Genetic Profile: Maghrebi populations are Not "Satellite-Europeans" in Africa

Introduction: 

Many observers will note that at some point in time or another, they witnessed (if not perpetrators themselves) internet inquiries around the origins of Maghrebi populations, whom as indigenous Africans (let's face it; they are not endemic to any other region outside of Africa), are notable not only for having elements among them who are relative "outliers" in terms of epidermal pigmentation, but also for their own original distinctive language family, properly called Tamazight—otherwise inconsiderately known in English lingo as "Berber". These populations have wittingly or unwittingly been subjects of racism, in the sense that their "African-ness" have been questioned primarily because of appearance of those populations among them with "light skin", "depigmented skin" or "white skin'—the terms change depending on who's speaking. If for example, they were unanimously dark-skinned like the remainder of African populations, it is highly doubtful that "western" researchers would even consider tacitly passing them off as some sort of lost-Europeans (or extensions of Europeans) in Africa, notwithstanding any genetic similarity to the patterns seen today of Maghrebi populations.

An Investigation into the "Mysterious" Mesolithic Maghrebi populations

The present discussion takes off from the trail of a previous one posted here on the examination of the "Mechta-Afalou" and the European "Cro-Magnons" as elements of a monophyletic unit constituting a taxonomic "type". That previous discussion had put a spotlight on the futility of such a thesis, which varied quite widely in its specifics as one went from the subjective taxonomic regime of one researcher to the next. In that same discussion, the following was noted:  "the Mechtoid concept and its supposed relationship with Cro-Magnon, is nothing more than another Eurocentric way of trying to relate North Africa to Europe...essentially a *wishful* desire to see north Africa as more an extension of Europe than the actual continent [Africa] its attached to".

Indeed, there has been a long indirectly-stated desire to make the coastal northern piece of the African continent an extension of Europe, and territory which has traditionally been THE domain of European-descended peoples than of autochthonous Africans, which are generally treated as "newcomers" to the region. In other words, some sort of a "natural" apartheid was put in place in the late Upper Paleolithic until some time in the early Holocene, preventing autochthonous Africans from inhabiting the place, while the same region was supposedly hospitable to "European immigrants". This theme blatantly makes its reappearance in another study that has fairly recently enjoyed elevated circulation and has become a citation-favorite within online 'white' supremacist cult gatherings, undoubtedly attributable to the French-published study becoming more accessible as English-translated versions became more available on the net. The study in question comes from Kefi et al., under the heading: Mitochondrial Diversity of the Population of Taforalt (12,000 years b.p. - Morocco): A Genetic Study Approach to the Peopling of North Africa, 2005.

Review — "Linking the sub-Saharan and West Eurasian gene pools: maternal and paternal heritage of the Tuareg nomads from the African Sahel"

As it has become customary on this site, we will hereby examine the material specifics of work that recently came out, by Pereira et al., giving us an idea of what aspects of the journal withstands scrutiny and what doesn't. Below, is the extract, to give an indication of what we are working with:

Linking the sub-Saharan and West Eurasian gene pools: maternal and paternal heritage of the Tuareg nomads from the African Sahel

Pereira et. al.

European Journal of Human Genetics advance online publication 17 March 2010.

DISCUSSION

The Tuareg have a nomadic lifestyle and according to some demographic reports they show reduced fertility in comparison with their neighbours.47, 48, 49 The data observed here for mtDNA and Y-SNP diversities are concordant with those independent reports, especially for the Tuareg living within the bend of the Niger.

The overall West Eurasian mtDNA gene pool in the Tuareg population as a whole (H1, H3 and V) seems to favour a North African heritage.50 The only exception is the absence of the otherwise rare U5b that might have rather come to Africa through the Near East, and then drifted to higher frequencies only in some isolated populations such as in the Egyptian oasis Siwa.51 The absence of U6 can further be explained by genetic drift during the expansion of this haplogroup within North Africa.51 Note that U6 was observed at low frequencies in several population groups from the Chad Basin, such as in the Nilo-Saharan Kanuri and the Afro-Asiatic Masa.35

Relationships with the peoples of Eastern Sudan (the Beja) as pointed to by the study of classical genetic markers2 cannot yet be disregarded here as there is still no mtDNA of the Beja people available for study. However, according to historical reports, the origin of the Beja is more likely to be traceable to the Arabian Peninsula52 and the West Eurasian mtDNA lineages seen in the Tuareg have a rather Iberian affiliation in the post-LGM, and probably expanded to North Africa first.30, 31 The weak Eastern African influence in Tuareg is further supported by the M1 haplotypes belonging to the lineages characteristic of the later Mediterranean expansion (M1b and M1a2a) and the presence of very few matches for sub-Saharan L haplotypes with East Africa. The main post-LGM Eurasian and M1a2a lineages found in the Tuareg favour North African origin with migration to its southern location in the Sahel between ~9000 and ~3000 years ago. The upper time limit is defined by the age of the M1a2a, (estimated here from the coding region diversity observed in the three Tuareg, two North and two south Mediterranean individuals at 8000±2400), and by the upper 95% confidence interval for the Tuareg V lineages having polymorphism 16 234 (8800 years ago); the lower limit is defined by the age of the Tuareg V lineages having polymorphism 16 234 (3600 years ago).

The dates obtained from the genetic data coincide well with climatic changes in the Sahara, which resulted in repopulation during the first half of the Holocene when by ~10 000 YBP (the Holocene climatic optimum) humid conditions and greening were established. The climatic optimum lasted until ~6000 YBP, when the shift towards more permanent aridity occurred, culminating with the formation of the current Sahara desert. This desertification could have entrapped Tuareg populations coming from North Africa to the Sahel belt together with other pastoralists such as the Chadic speaking peoples41 coming from East Africa and Fulani nomads6 coming from West Africa. In fact, by performing complete mtDNA sequencing of the L3f3 lineage, specific for Chadic-speaking groups of the Chad Basin, Černý et al41 estimated a local demographic expansion during the Holocene period at about 8000±2500 YBP. No doubt all populations arriving to the Sahel were further enriched by various admixtures of many other sub-Saharan lineages, an effect even more pronounced in the Chadic groups who adopted a sedentary lifestyle soon after their arrival to the fertile Chad Basin than in the Tuareg who remain nomadic until present.

It is curious that, at least for the Tuareg maternal gene pool, there are no mtDNA lineages connected with the Neolithic expansion from the Near East despite being present in considerable frequencies in other North African populations. For example, the conservation of the high frequency and remarkable internal variability of T1 haplotypes within the distant and relatively isolated Egyptian oasis of el-Hayez led to an estimation of local expansion at around 5138±3633 YBP.37 There are no indications yet of the ages of local expansions in the more central and western regions of North Africa, which could contribute further insights for its absence in the Tuareg population as a whole.

Interestingly, for the Y chromosome, the dominant haplogroup in North Africa as well as the Tuareg is E1b1b1b. This haplogroup was associated with Neolithic diffusion in North Africa, with an age estimation of 2800–9800 YBP,45 but the lower resolution of the Y chromosome tree did not allow us to investigate this issue further. Nonetheless, disregarding whether they are in fact Neolithic, the ages for the mtDNA and Y chromosome lineages of North African origin observed in southern Tuareg are consistent with the same period, between 9000 and 3000 years ago. - Extract ends.

Let's start with a statement that Pereira et al. make in the abstract provided for the journal at hand:

"On the other hand, the Y chromosome SNPs data show that the paternal lineages can very probably be traced to the Near Eastern Neolithic demic expansion towards North Africa, a period that is otherwise concordant with the above-mentioned mtDNA expansion."

What is one to make of the Pereira et al.'s (2010) reference to a "Near Eastern" origin above? The answer is that the said authors meant by it just as it sounds like: "Near Eastern". Obviously, the line of question at hand is not so much as a demand for learning the definition of the Euro-centered political/geopolitical construct  of  the so-called "Near East", but so much as a call to question, the far-fetched claim about "Near Eastern" origins of Tamazight speakers of western Africa, including Tamasheq (Tuaregs) nomads, whom they compare to the Beja, another group whose origins they dubiously trace to the Arabian peninsula. As ridiculous as it may sound, from the looks of it, the authors are interpreting the African haplogroup E1b1b1b (a.k.a. E3b2-M81) as a "Neolithic" marker of "Near Eastern" rather than the conventional understanding of its African origin. The present author of this blog supposes that this is the reason they find it "interesting". If so, conversely, the present author finds it interesting, how a typical African marker turned into a "Near Eastern" one.

As a simple matter of fact, neither the quadro-entity of hg E1b1b1b, Tamazights, Tamasheq (Tuareg) nomads, and Beja nomads are found in the Arabian peninsula, at least not as autochthonous groups. All are rare to absent in the Arabian peninsula. Hg E1b1b1b is generally absent in the Arabian peninsula, save for the few sporadic spots which became host to historic era immigrants from coastal northwestern Africa, like say the "Mizrachim" Sephardic Jews of coastal northwestern Africa, elements of which can be found in Israel. That is the only context in which any hg E1b1b1b subclade makes its presence in the so-called "Near East". The basal clades of the marker are restricted to Africa, not to leave out the haplogroup's diversity. Likewise, neither Tamazight nor Beja language are spoken in the Arabian peninsula.

Getting back to that matter of why Pereira et al. find it interesting that the "dominant" male haplogroup of coastal northwest Africans and the Tamasheq ("Tuaregs") happens to be of the characteristic "Berber" marker of hg E1b1b1b, when such an understanding is otherwise conventional wisdom and therefore, to be the expected turn of results; let's think about it for a moment: Getting an understanding of why these authors feel that way may also explain why they would even bother to entertain an Arabian peninsula origin of the Beja, even though no such group is native to that area. The invocation of the Beja has a role to play here: Since the Beja (Bedawi) have been claimed to show close relationship with the Tamasheq/"Tuareg" in one study [specifically by Sforza] that used select autosomal markers, the authors brought them up perhaps to support their idea of the supposed arrival of Tamasheq/"Tuaregs" first from coastal north Africa, and then their settlement in the Niger Bend areas upon the desertification of the Sahara in the mid-Holocene era.

They try to buttress this viewpoint, from what the present author can discern, by turning to what they call "Eurasian component" of Western African Tamasheq/"Tuaregs", again NOT the Libyan or northwestern Saharan Tamasheq/"Tuaregs", but the ones in the noted "sub-Saharan" African countries. The markers that they are leaning on are hgs H1, H3 and V.

Pereira et al. seem to be of the mindset that the Y-markers came to coastal north Africa via the "Near Eastern" Neolithic diffusion, and then in coastal northwest Africa, mtDNA from the Iberian peninsula were picked up, with some contribution from a "Near Eastern" Neolithic mtDNA gene pool [albeit not significant in the western African Tamasheq ("Tuareg"), according to the said authors' own observations], which they seem to identify with the expansions of M1, hence calling it a late "Mediterranean expansion".

Few other things to take into account:

Hg M1 markers...

The M1 markers implicated in the extract are actually of the Eastern African types, and not the typical coastal northwestern African examples. While the authors here do seem to entertain an ultimate eastern African origin for the M1 clades that they mentioned on the one hand, from the language in the extract provided, it appears that they were describing a coastal north African flow of the marker on the other hand, i.e. presumably before its heading for the Sahel in western Africa. In other words, notwithstanding the acknowledgment that their mentioned M1 sub-clades originate in eastern Africa, Pereira et al. are treating the M1 sub-clades here as "northern markers".

*More insight into hg M1 expansion across coastal northern Africa, eastern Africa and the so-called "Near East" had recently been posted here: Following Trails of the Cro-Magnon - II

The Beja...

While some genetic infiltration has taken place into the autochthonous gene pool of the Beja, the bulk of their Y-DNA lineage is of M78, BUT these are not the examples found in the Arabian peninsula; rather, these are the examples mainly found in the African Horn. The bulk of Beja M78 markers carry the V32 binary marker [see for example, Hassan et al. 2008].

The Beja language is considered to be a standalone sub-phylum of the Cushitic branch, and hence, not Semitic. The Arabian peninsula is dominated by Semitic languages.

The west African Tamasheq or "Tuaregs"...

The study that the present author posted on this site earlier about the largely west African component of west African Tuareg mtDNA related the following, to recall:

West African Tamasheq/"Tuaregs":

The mitochondrial data of the Northwest African populations (Berber from Morocco and Algeria, Moroccans, West-Saharans, Mauritanians, Tuareg) show a mosaic composition of mtDNA types, with a pronounced gradient of sub-Saharan lineages from north to south: at the one extreme, the Berbers from Morocco have a predominantly European (Iberian) affinity, while at the other extreme, the Tuareg are closely related to sub-Saharan West Africans as represented by several Senegalese groups in this study, whereas the West-Saharans and Mauritanians are somewhat intermediate. It is remarkable that the Tuareg bear little mitochondrial resemblance to the Berber populations, although they speak a Berber language. - Rando et al. 1998 [mtDNA analysis of Northwest African populations reveals genetic exchanges with European, Near Eastern and sub-Saharan populations]

One should not think for a moment that Rando et al. (1998) were alone in their findings about western African "Tuaregs"; the following from Claudio Ottoni et al. (2009) essentially tells the same story:

Of note is that the other Tuareg sample described in the literature (Watson et al., 1996) (Western Tuaregs) did not show a close genetic relationship with the Libyan Tuaregs, implying a genetic heterogeneity of the Tuaregs. This difference appears to be primarily caused by the low frequency (8%) of the European component in the Western Tuaregs, characteristic of northern African populations. After the removal of the H and V haplotypes, the Libyan Tuaregs showed a strong affiliation with the Eastern populations, while the Western Tuaregs associated more with the Central and Western African populations. - Claudio Ottoni et al. 2009, First Genetic Insight into Libyan Tuaregs: A Maternal Perspective.

*While Claudio et al. have their own shortcomings—a  matter that may be saved for another discussion, what is appealing about their findings is the fairly comprehensive range of their Tamasheq ("Tuareg") population samples, spanning western Africa to coastal central-north Africa, and thereby giving a noticeable mtDNA structuring that takes shape in the form west-to-east or vice versa gradients and south to north or vice versa gradients. In the west-to-east genetic structuring, western African "Tuaregs" more readily related to other western Africans and central Africans, while by removing the hgs H and V from the Libyan "Tuareg" sample, the Libyan-situated "Tuaregs" more readily related to eastern African populations. As noted here (click to the link) before, this pattern is suggestive of expansions of ancestors of contemporary Imazighen populations as a migrating group whose demography was likely biased towards male constituents; females of their destination points or from the vicinities thereof were then later integrated into the Imazighen nomads. So, we see distinctive patterns between Libyan "Tuaregs" and the western African "Tuaregs". The western African Tuaregs, just as Rando et al. found out, cluster with other western Africans. What does this then tell us? It tells us that the findings of authors in the intro post is the one that is out of character, if we are compelled to judge the results on such grounds.

Even Pereira et al.'s own observations hint on the very real possibility that they are only seeing a snapshot of a bigger picture...

The weak Eastern African influence in Tuareg is further supported by the M1 haplotypes belonging to the lineages characteristic of the later **Mediterranean** expansion (M1b and M1a2a) and the presence of very few matches for **sub-Saharan** L haplotypes with East Africa. - Pereira et. al.

What else do these "sub-Saharan" L types match with then? Why, very likely the central and western African examples!

Hint on what these L types are, may be found in the authors' emphasis on Chadic speakers carrying L types.

They theorize that the Tuaregs could have started in the North, and then descended down, based on H and V markers, which they date to around 8000 YBP or so, but guess what? That specific L haplogroup (L3f3) that they do mention in the extract provided, also dates to about the same time frame, or falls within the same window of time frame!

What does all this mean at this point? Their theory has little basis to it. This may very well be hinted on by Pereira et al.'s own puzzlement in the following:

It is curious that, at least for the Tuareg maternal gene pool, there are no mtDNA lineages connected with the Neolithic expansion from the Near East despite being present in considerable frequencies in **other** North African populations. - Pereira et. al.

It puts them in a difficult position, because their mental attitude seems to be one of a demic diffusion from the "Near East" in the Neolithic times along the coastal areas of North Africa, whereupon in the far west Maghreb, they'd pick up European mtDNA marker types. This then will have been followed up with the mid-Holocene aridity of the Sahara, prompting the ancestors of Tuaregs to seek refuge in the Sahel, where they would mix with "sub-Saharan" Africans. This absence of "Neolithic" mtDNA markers in Sahel/"sub-Saharan" west African Tuaregs upsets that theory, in addition to the other little details just noted. 

Additionally, they mistake an African-specific Y gene pool with "Near Eastern" Neolithic demic diffusion. Northwest African M78 markers cannot be from the "Near East", because northwest African M78 gene pool is actually more diverse than those in the Levant, where the "Neolithic farming" revolution is supposed to have started, i.e. if we are to go by data that Cruciani et al. (2007) give us. Not to leave it out here, but even E-M81 (E1b1b1b) is rare to absent in the "Near East". In this idea, Pereira et al. may well have drawn inspiration from Arredi et al. (2004): 

Arredi et al.(2004) ran off with the theory that hg E1b1b1 markers first arrived in northern Africa in tandem with "Afro-Asiatic" speakers moving in from the "Near East", having citing outdated linguistic theories [like that of Diamond and Bellwood (2003)] of "Near Eastern" origins of "Afro-Asiatic" as their source of inspiration; this was the thesis of their study. They seemed to be tacitly implying that the characteristic E-M81 emerged along the path of this migration, with relative diversity of the clade decreasing as one moved to the far western corner of the continent. Of course, Arredi et al. (2004) did not account for the microsatellite distinctions in hg J clades, the other marker that is generally implicated in "Neolithic farming" subsistence for the so-called "Near East" aside from hg E1b1b1markers, that Semino et al. pointed out the same year. Much of northern African hg J clades appear to be more of post-Neolithic historic extraction, esp. during Arabic expansions, than Neolithic extraction, even though the coalescent age of the J1-M267 sub-clades when considered together point to initial dispersions within the vicinity of Neolithic time frames;  this latter observation is true for the more recent "Near Eastern" chromosomes as well. The older clades were better preserved in European and Ethiopian samples, according to Semino et al., and hence, the most likely markers of Neolithic era expansions. Moreover, Arredi et al.'s hypothesis is contemptuous of the facts mentioned above, to reiterate, about the restriction of hg E1b1b1b markers to Africa, the absence of Tamazight as an autochthonous "Near Eastern" language and of any languages therein in the "Near East", that serves as the root stratum of Tamazight. Lastly, as pointed out here time and again, northern Africa did not adopt large scale farming subsistence at the same time or within the logical or expected time frame that it should have, in the event that one should entertain the idea of Neolithic demic diffusion in tandem with the spread of agricultural subsistence from the so-called "Near East".

What Pereira et al. may be seeing, if anything is to be made of it, is a portion of the "sub-Saharan" west African "Tuareg" gene pool that was attained from interactions between those Niger River bend "Tuaregs" and the more northern west Saharan groups. It is not inconceivable, since paternal and cultural relationship exists between these groups. The patterns seen in west African "Tuaregs" is obviously reflective of the Imazighen bio-history as nomads, and so, in order to get a fairly comprehensive picture of the uniparental DNA structuring in such groups, it is necessary to deal with large sample sizes that are not from just one or two spots, but spanning the general territories that are known for Tamasheq ("Tuaregs") communities. Rando et al. and Claudio et al. had done this to some degree or another, and hence, their relatively more consistent findings in DNA structuring. Pereira et al. may be suffering from what Cherni et al. described as "patchy" DNA sampling work. However, western African "Tuaregs" in the main, cluster with western Africans where mtDNA is concerned. As we've seen, two studies already attest to this, and as shown above, there are also hints in Pereira et al.'s own findings that this is the case.

Keep an eye on possible updates in future!
 ___________________________________________________________________
*References

—Pereira et al. (2010), Linking the sub-Saharan and West Eurasian gene pools: maternal and paternal heritage of the Tuareg nomads from the African Sahel.

—Rando et al. (1998), mtDNA analysis of Northwest African populations reveals genetic exchanges with European, Near Eastern and sub-Saharan populations.

—Claudio et al. (2009), First Genetic Insight into Libyan Tuaregs: A Maternal Perspective.

—Hassan et al. (2008), Y-Chromosome Variation Among Sudanese: Restricted Gene Flow, Concordance With Language, Geography, and History.

—Semino et al. (2004), Origin, diffusion, and differentiation of Y-chromosome haplogroups E and J: inferences on the neolithization of Europe and later migratory events in the Mediterranean area.

— Arredi et al. (2004), A Predominantly Neolithic Origin for Y-Chromosomal DNA Variation in North Africa. 

—Personal notes; 2009, 2010.

The So-called Tehenu Palette

The palette itself appears to have an alternative name given to it by contemporary researchers, perhaps underlying the uncertainty surrounding the basis on which the palette has been associated with the Tehenu and/or else "Libyans", along with other speculative explanation based on such. For instance, on one website, the following was offered as an explanation:

The Tehenu palette (or Towns palette) in Cairo (C.G. 14238) is named after a sign on its verso: this shows three registers with domestic animals files and a fourth lower one with plants (trees) and the hieroglyph of the throwing stick on an oval (which means 'region', 'place', 'island'), thus a toponym of Libya or Western Delta (THnw, Tjehenw). The recto of the palette is of great importance, showing the feet of some persons and, below the register line, two rows of four and three groups respectively; each group is constituted by an animal grasping the Mer-hoe on the crenellated wall of a town; the name of each town is written within the wall [see my Corpus of Late Predynastic Decorated Palettes]. - Courtesy xoomer.virgilio.it

...that alternative name, as provided above, is "Towns" palette. Given the air of uncertainty around preexisting rationales that associate the palette scenery with "Libyan" or "Tehenu booty", the relative neutral, if not ambiguous, moniker of "Towns palette" is not unreasonable. Coincidentally, the extract reaffirms what the present author had bumped into while researching the question of the actual underlying connections made between the "Tehenu Palette" and socio-ethnic entity known to us by primary Kemetic texts, i.e. prior to the above coming to immediate attention. This has to do with the said hieroglyphic sign of the "throwing stick". Accordingly, the present author put forth the following in a discussion elsewhere:

Managed to come across reference to Alexandre Moret's and Margaret Murray's interpretation of that "throw strick"/stick sign to the right of the plants-i.e. on the side of the "Tjehenu Palette" bearing the animals and plants--as "Thn", and therefore read it as "Tehenu" or "Libyans". But as far as I know, the "stick" is generally an ideogram for something "foreign" or a "foreigner", and not relegated to any one specific ethnicity or locality. On the other hand, Jean Capart put forth the case that the said plants are "Libyan", reportedly from having made comparisons to "a bas-relief depicting Libyan spoils in Sahure's temple".

Indeed, the "throwing stick" alone does not implicate nor is relegated to just one specific foreign socio-ethnic entity or foreign locality, unless, accompanied by other particulars or specificities that point to a specific foreign ethnic entity or locality. There is nothing there about the ovular-looking heiroglyphic sign [in the lowermost register on the "verso" side of the palette] accompanying the "throwing stick" in question either, that supposedly speaks to singular and specific foreign ethnicity or locality, which in the present case, is identified as either "Tehenu" or "Libyan". Therefore, one has to look for a more compelling explanation than the one suggested by either Moret or Murray, and pin it down to a specific ethnic entity and/or state. For instance, one can start by narrowing down the possible territorial confines in which the palette scenery is implicated. As an example, perhaps if there is any indication that the side of the so-called "Tehenu Palette" bearing registers of animals and plants describes a locality, then perhaps the donkey or ass figures therein could suggest that it is on the African continent, especially if one were to put faith in the age estimations given to the palette, the specifics of which we will shortly visit, which would make the domestication and widespread use of donkeys outside of Africa a relatively rare sight at the time in question. Jean Capart, as mentioned above, appears to have taken this latter route of correlating certain "spoils" [which presumably, happened to be made largely up of fauna] implicated in the endeavors of exotic personalities [vis-a-vis the Kemetian], as it relates to their dealings with Kemetic personalities in artistic scenery, with geography. To see how Capart may have made the "Libyan" connection to the so-called Tehenu palette, one might want to examine what is displayed in the Sahure temple wall renditions that he was presumably referring to; to this end, the following images come to mind:

 Click on the image for greater resolution

Indeed, here we see accompanying fauna and flora that are not that different from those seen on the so-called Tehenu palette. Visible are sheep-like figures, cattle and asses/donkeys, which are also featured on the so-called Tehenu palette. The only fauna shown here, not visible on the "Tehenu" palette fragment, is the goat or goat-like figures. Nor are flora featured in the particular rendering at hand; whereas, at the very bottom register of the "verso" side of the "Tehenu" palette, flora are featured. The wall relief above features human personalities just above and below the registers featuring fauna; The "verso" side of the "Tehenu"/"Libyan" palette on the other hand, displays no human figures, to the extent of what is visible there. There are human figures featured on the "recto" side of the palette though, but even these are not sufficiently visible, so as to make a logical statement about possible patterns of attire on not just a single figure, but several involved.

 Click on the image for greater resolution

Another standout, is the personalities accompanying the "spoils" [if these are indeed the spoils Capart was referring to] or else fauna; while they too are partially uncovered, where attire is concerned, the patterns invoked in their attire is markedly distinct from those featured in another palette wherein "Libyans" or else "Tehenu" have been implicated by some observers—the Narmer palette. What significance does the Narmer palette hold here? For one, it is supposed to be the temporally closer item to that of the "Tehenu" palette than the Sahure temple wall relief in question. Thus, if one is to have any faith on claims that "Tehenu" are featured on either the "Tehenu" or Narmer palettes, then it is only fitting to examine any connections between the attires featured on the human figures. For this purpose, the Narmer palette renditions are provided below:

Click on the image for greater resolution

The bearded fellows under captivity, who are supposed to be "Tehenu" or else "Libyans" according to some observers, do not feature the same attire as the figures in the Sahure temple wall relief. If the figures on the Sahure wall mural are considered to be "Tehenu", then taking that into account thereof and making a comparison to the "Libyan" or "Tehenu" figures on the Narmer palette does not immediately make apparent, the prospect of the latter "Libyan" figures of the Narmer palette being "Tehenu" as well, solely based on attire. The strapped-gear around the trunk of the human figures of the wall mural are visibly absent on the bearded-figures featured on the Narmer palette. It can be pointed out that the figures on the Sahure temple wall mural generally comprise of juvenile and feminine personalities, as opposed to adult male personalities. A close inspection of the 'verso' side of the Narmer palette, as shown above, also features what appears to be a juvenile personality to the left hand side of the authoritative figure [generally deemed to be Narmer, in this case], and here too, one cannot help but notice a strapped-gear around the trunk, with the straps crossing one another at the chest level—not unlike the manner in which the "strapped-gears" of the Sahure wall mural "Tehenu" figures feature. This figure though, sports some yet-to-make-out item on the chest area, where the straps are supposed to cross one another; this feature is absent in the Sahure wall mural counterparts. Furthermore, this same figure appears to be wearing a loincloth that is not too deviant from that of the aforementioned head figure (Narmer). This stands in contrast to the juvenile "Tehenu" figures of the Sahure wall mural, whom aside from the said "strapped-gear", wore nothing underneath, i.e. below the waist. On the other hand, the "Tehenu" figures in general, feature some sort of neck gear or ornament, which is visibly absent in the Narmer "juvenile" figure in question. Not to be left out in the midst of these observations, is that this Narmer palette "juvenile" personality does not appear to be an accomplice of the supposed "Libyan" figures, as he doesn't appear to be under any form of suppression as the said "Libyan" figures, nor do they share attire forms.

Visible on at least one of the "Libyan" figures, is a semi-exposing attire below the waist, only featuring a "penis sheath". The adult "Tehenu" figures of the Sahure wall mural also appear to be wearing similar line of clothing, although these are generally accompanied by some rope-like or semi-looped tool, weapon or device on the sides of the figures, which is visibly absent on the Narmer palette figures in question. At the top registers of the Sahure wall mural, there are some characters that might remotely be deemed to be those of male figures, but because of the tearing and wearing, and especially if one is faced with dealing with repros, cannot conclusively be reported as such. However, repro of a "Tehenu" figure, floating around the web, displays an adult male figure featuring attire not too different from the Sahure wall mural personalities:

Click on the image for greater resolution

This male figure above on the other hand, has gear that is visibly distinct from the personalities mentioned on the Narmer palette. Even the head-hair arrangement is distinct from those of the "Libyan" personalities of the Narmer palette. So, if these items are to serve as the parameters by which a connection between the Sahure wall mural figures and those of the "Tehenu" and the Narmer palettes are made, then such connection is not made apparent here. This in of itself doesn't disprove, or prove, that the figures on either the "Tehenu" or "Narmer" palette are of folks associated with territory now identified with Libya, but it does put a question mark on whether the figures of the palettes in question are in fact those of "Tehenu", i.e. if the Sahure wall-murals are determined to be those of "Tehenu". Nor does it get any clearer, when the recto side of the Narmer palette is considered:

Click on the image for greater resolution

Can it therefore be reckoned, that the attire of the Tehenu may have changed through the time frames in which the discussed art renditions belong? It is not impossible, but evidence for such is wanting.

On a passing note, the long-necked feline creatures on the recto side of Narmer palette are hard to ignore. Some observers have dubbed them as "serpopards". Some have interpret  them as supposed "imports" from the so-called "Near East", predicated on the flimsy standpoint that similar creature is featured in Uruk artifact dated to more or less contemporary time frame as the Narmer palette, like the Uruk seal below, reportedly dated to c. 3000 BC:

Click on the image for greater resolution

Notwithstanding such claims, the Nile Valley example is actually dated earlier than the "Near Eastern" examples. Anytime there are some parallel cultural/artistic expressions in the Nile Valley and the so-called Near East, these same characters tend to give priority to the so-called "Near East" as the most likely source, as if to tacitly say that the Nile Valley can never be conceived as the source (it is virtually impossible) and therefore, must always be the recipient of cultural diffusion. Note that the Uruk  serpopards have much longer tails than the Narmer palette rendition, with their tails almost as long as their necks.The mythic creatures themselves may well serve as "guardians" against enemies or adversarial elements, just as that served by other mythic creatures like the Sphinx. Now that the issue of dates has been brought up, we now visit the issue of the dating attributed to the palettes discussed here. On the basis of serekhs and iconography associated with successive designated rulers spanning the predynastic time frame, attempts had been made to place the age of the so-called Tehenu palette amongst several other palettes. For instance, G. Dreyer  [Umm el-Qaab I, Mainz 1998] sought to do this very thing, i.e. reading through a series of signs/serekhs on a single palette, check to see which pharaoh figures' signs, names or serekhs have been implicated and whose has been left out thereof, and extrapolate from there, the likely temporal place of the palette in question in the midst of several other palettes. Going by a website earlier cited above, we are informed how Dreyer and others went about doing this, in reference to the themes of the Coptos Colossi statues:

The colossal limestone statues of Min were found by Petrie in the temple of Koptos in 1894; they had been fashioned with hammering technique (no chiselling) and represented the god standing with erected phallus; only the torso and part of the legs was preserved and the head of one of the statues in Oxford, although almost entirely effaced; some signs in relief were noted on the statues showing animals, plants, shells and standards.

In an article published in 1988 [JARCE 25, 35-60], B. Williams suggested the presence of a fragmentary trace of the name of Narmer on the Cairo statue; this gave an important clue about the long disputed question of the date of the statues (which in the past had ranged from Predynastic to 1st Intermediate Period according to the opinions of different Egyptologists). [For reconstructions of the colossi and temple see this page in the Petrie Museum website: Digital Egypt].

In 1995 Dreyer (loc. cit. above) proposed that the graffiti on the statues were the names of older rulers, and Narmer had been the last one to make his name be carved on those statues; therefore the colossi probably dated well before his reign, down to Naqada IIIa, and the signs carved onto them would be perhaps something similar to a king list. I must notice that the Nar-fish and the Mer-chisel are very fragmentary -only the left end preserved- and, as suggested by Kemp, the upper sign is rather the tail of a bird than that of the Nar cat-fish, thus suggesting a falcon on a perch or on a standard [cf. B.J. Kemp, CAJ 10.2, 2000, 211-242, fig. 10; H. Goedicke, MDAIK 58, 2002, 253].

Basing on the reciprocal placement and superimposition of the signs on the colossi, Dreyer seems to have found a possible sequence of the stages in which the graffiti were incised (cf. table): Animal-head standard, Shell, Elephant, Bull, Stork, Canid, Min-standard, Plant, Lion and Narmer. - Courtesy xoomer.virgilio.it

A similar approach was taken by Dreyer in narrowing down the chronological order of the so-called "Tehenu" palette amongst several other predynastic era palettes. In the case of the "Tehenu" palette, the signs implicated are those that belong to the following figures: Falcon, possibly but not certain—Falcon II, Lion, Double Falcon, and Scorpion II. Other figures or names had been left out from what was available to read on the palette, but from extrapolation, it may be safe to say that the palette belongs to a chronological order that proceeds those of unlisted figures that predate the ones named or implicated, while preceding those unlisted counterparts that are considered to post-date King Scorpion II. Reading on, from the above mentioned website, and in a passing, the author gives us his personal opinion about the chronological placement of the Tehenu palette...

Dreyer has postulated that, given the mentioned late-Dynasty 0 manufacture of the Tehenu palette (which in my opinion is in fact later than the Battlefield and Bull palettes but earlier than the Plover and Narmer palettes, it could never neither celebrate nor narrate the foundation of the towns by those kings : the town of the Heron (Djebawty), probably Buto, which is currently being excavated by T. Von der Way, was founded much earlier than Naqada IIIA. Therefore the action performed by the royal entities on the palette could eventually be the foundation of fortresses in the respective centers (cf. the example from Elephantine), or more probably it was the (symbolic ?) destruction of the centers after their defeat by the Southern Kings; this progressive military expansionism and submission of the Delta by the Thinite sovereigns, which is echoed in the scenes of battle and of their aftermath represented on Naqada III palettes and ivories, was probably a relatively common scenario until the country unification (cf. M. Campagno 2002). As quoted above, and always with the due cautions, it can be supposed that a parallel warfare-pattern should have been followed by the Dynasty 00-0 kings in respect of the Nubian antagonists; perhaps also in the Delta the Maadi-Buto decline hadn't happened (Naqada IIC-D) without some conflict; the EB I Canaan colonization is instead a different matter (although some scholars hypothesized, in the past, massive military interventions of the Egyptians there): the difference between Egypt and Canaan at that time was too large to favour the assumption of any possible competition between them; Egyptians must have found no resistance in their infiltration into those territories, contributing to their evolution towards the EB II Urbanization [cf. the interesting and still valid synthesis 'The relations between Early Bronze I age Canaanites and Upper Egyptians' by Branislav Andelkovic, 1995].- Courtesy xoomer.virgilio.it

Indeed, there seems to be little agreement on, nor are any conclusive material backing available out there that sufficiently lean towards one interpretation or the other, what the signs for localities or towns specifically point out. Two common themes that the various interpretations seem to oscillate to and fro, is one either of "foundation" commemorative symbolism [acquiring territory] or "victory" commemorative symbolism [display of spoils of victory in conflict or war]. The issue of whether the  "Tehenu"/"Libyan" palette is justifiably named so has been explored over the course of this post, particularly as it pertains to the "Tehenu" socio-ethnic entity; but what about a relatively more generalized prospect of the palette featuring groups from "Libyan" territory? Well, again, some level of evidential specificity has to be first established that narrows the possible territorial candidacy down to a fairly restricted geographical confine, which in this case would be the area where modern Libya now lies, as opposed to evidence that might narrow the likely geographical location down to a sub-continental level or a wide regional level. Some of that has been looked at in terms of certain fauna, but that sort of evidence can only go so far in pinning down a specific restricted locality, since such fauna were likely found in multiple locals in the same general time frame. And even if one were to narrow the likely territory down to "Libya", that stills tells us little about the specific ethnicity or nationality involved, which in this case, some observers are inclined to implicate the "Tehenu". We've covered how J. Capart sort to make such connection via what is referenced as "spoils", between that seen in the Sahure wall mural—claimed to be associated with the Tehenu—and that seen in the "Libyan"/"Tehenu" palette; he does make some interesting comparisons around the fauna, but even then, as discussed above, questions continue to linger around human figures, what they feature in terms of gear and attire, and accompanying hieroglyphs of the scenery. The damages on the "Tehenu" palette for instance, make it difficult to ascertain any trends about the human figures on the recto side the palette; the Narmer palette on the other hand, provides a better opportunity to make a comparative analysis between the human figures and themes on that palette and say, those on the Sahure wall mural. The usefulness of the Narmer palette in our case study lies in its implication in featuring themes around "Libyan" or possibly "Tehenu" foe. Pressing issues around this implication have also been briefly covered here.

As further information comes to mind, this post will be correspondingly be modified.
_____________________________________________________________
*References:

— Sources as already cited.

— Retrieval of personal notes from elsewhere.

Review: Population Relationships in the Mediterranean Revealed by Autosomal Genetic Data (Alu and Alu/STR Compound Systems)

Herein, aspects of a recent publication by E. González-Pérez et al. (2009) under the heading described above, will be revisited and examined respectively.

The abstract goes as follows:

González-Pérez et al.

Am J Phys Anthropol. 2009 Nov 16.

The variation of 18 Alu polymorphisms and 3 linked STRs was determined in 1,831 individuals from 15 Mediterranean populations to analyze the relationships between human groups in this geographical region and provide a complementary perspective to information from studies based on uniparental markers. Patterns of population diversity revealed by the two kinds of markers examined were different from one another, likely in relation to their different mutation rates. Therefore, while the Alu biallelic variation underlies general heterogeneity throughout the whole Mediterranean region, the combined use of Alu and STR points to a considerable genetic differentiation between the two Mediterranean shores, presumably strengthened by a considerable sub-Saharan African genetic contribution in North Africa (around 13% calculated from Alu markers). Gene flow analysis confirms the permeability of the Sahara to human passage along with the existence of trans-Mediterranean interchanges. Two specific Alu/STR combinations-CD4 110(-) and DM 107(-)-detected in all North African samples, the Iberian Peninsula, Greece, Turkey, and some Mediterranean islands suggest an ancient genetic background of current Mediterranean peoples. - abstract ends

A run down of the Alu markers and 3 Alu-linked STRs, is as follows:

18 autosomal Alu markers: CD4, TPA25, APO, ACE, Yb8NBC120, Yb8NBC125, B65, D1, FXIIIB, A25, PV92, HS2.43, Sb19.3, Sb19.12, HS4.32, HS4.69, DM, Ya5NBC221

Tandem Repeats linked to sites identified with specific Alu insertions or deletions designated by three designators: CD4+, DM+ and FXIIIB-. These reportedly represent the "ancestral" states of the sites in question, and the nature of said site-states are indicated by either the "+" or "-" symbols respectively.

Y-chromosome Alu insertion: The YAP+ Y-chromosome Alu insertion, fairly common in African populations, particularly in the form of Hg E, serves as an addition to the collection of Alu markers cited above.

Populations sampled:

Click on the image to get a better resolution.

The study opens with the following lines:

As far as the origin of human populations in the Mediterranean is concerned, it is commonly accepted that their roots can be traced back to the Upper Paleolithic with the expansion of human groups from the Near East or Central Asia, or some millennia later with the westward and northward spread of Neolithic populations from the Fertile Crescent. Although there is little doubt regarding the human entrance route to the Mediterranean, controversy appears when different studies try to determine to what extent their current genetic background preserves traces of Paleolithic people and in which degree the almost continuous cultural and political contacts have influenced present genetic affinities.

Indeed, the authors are correct in their assessment about controversy in terms of how different studies interpret their observations, with regards to the entrance or exit of certain lineages, particularly in relation to the time of event and how extensive. Their own opening assessment attests to this, recalling the bit about origins of "Mediterranean" populations being traced back to the so-called "Near East" or Central Asia. This implies that northern Africa was a barren region for a long period of time, where no autochthonous African population ventured, even as humanity spent the bulk of its socio-biological evolution exclusively on the continent until ca "50-60 ky ago era" when a subset of anatomically modern humans successfully left the African continent for refuge elsewhere. This begs the question: human beings in Africa did not see fit to populate the northern areas of the continent, yet non-Africans were supposedly the first to see fit to do so? As a matter of fact, Maghrebi paleontological record stretches back to as far as the Middle Paleolithic era, preceding anatomically modern human occupation outside of Africa.

The authors appear to have been influenced in their assessment by the likes of Olivieri et al. (2006), whose work has been a subject of discussion (clickable link) on this site, as it relates to the Upper Paleolithic demic diffusion episodes in the northern sections of the African continent, and by the likes of Arredi et al. (2004), as it relates to theories surrounding Neolithic demic diffusions in that same region. Apparently, the authors are working with outdated concepts in their assessment, as earlier theories about Upper Paleolithic northern African complexes [see for example, the so-called "Ibero-Maurusian"] being manned by people from outside of the continent, based on erroneous assumptions built around archaeological finds on lithic artifacts, have now been rectified and updated with research that link origins of certain Upper Paleolithic lithics-oriented innovations to northern Africa, which were subsequently diffused into neighboring extra-African territories. Other erroneous assumptions about the earliest Upper Paleolithic northern African anatomically modern populations coming from outside had been based on shabby and flimsy reliance on outdated bio-anthropological concepts built around cranio-morphometric examinations. A notable example that immediately comes to mind, is the idea of Mechtoid populations [see: Mechta and Afalou: Do they and the so-called "Mechtoids" constitute a type with the "Cro-Magnon"?] , who were almost considered to be synonymous with the Cro-Magnon of Europe. Outposts of lingering cult-like Eurocentric elements continue to rely on ideological concepts of the Cro-Magnon as some sort of embodiment of "Caucasoids" or "Caucasians", and even that, has been discredited by more recent and refined analysis of cranio-morphometric data [See: Brace et al. (2005) and Chris Stringer (click), for example]. Even earlier bio-anthropologists tacitly took note of differentiations between the African Mechtoid variants and those of the European Cro-Magnon, even as superficially-invoked links were being insinuated. The bottom line is that the authors' presumptuous assessment, that the theory of northern Africa being first populated by people from the so-called Near East is a "commonly accepted" understanding, has little basis to it, as no prevailing evidence backs up such a notion. It implies that this is an understanding that has harmonized the various scientific multidisciplinary applications at our disposal, when no evidence has been brought forth to suggest such status quo. It is certainly not the message harmoniously relayed by either genetics or paleontology, nor by Upper Paleolithic complexes, as just mentioned a few comments ago. Then by what, aside from wishful thinking? Notably, the authors' own data does not lend support to such thinking. On the other hand, the flaws of Olivieri et al.'s (2006) and Ana Gonzalez et al.'s (2007) Upper Paleolithic demic diffusion hypothesis have been touched upon on this site before. U6 is undoubtedly Upper Paleolithic by most accounts, but it makes up very little of the contemporary northern African gene pool, while major M1 expansions are mostly linked with spread of proto-Afrasan or Afrasan-affiliated speaking groups some time in the late Paleolithic and early Holocene Neolithic time frames. Again, Olivieri et al. (2006) are emphasized here, because the authors of the present study appear to be relying on them, with regards to so-called Near Eastern sourcing of Upper Paleolithic northern African populations; see for example:

Recent mitochondrial DNA data (Olivieri et al., 2006) suggest a common Levantine source for the Upper Paleolithic cultures that occupied the European (Aurignacian) and North African (Dabban) shores of the Mediterranean. A more recent origin for these populations associated with the demic diffusion of Middle Eastern groups in the Neolithic has been suggested by studies of Y-chromosome (Arredi et al., 2004) and autosomal data (Myles et al., 2005; Tomas et al., 2008).

The authors of the present study themselves reference research that contradicts the idea of either the Upper Paleolithic or Neolithic sourcing from the so-called Near East; see for example:

A detailed survey of the E-M78 Y-chromosome haplogroup (Cruciani et al., 2007) indicates the Northeast African origin of this variant and its involvement in trans-Mediterranean migrations from North Africa to Europe during the last 13,000 YBP.

The predominant paternal markers of coastal northwestern-central African markers are comprised of E-M78 and E-M35 markers. This being the case for the northern African populations that the authors sampled here, the fact serves as a major contradiction to the so-called Near Eastern sourcing of northern African populations, who have supposedly persisted into contemporary times, if we are to go by conclusions drawn by the authors of the present study. Yet, we are suppose to buy into some presumption of the Upper Paleolithic "Near Eastern" sourcing of northern African populations as some sort of a "commonly accepted" understanding or truth.

It should be reiterated, as explained before on this site, that contemporary Imazighen-groups , who predominate much of northern Africa today, don't have TMRCAs—deemed to be "characteristic" of Tamazight or "Berber" speaking populations—that date to the Upper Paleolithic. Yes, these lineages derive from lineages of Paleolithic provenance, but they themselves, don't—at least not according to patrilineal lineage. The E-M81 mutation of the E1b1b lineage—which is predominantly found in Imazighen populations—has at most, been implicated in expansions that only go back as far as 8 ky or so ago. This falls short of the ages associated with Upper Paleolithic/Epi-Paleolithic or earliest Holocene cranial specimens uncovered in coastal northwestern Africa. Furthermore, none of the cranial specimens tied to contemporary northern African populations remotely tie in with the Cro-Magnon specimens of Europe, as Brace et al. (2005) had found out; whereas we are pressed to believe in ties between the so-called Mechtoid variants and the European-based Cro-Magnon.

Furthermore, the authors note:

Similarly, specific Mediterranean haplogroups or clades (U6 and M1b in the mtDNA; EM78 and EM81 in the Y-chromosome) have also been described for these populations and dated in Paleolithic times.

Nothing in the above supports "Near Eastern" sourcing of northern African populations examined here. None of the above markers are known for being quintessential indications of "Near Eastern" ancestry, as opposed to African ancestry. Not even U6, whose "non-African" ancestor remains ever so elusive, is a marker of the so-called "Near East"; it is quite rare in that region and its presence there can only be spoken of, in terms of back-migration from northern Africa, even if it is assumed that a proto-U6 ancestor was "Near Eastern" in origin. None of the markers above are even confined to the "Mediterranean" regions, so as to justify the use of the moniker of "Mediterranean haplogroups or clades".

One issue that stands out like a sore thumb, is the comprehensiveness of the authors' so-called "sub-Saharan" collection; see:

In search of new insights into these questions, this study analyzes a relevant set of Mediterranean populations including eight European samples (from Spain, France, Greece, and Turkey), seven from North Africa (Morocco, Algeria, and Egypt), plus two samples from Central Europe (Germany) and sub-Saharan Africa (Ivory Coast) as external references.

Granted, the Ivory Coast sample is representative of sub-Saharan gene pool, but it only serves as a part of that gene pool, not the whole of it. It is highly questionable that this Ivory Coast sample will contain all that that is present in sub-Saharan Africa, as opposed to giving a snapshot of what is present in sub-Saharan Africa. Furthermore, what purpose does it serve, to ignore populations situated between Ivory Coast and those in the northern African territories sampled? Common sense intimates that such an undertaking will ensure more abrupt changes in DNA marker distribution trends; but then again, the authors could be gunning for just that. From their frame of thinking, such relative abrupt change in pattern could serve to sift out what they think could be representative of the autochthonous northern African patterns. After all, the authors reckon:

This pattern identifies Mediterranean populations as genetically separate from both sub-Saharans and Central Europeans and allows the identification of a certain genetic structure between the two shores of the Mediterranean region.

There is still a problem with that perception, because while genetic exchange is expected between coastal northern African and the more-inward African populations, their primary ancestry has been liked to northeastern Africa [the Sahel region or southern confines of eastern Sahara; in other words, the belt or areas that seem to have been neglected in the study] and ultimately sub-Saharan eastern Africa. Certain information is bound to escape the authors' observation, with such sampling choices. Let's examine the sampling particulars, visually:


Click on the image for better res.

It is highly questionable that the Ivory Coast sample will be representative of all that which is part of the southern Sahara or Sahel belts. It is any wonder the Siwa sample assumed an "outlier" position on the authors' admixture analysis mapping, even though the Siwa, like the rest of the northern African populations sampled, are largely Imazighen and also live on the coastal areas of northern Africa. The Siwa sample noticeably maintains a good deal of distance from the Ivory Coast sample as well, not withstanding observations that sub-Saharan gene flow appears to be most significant amongst them vs. the other northern African samples.


Click on the image for better res.

Image caption: Multidimensional scaling plot (stress 0.036) applied to the Reynolds’ genetic distance matrix based on 18 autosomal Alu markers.

This sampling choice might account for the seemingly discordant observations in the 18 Alu "admixture" estimations and that of the Alu-STR combination "admixture analysis". The authors note:

In this general view, it is worth noting the particular position of two populations (the Spanish Pas Valley and the Egyptian Siwa Berbers) (see Fig. 2). These two populations have previously been described as genetic outliers (Esteban et al., 2006; Moral et al., 2006; Coudray et al., 2009) due to the orography of the Pas Valley and the desert surrounding the Siwa Oasis. This isolation could explain their differentiation by the action of the genetic drift associated with episodes with low effective population size, which in the case of Siwa Oasis, could have enhanced the effect of sub-Saharan flow (51% from Alu/STR data) through the Nile River (Fakhry, 1973).

Assuming one went by the earlier theory of the so-called Near Eastern Upper Paleolithic origin for coastal northern African populations, shouldn't the basic genetic structure of these populations therefore be the same, even when the effects of genetic drift are accounted for? Not only does the Siwa sample cluster away from the lone sub-Saharan sample of Ivory Coast, but also considerably does so from the coastal north African bunch, just going off on the 18 autosomal Alu markers alone...

MDS representation of the genetic distances (see Fig. 2) based on autosomal Alu data stresses the main differentiation of sub-Saharans, the clustering of Mediterraneans in two different groups corresponding to northern and southern populations, and the distant position of the Egyptian Siwa and the Spanish Pas Valley samples from their corresponding population clusters. The Siwa oasis sample presents a relatively extreme position, with respect to the other populations. In fact, the first genetic boundary in the Mediterranean separates Siwa Berbers from all remaining groups.

For visual aid, we have the following:


Click on the image for better res.

Image caption: Fig. 3. Multidimensional scaling plot (stress 0.049) applied to the Reynolds’ genetic distance matrix based on three Alu/STR compound systems.

Furthermore, if "sub-Saharan" gene flow was able to reach the Siwa, then how could they be considered "isolated"? Certainly the desert areas in northern Africa have not "isolated" the other coastal northern African groups. "sub-Saharan" gene flow reportedly finds expression in many of the coastal northwestern African Imazighen populations sampled, as reaffirmed by the pattern seen in the Alu-STR clusters; however, we are told that in the Siwa sample's case, "sub-Saharan" gene flow is virtually negligible in Alu pattern alone. Yet, the same Siwa sample is supposed to be indicative of the highest "sub-Saharan" gene flow amongst the coastal northern African Imazighen groups, going by STRs linked to certain Alu sites.

As for individual populations, the sub-Saharan gene flow in North Africa based on the Alu data collection ranges between 6 and 17% (Table 3), except the Siwa Berbers where that influence was negligible. Admixture values based on Alu/STR combinations indicate that sub-Saharan flow in North Africa ranged from 16% (North East Moroccan Berbers) to 35% (remaining samples) with the exception of Siwa Berbers who showed the highest admixture value (51%).

How was the relatively lower "sub-Saharan" contribution able to find expression in Alu markers of the other coastal northern African populations, but the more significant "sub-Saharan gene flow"— as communicated in the Siwa Alu-STR combinations— almost not represented at all in the Siwa Alu markers alone? Are we to assume that genetic drift enhanced "sub-Saharan" STR patterns but minimally did the same for Alu markers? The authors attribute this phenomenon of their finds in the following manner:

The disparity between the results from Alu loci and Alu/STR haplotypes, apart from the potential effect of the different number of independent markers examined (18 vs. 3), could be related to different mutation rates and therefore the power to detect ancient or more recent demographic events. Similar disparities between these two kinds of markers were found in the admixture analysis (Table 3).

Now of course, only three types of autosomal Alu loci were selected for examination along with flanking STRs, which tells us little about change in mutation rates across the genome types used here, and to what extent STRs on the other locations are useful enough in determining gene flow, along with whether this is in line with the data provided by the three type of sites used here. Undoubtedly different mutations rates between STRs and Alu markers could be a factor at some level, but the pattern we see in the extent of "sub-Saharan" gene flow across the full range of markers used in this study, may be more explainable in the sense, that 1) if the Siwa sample sufficiently comprised of identical Alu markers on chromosomes that share their immediate TMRCA nodes with sub-Saharan counterparts, then it could be distributed in such a way that it would be hard to ascertain gene flow from "sub-Saharan" populations with any degree of precision, or 2) some differentiation in Alu allele representation and nucleotide manifestation could be the product of within-population mutational events of markers with a "sub-Saharan" background in the Siwa, possibly in an interplay with that "action of genetic drift" that the authors mentioned in a piece cited above and some level of external gene flow from neighboring non-African territories, or yet 3) if the basic genetic structure of the Siwa stemmed from a non-African source, but then got introduced to "sub-Saharan" gene flow in an ancient period, and the population had since then remained relatively isolated from such influences ("sub-Saharan"). Only here, in either scenario, some visible level of Alu allele similarities would have come to the surface within those samples that reportedly tested positive for said "gene flow".

The first scenario doesn't seem to be likely, based on the 18 Alu makers multidimensional plot, given the position the Siwa sample assumes. While the third scenario could be presumptuously insinuated from the 18 Alu markers plot, given the considerable distance between the Siwa and that lone sub-Saharan African sample from the Ivory Coast, not to mention the possible case of the relative narrower distancing from the European clusters when compared to the African counterparts, including the so-called "southern Mediterranean samples" (coastal northern African samples), it is doesn't seem likely either; why? One would have to assume that while the Siwa might have been introduced to "sub-Saharan gene flow" at some point in time, it would have likely been a very occasional affair, and/or a very ancient one in the ethnogenetic history of the population, because this gene flow would otherwise not be negligible from across just the 18 Alu markers standpoint, even if the Siwa were of a small effective-population size subjected to heightened "action of genetic drift", and would therefore find expression as it did, in the other coastal northern African samples and European counterparts. Furthermore, the problem with that assumption is that the Siwa would likely have assumed a position more extreme than that of the "northern Mediterranean" samples in the 18 Alu markers multidimensional plot, from the lone sub-Saharan sample of Ivory Coast. The reason for this, is that elements of the "northern Mediterranean" samples would have become continued recipients of "sub-Saharan gene flow" either directly from sub-Saharan emigrants, and/or indirectly through continued contact with the "southern Mediterranean" populations aka coastal northern Africans. So the inclination here, is to go with the second scenario, and here's why: If the range of Alu markers were one or several step derivatives of autochthonous African counterparts, developed within the Siwa population during its ethnogenesis, then naturally, these markers would stand in contrast to ancestral sub-Saharan counterparts. As such, one would expect some level of persistence in some areas of the genome type selected for this study, particularly given that the sites that were picked for STR analysis happen to be those under linkage disequilibrium, according to the authors. So, while subsequent "sub-Saharan gene flow" cannot be ruled out in this scenario, it need not be the sole explanatory factor for the sub-Saharan inclinations of Siwa Alu-STR combinations, i.e. if the Siwa were treated as group that has been socio-culturally isolated from other external groups for some reason or the other. Possible additional external gene flow from nearby "non-African" territories, again likely ancient, cannot be ruled out under this scenario, in which case, such element would only serve to further contrast the Siwa Alu distribution from the sub-Saharan counterpart examined here...

The Siwa oasis sample presents a relatively extreme position, with respect to the other populations.

Under this scenario (2nd scenario), one can see why the so-called "southern Mediterranean" samples would assume intermediary positions along both types of multi-dimentional plots provided by the authors. The polarity here likely stems from a mix of continued, and hence more recent gene flow from external populations both African and non-African, along with in situ autochthonous within-population evolutionary events in said "southern Mediterranean" populations. Either of these factors would ensure that their socio-cultural and geographic distance from the Siwa would contribute to the differentiations in general Alu marker genetic structure, while at same time clustering them away from European clusters and the lone sub-Saharan sample. See:

the clustering of Mediterraneans in two different groups corresponding to northern and southern populations, and the distant position of the Egyptian Siwa and the Spanish Pas Valley samples from their corresponding population clusters.

 Furthermore,...

This pattern identifies Mediterranean populations as genetically separate from both sub-Saharans and Central Europeans and allows the identification of a certain genetic structure between the two shores of the Mediterranean region. This genetic picture of populations may be related to geographic factors as indicated by the high correlation (P < 0.002) between geographic and genetic distances (based on Alu markers) found under the isolation by distance model. The genetic distinctiveness of Mediterranean populations, as well as the distinction between Northern and Southern Mediterraneans, coincides with results in previous studies (see for instance, Simoni et al., 1999; Comas et al., 2000; Boschet al., 2001).

Furthermore...

The estimates of sub-Saharan gene flow in Southern Mediterraneans oscillated between 12.9% (Alu loci) and 39.5% (Alu/STR haplotypes), a wide range probably related with the different mutational nature of the markers analyzed and with the effect of repeated homoplasic mutation in STRs.

One might expect the effect of genetic drift to pick up these elements in the Siwa as well, if one is to treat said "sub-Saharan" gene flow level as largely the product of action of random genetic drift in a population of small effective-population size.

The presence of sub-Saharan African traces in the gene pool of North Africans supports the idea of the permeability of the Sahara desert to human migrations as reported in other studies for different kinds of markers (see for example, Plaza et al., 2003; Arredi et al., 2004; Myles et al., 2005; Coudray et al., 2006).

In the above, the authors seem to have no problem in acknowledging the fact that a desert environment, of the Sahara, has not restricted or barred gene flow. This means that these groups are not isolated by the desert; so why couldn't the same logic be approached with regards to the Siwa? It may well be the case, that the Siwa have socio-culturally isolated themselves from other coastal northern African Imazighen groups on their own terms, not to mention the considerable distance between them and the other coastal northern African populations sampled, and has little to do with the desert environment. One will note that even as far as neighboring territories go, which here are apparently European territories bordering the Mediterranean sea, the territory that the Siwa are identified with is relatively more distant from the nearest such territory than those associated with the other coastal northern African populations, respective to their nearest neighbor, This could explain the differentiation in genetic structure and their relative "outlier" position. The authors add:

Interestingly, data from mtDNA and Y-chromosome estimates of sub-Saharan gene flow in North Africa are similar to that obtained from our Alu loci set, a value also concordant with that corresponding to Mozabites in the recent survey of Li et al. (2008) based on more than 500,000 SNPs. The interpretation of the disparity in gene flow estimates according to the kind of marker is difficult, but it might be presumably be related to the different mutation rates of Alu and STRs.

The need to confide in uniparental lineage is not obviously underestimated, but the authors allude once again to the unpredictability characterizing their choice of markers, autosomal markers in the form of Alu sites and flanking tandem repeats at certain designated sites. As we have seen in an earlier piece, the chiming in of homoplasic tendencies in STRs does not dampen this unpredictable character. Of course, since we are dealing with autosomal sites, the question of recombination cannot be avoided. We are assured here, at least with regards to the Alu-STR combinations, that these are perceived to be the types in linkage disequilibrium.

Alu/STR linkage disequilibrium was present in all systems and samples.

This is a sure sign of non-random associations here, which means that the odds against random reshuffling by recombination are high and hence, possibly of some selective pressure advantage of the Alu/STR association. This naturally factors further into that matter about "different mutation rates" and no less, contributes to the unpredictability character of the change in mutation rates in different parts of the genome.

Notwithstanding the lone sub-Saharan sample of Ivory Coast, upon revisiting the matter, one notices that it still managed to give a snapshot of the fact that non-African populations are just representative of a subset of African gene pool: 

When STR variation has been analyzed separately in Alu(+) and Alu(-) chromosomes, larger variances are observed in chromosomes carrying the ancestral Alu variant: CD4(+), FXIIIB(-), and DM(+). In humans, the ancestral stage of the CD4 and DM loci is the presence of the Alu insertion, whereas the absence of the insertion is the ancestral stage for the FXIIIB locus (Brook et al., 1992; Nishimura and Murray, 1992; Tishkoff et al., 1996 ). Alu/STR linkage disequilibrium was present in all systems and samples.

The most obvious pattern of haplotype variation is observed in the CD4 system. The ancestral CD4(+) chromosomes show a decreasing pattern of copy number variation from sub-Saharans to Southern and Northern Mediterraneans. Among these latter populations, the 85(+) and 110(+) haplotypes are the most frequent (Supporting Information Table 2). The derived CD4 Alu(-) chromosomes present a lower variation than the ancestral Alu(+) chromosomes, which is statistically significant for Northern Mediterraneans (P < 0.01) and Southern Mediterraneans (P < 0.05), but non-significant for the sub-Saharan sample. This reduction trend is considerable in Northern Mediterranean samples (gene diversity: 0.174 for derived chromosomes vs. 0.554 for ancestral ones), moderate in Southern Mediterraneans (0.458 vs. 0.705), and less marked in sub-Saharans (0.721 vs. 0.778).

The ancestral markers are disproportionately higher in "sub-Saharans", which in this case as we know, is based on that lone sample from the Ivory Coast, and then, they are moderately represented in "southern Mediterraneans", which would be our coastal north African samples here, and least represented of all the groups herein, in the "northern Mediterranean" samples, which here would be the southern European samples. Respectively, greater nucleotide variation is found in "sub-Saharans", as characteristic of the ancestral markers, moderate diversity in coastal northern African, and least diversity in Europe. This seems to find some expression in the general positions assumed by the samples in the plots respective to each marker-format type; in each case, the northern African groups appear to be in the intermediary positions between the African samples in the extremes and the European ones on the other hand. Along the x coordinates of the multidimensional scaling plots provided to us by the authors of the present study, the Ivory Coast sample consistently attains the most extreme position on one end. Please refer back to the plots or maps provided earlier in the body of this post.

Of the derived examples of the Alu-STR clusters, the distributions patterns found in the present study suggest possible "southern Mediterranean" or coastal Northern African origins (or at least, populations ancestral to them) for the following types: CD4 110(-) and DM 107(-)

The highest frequencies of CD4 110(-) and DM 107(-) have been found in the High Atlas region (7 and 5.5%, respectively) of Morocco, reaching polymorphic frequencies in all the North African samples [barring the Mozabites for the CD4 110(-) combination]. They have also been found in the Iberian Peninsula, scattered along the northern Mediterranean shore to Greece and Turkey, and on the main islands of the western Mediterranean (Majorca, Corsica, Sardinia, and Sicily; González-Pérez et al., 2007). The CD4 110(-) haplotype (Flores et al., 2000) and has also been reported in West Saharans and Mauritanians on five of the seven Canary Islands (Flores et al., 2001), as well as in Adygei from the Northern Caucasus (Tishkoff et al., 1996). Assuming from their frequency distribution that the place of origin of these particular haplotypes is located in the westernmost extreme of North Africa (Fig. 4A,B), their current ample distribution along both shores of the Mediterranean most likely reflects the effect of gene flow across the region since ancient times, even though specific ages cannot be accurately estimated with our data. Similarly, specific Mediterranean haplogroups or clades (U6 and M1b in the mtDNA; EM78 and EM81 in the Y-chromosome) have also been described for these populations and dated in Paleolithic times.

Last but not least, in keeping with pointing out the recurring theme of the lingering onto outdated or outmoded and subjective concepts by the authors of the present study, the following serves as further example:

Concerning Northern Mediterraneans, the gene flow from sub-Saharan Africa was inappreciable for Alu markers and swung from 6 to 15% for the Alu/STR haplotypes data calculations. When gene flow in Northern Mediterraneans was tested, taking Central Europe and Southern Mediterraneans as parental populations, the results were statistically inconsistent, indicating the limited power of our markers to discriminate gene flow within Caucasoid populations. Nonetheless, the distributions of frequencies for the Mediterranean haplotypes CD4 110(-) and DM 107(-) (Fig. 4A,B) are suggestive of gene flow processes across this geographical region.

Such recurring themes throughout different parts of the study do not bode well for the authors at hand.

*Keep an eye on possible future updates.
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*References:

— As already cited in the body of the post.