speaking groups. Our results show that the genetic heterogeneity found in Algeria is not correlated with geography or linguistics, challenging the idea of Berber groups being genetically isolated and Arab groups open to gene flow. In addition, we have found that external sources of gene flow into North Africa have been carried more often by females than males, while the North African autochthonous component is more frequent in paternally transmitted genome regions. Our results highlight the different demographic history revealed by different markers and urge to be cautious when deriving general conclusions from partial genomic information or from single samples as representatives of the total population of a region.

Y-DNA frequencies

Supplementary Table 2: Y chromosome haplogroup frequencies among the studied populations (% in parentheses)
Population	Algiers1	Oran1	Reguibate1	Zenata1	Mozabite2	Oran3	Algiers4	Tizi Ouzou4
Abreviations	ALG1	ORN1	RGB	ZNT	MZB	ORN2	ALG2	TZO
Number of individuals	26	80	60	35	20	102	35	19
A -M91	(-)	1 (1.25)	(-)	(-)	(-)	(-)	(-)	(-)
C-M216	(-)	1 (1.25)	(-)	(-)	(-)	(-)	(-)	(-)
E1a-M33	1 (3.84)	(-)	(-)	1 (2.86)	(-)	(-)	1 (2.86)	(-)
E1b1a-M2	(-)	8 (10)	2 (3.33)	8 (22.86)	2 (10)	8 (7.84)	(-)	(-)
E1b1b1a-M78	4 (15.38)	2 (2.50)	(-)	1 (2.86)	(-)	6 (5.88)	4 (11.43)	(-)
E1b1b1b -M81	14 (53.85)	33 (41.25)	48 (80)	17 (48.57)	16 (80)	46 (45.10)	14 (40)	9 (47.37)
E1b1b1-M35	(-)	3 (3.75)	3 (5)	(-)	(-)	(-)	1 (2.86)	2 (10.53)
E2 -M75	(-)	1 (1.25)	(-)	(-)	(-)	(-)	(-)	(-)
F -M89 (xJ, K, Q, R1)	2 (7.69)	4 (5)	1 (1.67)	(-)	(-)	(-)	4 (11.43)	2 (10.53)
J -M304 (xJ2)	5 (19.23)	18 (22.50)	6 (10)	4 (11.43)	(-)	23 (22.55)	8 (22.86)	3 (15.79)
J2 -M172	(-)	1 (1.25)	(-)	(-)	(-)	5 (4.90)	2 (5.71)	(-)
K -M9	(-)	(-)	(-)	(-)	(-)	(-)	1 (2.86)	(-)
Q -M242	(-)	2 (2.50)	(-)	1 (2.86)	(-)	1 (0.98)	(-)	(-)
R1 -M173	(-)	6 (7.50)	(-)	3 (8.57)	2 (10)	13 (12.75)	(-)	3 (15.79)
Y Haplogroup Diversity GD (h +/- sd)	0.6677 +/- 0.0806	0.7674 +/- 0.0356	0.3520 +/- 0.0757	0.7092 +/- 0.0625	0.3579 +/- 0.1266	0.7245 +/- 0.0325	0.7782 +/- 0.0499	0.7427 +/- 0.0831
1 Present study
2 Shi et al. 2010
3 Robino et al. 2008
4 Arredi et al. 2004

The most common lineage is E1b-M81, which is centered around Morocco and has a mostly NW African distribution. The Reguibate sample (Arabic speakers from near Southern Morocco and West Sahara) shows extremely high frequencies (80%) of it. This is also true of the Mozabites. Otherwise the frequencies range between 40% and 54%.

Tropical African lineages are mostly represented by E1b-M2, which peaks among the Zenata Berbers of the Southern Atlas and Northern Sahara but has also some notable presence in Oran, Mozabites (North Sahara) and Reguibate (West Sahara). However these lineages are nearly absent in the Northeast Kabyle Berbers (Tizi Ouzu) and only have a token presence in Algiers (E1a). 

E1b-M78, a lineage centered in NE Africa, seems to peak in Algiers, with low frequencies in Oran and effectively absent in other populations. 

J1, presumably the same as J(xJ2), is strongest in the coast (Algiers, Oran) but has significant frequencies in other populations (excepted Mozabites). 

J2, although quite rare, is worth mentioning because its presence may indicate areas of true Arabic settlement (of course J1 is more common in Arabia but it is unthinkable that one goes without the other in such a recent time frame). It seems that Oran has the strongest such settlement, although some is also apparent in Algiers.

R1 peaks among Kabyles (16%) and is also present in Oran and among the Mozabite and Zenata Berbers. Sadly it is not analyzed what fraction of it is R1b-M412 (Western European) or R1b-V88 (Afro-Mediterranean), as both lineages have been detected in North Africa in previous studies but almost certainly have different histories. 

Other F is quite intriguing. The few Q and K* individuals are within expectations (at least my expectations) but there are a lot of F* people, notably in Kabyle and Algiers that are most intriguing. Are they within haplogroup G or is it something else? G reaches almost 10% in Egypt but previous studies had not found more than 6% in NW Africa (Bouhria Berbers, see here).

Update (Nov 4): Chris makes a very interesting suggestion in the comments section about all this F*: what if it is (partly or in full) haplogroup I, a typical European Y-DNA lineage that is clearly rooted in the Paleolithic of the region? The lineage has been documented in ancient Berbers from Canary Islands and, for what Chris says, also in Sudan. It would make perfect sense if it was also present among modern NW Africans, being consistent with other genetics that seem to originate in Paleolithic Europe (~30% of mtDNA, a good share of autosomal DNA, maybe also part of the Y-DNA R).

Mozabites are close to "pure North Africans"

Autosomal analysis shows that this Berber population of the Algerian Atlas has the lowest range of admixture form any external source, be it Europe, West Asia or Tropical Africa. Some individuals appear extremely unadmixed.

Fig 3. Plots for the analysis of genome-wide SNPs.

PC analysis (upper figures) based on autosomal data, and X-chromosome SNPs. ADMIXTURE proportions (bottom figures) at k = 2,3, and 4 based on autosomal data and X-chromosome SNPs. Algeria, stands for general Algerian sample [3]; Mozabite, stands for the Algerian Berber Mozabites [32]; and Zenata, stands for Algerian Berber Zenata (present study).

X-chromosome conundrum

It is not common that genetic studies analyze the X-chromosome. A reason is probably that its interpretation can be confusing. Intuitively it seems true that the X chromosome is passed down by a mostly female line but this is not really correct, as (ignoring partial recombination) a man can have an X chromosome from either the maternal grandfather or grandmother, while a woman will have one from her father and another from the mother. Ironically only a woman's father-inherited X-chromosome can be automatically traced to a woman two generations back: that of the paternal grandmother. Complicated, right?

As probably apparent in fig. 3 above but made more clear in fig. 4 below, the study detected differences in autosomal (overall) ancestry and X-chromosome one.

Fig 4. Correlation plots of the ancestry proportions at k = 4 in the ADMIXTURE analysis comparing autosomes and X-chromosome SNPs.

North African, sub-Saharan, Middle Eastern, and European ancestry proportions are shown in different plots. Solid black lines represent linear correlations between autosomal and X-chromosome components.

The authors interpret these results as indicating female bias in the European and West Asian components. This may be true at least in the European case because it correlates well with the differential between European mtDNA (~30%) and Y-DNA (<10%), which suggests that European ancestry used to be more important in the past and that male-biased migrations (Capsian culture is probably one of the culprits) altered this. 

But is it also true for the West Asian ancestry? I can't say, really. I remember a study from a decade ago (don't have the reference right now, sorry) or so that showed that in a Colombian coastal town, X-chromosome ancestry was almost only European, while mtDNA was instead almost exclusively Native American, and that it should be interpreted as continuous influx of men from Europe, marrying local women, who managed to retain, generation after generation, the aboriginal mtDNA (which does never leave the strict maternal line) but not the X-chromosome line, once and again altered by male immigrants. 

I don't really dare to subscribe the authors' interpretation without a more nuanced analysis, analysis that I don't feel able to perform myself at the moment either. If they are correct, anyhow, it means that there were important male-biased demographic expansions of African specific origin, either in NW Africa itself (what could well be supported by the vigor of E1b-M81) or in NE Africa prior to migration to the West within Capsian. Or both. 

Mitochondrial DNA data

In case anyone wants to try their luck at this complicated analysis (North Africans are indeed a complex and most intriguing population), I'm adding here the raw mtDNA table:

Supplementary Table 5: mtDNA haplogroup frequencies (%) distribution among Algerian populations
Populations	Algiers	Oran	Zenata	Reguibate	Oran (Bekada et al. 2013)	Mozabite (Corte-Real et al. 1996)
Abbreviation	ALG	ORN1	ZNT	RGB	ORN2	MZB
Number of samples	62	93	73	108	240	85
H/HV	19.35	35.48	12.33	30.56	30.83	23.53
HV0	4.84	2.15	5.48	6.48	3.75	8.24
I	1.61	-	1.37	-	0.83	-
J (16069 16126)	14.52	3.23	2.74	0.93	3.33	3.53
K (16224 16311)	-	4.30	4.11	3.70	1.67	-
L	-	-	-	0.93	-	-
L0	1.61	3.23	1.37	-	0.42	-
L1b	1.61	2.15	9.59	6.48	3.75	-
L1c	-	-	1.37	0.93	0.83	-
L2	-	-	5.48	4.63	0.83	-
L2a	9.68	5.38	15.07	3.70	5.42	5.88
L2b	1.61	2.15	5.48	-	0.42	1.18
L2c1	-	-	1.37	-	1.25	-
L2d	-	-	-	1.85	-	-
L2e	-	1.08	-	-	-	-
L3b	1.61	3.23	2.74	3.70	1.67	2.35
L3b/d	-	-	4.11	-	-	1.18
L3d	-	-	4.11	-	1.25	-
L3e1	1.61	-	-	-	0.42	-
L3e2	4.84	-	5.48	-	0.83	2.35
L3e3	1.61	-	-	-	-	-
L3e5	11.29	-	-	-	0.42	-
L3f	-	4.30	8.22	3.70	2.08	-
L3h1b1a	1.61	-	1.37	-	-	-
L4b2	-	-	-	-	0.42	-
M1	3.23	5.38	-	1.85	7.08	4.71
N	1.61	1.08	-	0.93	0.42	-
R	-	-	-	0.93	-	-
R0a	-	-	-	0.93	1.67	-
R0a1a	-	-	-	8.33	-	-
T*	-	-	-	0.93	1.67	-
T1a	1.61	2.15	2.74	-	3.33	4.71
T2	-	1.08	-	0.93	0.42	-
T2b	-	-	2.74	-	2.92	-
T2c	-	-	-	-	0.83	-
U	-	1.08	-	0.93	0.42	-
U1	-	1.08	-	0.93	0.83	-
U3	-	1.08	-	-	1.25	10.59
U4	1.61	-	-	-	1.67	1.18
U5	-	-	-	-	0.42	-
U5a	1.61	3.23	-	-	1.67	-
U5b	1.61	1.08	-	2.78	0.42	-
U6a	-	4.30	-	7.41	6.67	-
U6a1a	-	1.08	-	-	-	12.94
U6a1a1	-	3.23	-	3.70	-	14.12
U6a1b	-	1.08	-	-	-	1.18
U6a5	-	-	-	-	0.83	-
U6c	-	-	1.37	-	0.83	-
U8b1	-	1.08	-	-	-	2.35
V	-	-	-	-	3.75	-
V7a	-	1.08	-	1.85	-	-
W	3.23	1.08	-	-	1.25	-
X	8.06	2.15	-	-	-	-
X2	-	1.08	1.37	-	1.25	-
mtDNA haplogroup diversity (h+-sd)	0.9175 +/- 0.0174	0.8630 +/- 0.0325	0.9376 +/- 0.0117	0.8823 +/- 0.0236	0.8853 +/- 0.0166	0.8891 +/- 0.0169

 

Good luck (and feed me back if you have some idea).