Sunday, June 21, 2009

Reviewing: The Genetic Structure and History of Africans and African Americans

This is a carryover from the "miscellaneous notes" of the last blog posting (The Creation of a Fake Controversy: The Fula Origins). It pertains to extracts taken from: The Genetic Structure and History of Africans and African Americans, a publication by Tishkoff et al (2009). that just came out recently. Here, we will examine the work a bit more:

Sampling entailed...

We studied 121 African populations, four African American populations, and 60 non-African populations for patterns of variation at 1327 nuclear microsatellite and insertion/deletion markers. We identified 14 ancestral population clusters in Africa that correlate with self-described ethnicity and shared cultural and/or linguistic properties.

And mentions of interest here...

Ethnic groups, sample size, language classification, and subsistence classification are given in Table S1. For the Kenyan, Sudanese, Nigerian, and Cameroonian samples, white cells were isolated in the field from whole blood with a salting out procedure modified from (S1) and DNA was extracted in the lab with a Purgene™ DNA extraction kit (Gentra Systems Inc., Minneapolis, MN). Dogon samples were obtained from blood spots donated by participants in a cohort study of malaria incidence in Bandiagara, Mali. Ghanaian DNA was extracted onsite from whole blood, with the Purgene™ DNA extraction kit. The Juspeaking !Xun (a.k.a.Vasekela) and Khoe-speaking Khwe samples were collected from individuals in the area of Schmidtsdrift in the North-West Cape of South Africa. The Cape Mixed Ancestry (CMA) population, commonly referred to as Cape Coloured in South Africa, was collected in the Western Cape Province. The Dogon sample was collected in Bandiagara, Mali. Nigerian samples were collected in Abuja and Adamawa State districts. Cameroon samples were collected from the Eastern Province (Baka Pygmies and neighboring Bantu groups), Southern and Ocean Provinces (Bakola Pygmies and neighboring Bantu groups and coastal groups), Center Provinces (Medzan Pygmies and neighboring groups, mostly Bantu populations), Western Province (Bamileke and Mbororo Fulani groups), Extreme North Province (Mandara mountains and northern plains; Fulani and Afroasiatic/Nilo-Saharan speaking populations). All Cameroonians were sampled in their native village; the Hausa sample (a population who emigrated mainly from the Kano area two generations ago) was sampled in the city of Yaounde. Samples from Chad, CAR, Congo, DRC and Rwanda were obtained from individuals who recently immigrated to Cameroon. Tanzanian DNA samples were collected from individuals residing in the Arusha and Dodoma provinces of Tanzania. Samples from Kenyan populations of southern Ethiopian origin (Burji, Konso) samples were collected in the Rift valley, Nyanza, and Eastern provinces of Kenya. Sudanese samples were collected in the Khartoum and Kasala provinces of the Sudan. The Yemenite Temani and Ethiopian Beta Israel samples were purchased from the National Laboratory for the Genetics of Israeli Populations (S2) . The South African !Xun/Khoe, Xhosa, Vende, Cape Mixed Ancestry (CMA), Yemenite Temani, Beta Israel, and the Malian Dogon DNA samples were amplified by Whole Genome Amplification (WGA) with Illustra GenomiPhi HY™ kits provided by GE-Healthcare (Buckinghamshire, UK). It should be noted that the DNA for the Dogon population extracted from blood spots appeared to be of lower quality and microsatellite markers did not amplify as well as other samples obtained from whole blood (43% of markers had missing data).


Abstract:

Africa is the source of all modern humans, but characterization of genetic variation and of relationships among populations across the continent has been enigmatic. We studied 121 African populations, four African American populations, and 60 non-African populations for patterns of variation at 1327 nuclear microsatellite and insertion/deletion markers. We identified 14 ancestral population clusters in Africa that correlate with self-described ethnicity and shared cultural and/or linguistic properties. We observed high levels of mixed ancestry in most populations, reflecting historical migration events across the continent. Our data also provide evidence for shared ancestry among geographically diverse hunter-gatherer populations (Khoesan speakers and Pygmies). The ancestry of African Americans is predominantly from Niger-Kordofanian (~71%), European (~13%), and other African (~8%) populations, although admixture levels varied considerably among individuals. This study helps tease apart the complex evolutionary history of Africans and African Americans, aiding both anthropological and genetic epidemiologic studies.

Right off the bat, the first thing that probably jumps at any reader, is the heavy reliance on microsatellite repeats, as opposed to uniparental binary markers. Restricted fragment length nucleotide sequences or polymorphisms [relying on insertions/deletions] can add precision to identifying discrete monophyletic units, but even here, the results are not always as clear cut as the case generally is with "classic" uniparental binary markers of Y-DNA and mtDNA respectively, relying on unique event SNPs. For instance, certain RFLP haplotypes have been linked to two different haplogroups; case studies on RFLP haplotypes on this website have shed light to such occurrences: see Lucotte et al.'s haplotype IV, RFLPs: Lucotte et al., A case study — Pt. 1, and RFLPs: Lucotte et al., A case study — Pt. 2. In the first link of the three for instance, we were told:

A total of 21 different 49a,f haplotypes were found and are illustrated in Fig. 4 as a sub-classification of the Iraqi Y-chromosome haplogroups. The most represented haplotype of haplogroup E is haplotype 5 (A 2 C 0 D 0 F 1 I 1 ). This is followed by haplotype 11 (A 3 C 0 D 0 F 1 I 1 ) at a much lower frequency. Haplotypes 5 and 11 were observed both in Africa (Lucotte et al., 2001; Passarino et al., 1998; Persichetti et al., 1992; Santachiara-Benerecetti and Semino, 1996; Spurdle and Jenkins, 1992; Torroni et al., 1990) and Eurasia (Passarino et al., 2001; Semino et al., 2000b) but in Africa they belong to haplogroup E, whereas in Eurasia, particularly in *Northeastern Eurasia*, they belong mainly to the haplogroup R-M17. Interestingly, the proportion of haplotypes 5 and 11 in haplogroups E and R-M17 is reversed, with haplotype 5 prevalent in haplogroup E and haplotype 11 in haplogroup R-M17. By considering that the two haplotypes differ by a single band change and their different proportion in the two lineages, it is likely that haplotype 11 is a derivative of haplotype 5 in haplogroup E and just the opposite in haplogroup R-M17. - N. Al Zahery et al. 2003, Y-chromosome and mtDNA polymorphisms in Iraq, a crossroad of the early human dispersal and of post-Neolithic migrations.

In cases where it might be difficult to link a RFLP haplotype to a specific sub-clade or else superclade, classic uniparental (Y-DNA or mtDNA) SNP and/or UEP markers are called on, as we see in the aforementioned link 2 and link 3 on Lucotte et al. Questionable allocation(s) of ancestral origin based on "shared" microsatellite markers and/or insertion/deletion markers is therefore bound to occur, as one comes across in the study at hand; ancestry, which would otherwise actually be more a reflection of an ultimate shared common recent ancestry prior to the ethnogenesis of the implicated contemporary groups into recognizable ethnic or socio-geopolitical entities they are known today, as opposed to some recent unidirectional gene flow from one designated contemporary group to another.

Moving onto excerpts of interest...

As some have correctly picked up on, and as the authors themselves openly admit, there are issues with their Dogon sample that need to be ironed out...

Detection of relative pairs: Relative pairs and duplicated samples in the dataset were inferred from the pattern of shared genotypes and population allele frequencies with RELPAIR 2.0.1 ( 6-8) . Because the inclusion of closely related individuals can impact population genetic inferences [e.g. (S9)], we took the conservative approach of excluding individuals inferred to be third degree or more closely related, including inferred relative pairs between regional ethnic populations (e.g. all Tanzanian populations). An exception was made in the case of the Dogon as it is difficult to reliably infer relative pairs in a small sample and the Dogon are highly distinctive and could not be readily merged with other populations to improve allele frequency estimates. Merging the Dogon with other non-Pygmy West African populations inferred four unrelated individuals in the sample, but this may be overly conservative given the distinctiveness of the Dogon sample from other West Africans. Also, the Dogon are the only representatives from Mali in our study and since the sample size is already small we did not want to further reduce the sample size in the analyses, especially if the relative pair estimates were questionable. Therefore, RELPAIR inferred relative pairs among the Dogon were not excluded. In total 737 individuals were removed. Networks of relatives, which in some cases were quite complex, were plotted with neato from the GraphViz software package (S10), which was used to select the minimum number of individuals to exclude to break up networks of relative pairs.

Indeed, the Dogon sample size, which to reiterate what the authors tell us consisted of candidates who could also possibly be closely related and the only sample that represented Mali, was only one of 9 individuals, a fairly modest size. Keeping this in mind, we are told:

The Dogon from Mali, who speak a Niger-Kordofanian language, cluster with the Saharan and eastern Africans in the NJ tree (Fig. 1), consistent with the results from STRUCTURE analysis, showing considerable Saharan (blue) and Afroasiatic (purple) ancestry, and consistent with oral history of a northern African origin (although it should be noted that the sample size for this population, 9 individuals, is very small and many markers did not amplify well) (Fig. S13; Table S9).

"Saharan" without regional specificity is next meaningless. Just about any of the groups on the Sahara, Sahel or adjoining "sub-Saharan" territories have "Saharan ancestry". Furthermore, as has been the talk of this site recently, relics of "fertile-Sahara" era or earlier east African ancestry is scattered along the length of the Sahara, and generally finds expression in Sahelian and "Sub-Saharan" populations in relatively low incidences, if not somewhat patchy distribution. Such shared Saharan ancestry is bound to have some sort of manifestation across bi-parental microsatellite markers as well. As we've just seen, the authors note that "many markers did not amplify well", and indeed, if there is any further indication of an air of dubiousness about the results of the amplifications, then perhaps it is notion of some 45% ancestry, or specifically 45% of the sample identified with an AAC (Associated Ancestral Cluster) , being suggestive of European ancestry in that small sample of Dogon, with 9 individuals [see table S8 of the publication]; and even if one were to remotely entertain the possibility as a hypothetical scenario for argument's sake, it would implicate these individual as essentially "outliers", if anything, based off on previous information published on samples taken from sections of Malian society, including the Dogon. Additionally, with regards to such an outlier situation, one might even ask if this is not a function of "third party" interaction, with that "third party" being coastal north African Imazighen and Arabized groups. The authors themselves base AAC assignment on where said markers are allegedly the most maximum in 'frequency'...

African and non-African Associated Ancestral Clusters (AACs; labeled based on the populations showing the highest levels of ancestry for each inferred ancestral cluster)are highly divergent.

One can suppose 'frequency' could be a telltale sign of geographical origin of certain biparental microsatellite clusters, if only tenuously by itself, but as a matter of precision, it effectively plays that role only when taken into account with other molecular genetics information, like the internal molecular variation of a designated clade, and in association, how much of this variation is paraphyletic, along with definite geographical patterns of distribution. For instance, one might surmise that Hg R1b originated in Europe, merely on the account of the haplogroup's unparralleled prevalence in that region, and place the maximum likelihood of finding ancestral R1b clades in Europe before finding them elsewhere. However, as a matter of merely "frequency", Hg R1b's incidence in Europe in no way allows inference that either Hg R1* or its descendant, the upstream clade R1b* originates there; in fact, R1b* clades bearing upstream SNPs that are nearer to the root of that sub-clade, but lacking the well-known downstream SNPs, have so far only been identified in non-European samples. Interestingly, these non-European samples happen to be African ones. At the same, ironically, R1b itself is relatively rare in Africa, even in the coastal north African areas which are closer to Europe. So frequency as a single factor for adjudging origin can be misleading. Neutral factors like random genetic drift, for example, can elevate a marker's frequency in a designated locale, and yet, this locale in of itself need not be the actual location of origin for said marker. For microsatellite repeats, in absence of binary or UEPs, the assignment of origin based on frequency alone becomes even more uncertain, than the case is for binary or SNP markers. Microsatellite repeats could even be shared both between different and within single monophyletic clades in some occasions, due to homoplasy.

Many questions abound the objectivity of result inferred for the small Malian sample. In a related matter, the Beja samples were implicated in some ~ 32% and ~ 34% AACs [the Banuamir and Hadandawa samples respectively] placed under "European" [see: Table S8 of the journal]. Well, to the extent that ancestry suggestive of "possible" European origin has been implicated in a Beja sample before, it is reasonable to assume that this may render a section of the Beja to display shared clusters with Europeans in a segment of nuclear DNA that is outside uniparental binary markers; however, it is worth noting that the uniparental gene pool, the paternally-inherited one in particular, featured fairly low incidence of ancestry suggestive of ultimate European ancestry. Citing Hassan et al. (2008), the Beja male gene pool comprised of only ~ 5% ancestry that could *potentially* be linked to European origin [Hassan et al. (2008) provide little information on specific downstream clades of Hg R1b]; this is fairly consistent with R1b's general distribution in northern Africa, where it is generally low, in contrast to the fairly considerable representation of European maternal ancestry in coastal northwest Africa. So, the 32 or so % of microsatellite AAC linked to a possible European origin is out of character, when uniparental ancestry information is taken into account. On the other hand, substantial AAC sharing between coastal northwest African samples and the European ones is consistent with uniparental DNA examinations, which to reiterate, mainly finds expression in the maternally-inherited gene pool.

Mitochondrial DNA analysis indicates that Fulani have lineages of predominantly West African origin and that they cluster together and close to the Mandenka population from Senegal [S93].

By contrast, Y chromosome analyses of Fulani
sampled in the Sudan indicates shared ancestry with Nilo-Saharan and Afro- Asiatic speaking populations [S89].

The line about the mtDNA is essentially a reversion back to what has been cited in the main post above, as observed by Cerny et al. (2006). The line about the Y chromosome analysis however, is quite misleading; one only needs to refer to Hassan et al. (2008) [see: Y-Chromosome Variation Among Sudanese] to see how: The Sudanese Fula sample here, as noted in the main post above, predominantly featured Hg R1*-M173 markers in the uniparental paternal gene pool. This marker is in fact generally quite rare in "Afro-Asiatic" speaking populations; rather, its highest incidences has thus far been reported in Niger-Congo, Nilo-Saharan and "Afro-Asiatic" (Chadic) speaking groups in northern Cameroon, with smaller occurrences in the rest of central Africa [see Luis et al. 2004]. Nilo-Saharan groups in Sudan itself, it should be noted, did NOT share this ancestry with the Sudanese-based Fulbe at all. This reverts back to what had already put forward in the main post, about the implication being that these Sudanese Fula derived from a section of the northern Cameroonian Fula, wherein the Hg R1*-M173 is featured in a considerable but not the predominant frequency.

The only marker that the Sudanese Fula shared with Nilo-Saharan and Afrasan ("Afro-Asiatic") speaking Sudanese groups was Hg E3b1 (M78). Common sense however, intimates that this aspect of the paternal gene pool was largely picked up from neighboring populations, only after the Fula situated themselves in Sudan. It makes sense, since after all, E3b1 is one of the prominently featured markers found in the Sudanese sample.

To recap, there's something still unmistakable about the Sudanese Fula sample: they retain western African ancestry, as signified by Hg E1-M33, which was absent in all autochthonous Sudanese groups; the only other groups where this marker was implicated in Hassan et al.'s (2008) work, were, well, the well-established west African groups like the Hausa and Wolof. When pressed on to explain away this anomalous feature of the Sudanese Fula's paternal gene pool, dissidents of the west African origin noticeably become mute on the matter.

These results raise the possibility of differential patterns of male and female gene flow into this population.

Whatever may be "differential" about male and female gene flow patterns in Fula sample, no doubt influenced by matters like life style recalling on nomadic vs sedentary, both the maternal and paternal gene pools communicate the same thing: west African origin!

Our analysis, using genome-wide nuclear markers and STRUCTURE, indicates that the Fulani have distinctive ancestry [fuchsia] at K = 14 in the global analysis [Figs. 3,4] and at K = 9 -14 in the Africa analysis [Fig. S13]. The Fulani cluster with the Chadic and Central Sudanic speaking populations at K <13> They also cluster near the Chadic and Central Sudanic speaking populations in the NJ tree based on population genetic distances [Fig. 1].

Fula samples taken from either central Africa or Sudan are expected to cluster with groups in central Africa-Chad, as the Fula would have arrived in Sudan via a central African and/or Chadic corridor, having experienced a bottleneck [see main blog post]. The central African Fula in turn would have arrived ultimately from Atlantic-coastal regions of western Africa, where their ethnogenesis as the Fula-"proper" likely occurred.

The pattern of the Sudanese Fula's Y-DNA composition is for instance, undoubtedly different from their western African brethren, but continuity still exists via recurring typical western African markers that unite all Fula groups, regardless of where they spread to; e.g. primarily Hgs E1 and E3a, and a lesser extent, Hg E2. Either of these clades are telltale signs of so-and-so Fula groups' western African ancestry.


In the global STRUCTURE analysis, the Fulani show low to moderate levels of European/Middle Eastern ancestry [blue], consistent with mtDNA and Y chromosome [S89] analyses, as well as the presence at low frequency of the -13910 mutation associated with lactose tolerance in Europeans in this population [S94].

Again quite misleading. Yes, understandably, "low" levels of shared ancestry attributable to potential "European" ancestry, might be correlated with "low level" incidences (~1.7 %) of R1b [pertaining the Rosa et al.'s (2007) Guinean sample] or relics of this ancestry attained from coastal northwestern populations [Imazighen] seen on the mtDNA side [Cerny et al. 2006], as cited in the main post. However, in the case of Fula samples, from northern Cameroon through to Sudan, a good degree of presumed shared "European/Middle Eastern ancestry [blue]" may very well actually be linked to common "distant" ancestry, as the paraphyletic R*-M207 and R1*-M173 markers suggest; these latter two are essentially rare to absent in "Middle Eastern" and "European" populations, which are mainly characterized by more downstream markers, especially European populations, who have no upstream markers for R1. Now, because R1b and R1a markers are still 'molecularly' linked to the upstream R*, any group bearing the latter or else any intermediary nodes between the upstream R* and the downstream R1a and R1b will "appear" to share ancestry with Europeans and/or possibly "Middle Easterners", which could misleadingly be misread as ancestry from "European/Middle East". Given the features of northern Cameroonian and Sudanese Fula sample uniparental male gene pool, it is a safe bet that this is the underlying issue at work, especially when the red flag of "moderate" levels is evoked; the Fula uniparental gene pool specifics, whether maternal or paternal, simply do not bear out "moderate" ancestry from "Middle East" or "Europe".

Recalling a previous blog post here
, R1*-M173 in Africa, we have:

Interestingly, upon revisiting Wood et al. (2005), it should be pointed out that paraphyletic clade of R*-M207 was detected amongst some "Afro-Asiatic" African groups, along with the paraphyletic clade R1*-M173 [it is worth noting that Wood et al. implicate the Egyptian sample here as something other than that of Semitic speakers (Arabic)], while some Niger-Congo groups — though in small frequencies [pooled] — tested positive for the paraphyletic R1b*, lacking the established downstream R1b markers. Henceforth, R*-M207, lacking downstream mutations have been identified in African groups via this study; and yes, the basic nodes of all presently known Hg R's downstream clades had been accounted for, which means that R*, as predicted above, is NOT relegated to the Indian sub-continent. All in all, this suggests that African Hg R pool is actually more diverse than many seem to think.

As for the matter concerning lactose tolerance, it has been pointed out here before, how this has nothing to do with European ancestry. Recap, see:
More on R1*-M173 bearers

In summation, the piece essentially rehashes the well-understood facts of primary west African ancestry of the Fula, both in terms of genealogy and language, along with genealogical "admixture" elements picked up over the course of migratory events, but suffers from matters, as laid out in detail in the main blog post, pertaining to idealistic urges for mystification where none is necessary or warranted. To provide an example, consider the following line:

Our analysis, using genomewide nuclear markers and STRUCTURE, indicates that the Fulani have distinctive ancestry (fuchsia) at K = 14 in the global analysis (Figs. 3,4) and at K = 9 -14 in the Africa analysis (Fig. S13). The Fulani cluster with the Chadic and Central Sudanic speaking populations at K <13>Fulani show low to moderate levels of European/Middle Eastern ancestry (blue), consistent with mtDNA (S93) and Y chromosome (S89) analyses, as well as the presence at low frequency of the -13910 mutation associated with lactose tolerance in Europeans in this population (S94). Additionally, we observe moderate to high levels of Niger- Kordofanian ancestry in the Fulani populations (Figs. 3, 4, S13;Tables S8, S9). These results do not enable us to determine the definitive origin of the Fulani, although they indicate shared ancestry with Saharan and Central Sudanic populations and suggest that the Fulani have admixed with local populations, and possibly adopted a Niger-Kordofanian language, during their spread across central and western Africa. The origin of European (possibly via the Iberian peninsula) and/or Middle Eastern ancestry in the Fulani requires further exploration with additional genetic markers.
 

The words "admixed with local populations" is particularly interesting, as it is naturally inconsistent with mtDNA report [and Y-DNA] where western Africa is concerned, recalling Cerny et al. (2006) for example, who point out that their results show a primarily or predominantly west African ancestry of "charasteristic" markers of the Fula, but wherein the gene pool of nomadic Fula samples show very little genetic contribution from neighboring sedentary populations. This, to put it simply, means that the only way for the Fula gene pool to be primarily west African, is if they were autochthonously west African.

Another interesting choice of words, is the mention of "adopt" in Tishkoff et al.'s piece above, in relation to the Fulani language. "adopt" is usually suggestive of one taking on something that wasn't originally his/her's. Supposing one remotely entertained said insinuation of the authors for just a minute, their very idea that a small band of foreigners adopted the "Niger-Kordofanian" language along their migratory path, means that the locals along that path would have had to have already been Fula speakers; that is the only way a wandering group of immigrants/foreigners could have become speakers of a Niger-Congo language a language family that supposedly did not characterize their own which by chance, happens to be Fula. In other words, the Fula as an ethnic group was already established on the continent prior to the intrusion of this small band of immigrants; it's just common sense. Henceforth, this band of immigrants cannot be judged as proto-Fula antecedents by any stretch of the imagination; rather, they should be viewed merely as a foreign element that integrated into an already well established ethnic-group, i.e. the Fula, as opposed to assuming the role of a "prototype" for the group into which they allegedly integrated. To somehow use this [unsupported] scenario as something that mystifies Fula origins, is clearly not rational or logical deductive reasoning; at best, it is quite comical. The matter of Fula origins was recently posted here, in detail; recap: The Creation of a Fake Controversy: The Fula Origins

It is interesting to note that while the authors carry themselves as very confused individuals about Fulani ancestry, their own tables (see S8) show very little AAC sharing with Europeans; their Nigerian Fulani sample supposedly shared ~5 % or so AACs placed under "European", while the Cameroonian Fula samples shared about 2.5% and 2.6% respectively. Nothing "moderate" about these.

From language, culture to genealogical heritage, all indicators resolutely point to a west African ethnogenesis of the Fula. This has been supplemented with archaeological evidence, as that from Tassili, on rock art. Here, are renditions of a cattle-rearing group, bearing features reminiscent of contemporary Fula and/or western Sahel groups in general. One scholar often cited on the internet, regardless of how objective or otherwise the sites themselves are, for identifying definite links between the Tassili n'Ajjer rock painting and contemporary Fula, is Amadou Ba. For instance, one site notes:

At the Tin Tazarift site, for instance, historian Amadou Hampate Ba recognized a scene of the lotori ceremony, a celebration of the ox's aquatic origin. In a finger motif, Ba detected an allusion to the myth of the hand of the first Fulani herdsman, Kikala. At Tin Felki, Ba recognized a hexagonal carnelian jewel as related to the Agades cross, a fertility charm still used by Fulani women. With the disappearance of many traditions and other aspects of African culture, works of traditional African art are becoming more and more scarce. - This particular piece is from Jamtan.com

The following are from Andrew Brown Smith [African Herders (2005)], who seemingly attempts to portray himself as a "voice of moderation":






Of note here, is the observation that Smith as any other person guided by reasoning could not pass over the significance of linguistics, i.e. recalling the Niger-Congo familial ties of Fula language.



While briefly citing examples of the sort of reception Ba's definite correlations got in certain quarters, mainly amongst "western" observers, even Smith could not resist pointing out that the "coincidences" surrounding the reminiscence of Fula socio-cultural traits in definite features of the rock arts are simply too considerable to not draw a link between the two, i.e. between the figures in the rendition and contemporary Fula. Notwithstanding this openly-admitted acknowledgment, Smith of course, goes onto characterize Ba's demonstrations as "perhaps what they wanted to see to support their thesis", but offers no substantive counter-thesis that puts to question the "specifics" of what Ba "sees", other than a mere opinionated presumption of Ba's "possible" motive behind making the definite links that he made.





Click on the above images to get higher resolution versions.

In the lower images for example, one notices the head gear, which is not unlike that featured on a contemporary Dogon dancer in the image on the right hand side.

Other notes: Fulani cattle are also deemed to be of west African origin, as a sub-phyla of the west African breed of Zebu cattle called the "West African Zebu", and they fall into the following two groups -

The Fulani have been classified further into two groups: the lyre-horned subgroup consisting of Senegalese Fulani (or the Gobra), the Sudanese Fulani, and the White Fulani (or Bunaji); and long-horned subgroup represented by the Red Fulani (or Rahaji). Diali (or Djeli) is a strain of Fulani found on the flood plains of Niger river in Niger and south-west Nigeria (Rege 1999; Rege and Tawah, 1999). - courtesy cdad-is.org

Dictionary description...

Sudanese Fulani cattle
West African, lyre-horned, milking cattle, usually light gray.- Medical-dictionary

Continuing with examination of the study at hand, the authors note:

Individuals from Saharan and Eastern Africa show heterogeneous ancestry, reflecting descent from populations ancestral to non-Africans and/or gene flow from non-Africans into Africa.

Well, "exotic" gene flow as function of said heterogeneity, while reasonably a contributing element to some extent, is really quite trivial in the big scheme of things. African populations would still be quite heterogeneous *regardless* of gene flow from non-Africans
. In relation to this, a subset of Africans who formed the basis for non-African groups, would assume an intermediary position in a case study involving a "comprehensive" collection of African and some non-African samples, because quite simply, that subset of Africans only represents a portion of the overall African diversity. Non-Africans in turn, only have a portion of the diversity of the subset of Africans from whom they emanate.

*Watch for occasional updates or modifications of this post in future.



*Last edited on 10/26/2010.

Monday, June 15, 2009

The Creation of a Fake Controversy: The Fula Origins

From time to time, one comes across mystification of certain western African ethnic groups by certain reactionary cults and like ideologues, whether it is in terms of certain intra-regional phenotypic variation and/or origins. We've seen an example of this sort of mystification in an earlier posting right here, although in this case, the authors point to definite facets of evidence that yet again underlie fictitious underpinnings of such mystification:

Recalling...

The origins of the Fulbe are the subject of debate, but the group is thought to be from outside Cameroon; on the basis of ethnic traditions and linguistic similarities between Fulbe languages and Tukulor (Toucouleur), an origin in the Futa Toro region of the Senegal river basin has been proposed (Newman 1995). - See: More on R1*-M173 bearers

Two notorious ideological camps that are known for propping up such controversy are Eurocentric extremists [and their ideological sympathizers, be it so-called "moderates" or "liberals"], and radical self-proclaimed "Afrocentric" elements, generally made up of non-African citizens. These two camps will for instance, insist that the counterpart ideological camp doesn't influence its idealistic approach to matters such as the subject of this post, but in fact, an attentive examination of their respective propaganda shows otherwise. Radical Eurocentric elements for instance, generally point to the so-called "anomalous" facial features of the Fula (Fulbe/Fulani) and conclude that it can only be an expression of their "mixed" ancestry, i.e. between the "stereotypical Negro" archetype and the "caucasoid/Caucasion" archetype.

Interestingly enough, Cerny et al. (2006) in a passing note:

HVS-I analysis of four Fulani populations revealed the different proportions of the mtDNA gene pool. A major role is played by West African mtDNA haplogroups, such as L1b, L3d, L3b, L2b, L2c, and L2d, which together make up 79.6% of the whole. The far from negligible presence of some haplogroups from western Eurasia (8.1%), such as U5, U6, and J1, is not particularly surprising in a sub-Saharan context because these haplogroups currently appear in North Africa. This may suggest an ancient origin of the nomads in the more northerly mountain massifs of the Central Sahara (Dupuy 1999). According to our own anthropological examination (data not shown), the non-sub-Saharan haplogroups are not carried by "West Eurasian-like" individuals, as might be anticipated, but were rather detected in common "Fulani type" peoples.

- Cerny et al. (2006), mtDNA of Fulani Nomads and Their Genetic Relationships to Neighboring Sedentary Populations

No doubt, the excerpt highlights the totally fallacious and highly idealistic nature of the faulty logic pushed forward by extremist ideologue cliques, in trying to attribute intra-regional west African diversity to gene flow from outside that region.

The "true" western African is not supposed to sport variation, according to radical Eurocentric extremist thinking. On the other hand, radical self-proclaimed "Afrocentric" elements feed off on this Eurocentric viewpoint of the "special" or supposed "outlier" orientation of the Fula, which no doubt also influences their all-too-readiness to preemptively claim primary descendancy from the Fula; you see, sections of the Black American community have long been told, or else given the impression, that they are of the "true Negro" archetype and naturally, that they descend from the so-called "true Negro" western African "types". At the same time, word is that the "caucasoid" archetype entails "fine features". What then does this mean? If the "caucasoid" archetype embodies "fine features", then the so-called "true Negro" archetype must be its opposite counterpart, i.e. "undesirable features". This causes "reaction" in elements of Black American communities who are highly sensitive to such perceptions, which have implications about their own physical looks. Henceforth, such elements rush to claim primary descendancy from *mainly* such groups bestowed [by elements of Eurocentric or European academia] with the caricaturization of "fine features"; of course said ideological elements, when pressed on it — as noted, will vehemently deny the influence of said Eurocentric concepts on their own reactions. Likewise, Black American communities have long been told that they are not related to ancient Egyptians. This adds another element to the reaction just now mentioned, because not only are elements of the Black American community ever eager to be primarily associated with those groups "blessed" with said "fine features", but also passionate about showing their Eurocentric ideological counterparts that they also have direct ties to the ancient Egyptians. This is where the ideological "opposites" converge: Eurocentric elements look for "non-western African" origins to explain away the supposed "anomalous" facial features of certain western Africans, and the common choices of the area of such origins are either in the Nile Valley or "western Eurasia"; similarly, elements of Black American communities and like-minded "Afrocentrists" link origins of certain"honorary" western Africans [you know — said to have said "fine features"] with the Nile Valley, albeit for a different reason: to lay claim to said group as their primary "representative" ancestral group, and one which links them directly to the Nile Valley, hence in direct rebuttal to the Eurocentric notion of Black Americans being "true Negroes" and descendants of "true Negroes" almost exclusively from "western Africa". Please note the emphasized words (italicized), because they are central to what drives these ideological cliques. In relation to the point just noted, it's worth recalling this from the extract posted above:

According to our own anthropological examination (data not shown), the non-sub-Saharan haplogroups are not carried by "West Eurasian-like" individuals, as might be anticipated, but were rather detected in common "Fulani type" peoples. - Cerny et al. (2006), mtDNA of Fulani Nomads and Their Genetic Relationships to Neighboring Sedentary Populations

An example of one line of argument the latter ideological camp put forward, is:

According to Diop and a few Wolof people I know, they came to West Africa from the "East", even though they may have originated somewhere in the North/West present Sahara region long ago. Even many of their customs and culture is similar to that of Northern Sudanese. I've seen many times from Sudanese/horn Africans how they feel Hausa/Fulbe/Songhai music is so similar to their own, but with a different language.

Naturally, the line above is suffering from internal contradiction, featuring an ad hoc hypothesis stemming from an underlying intention to make the claim less arguable or vulnerable to refutation, but the implications are clear. It is worth noting that the specifics of said customs and culture remain elusive, notwithstanding being put forth as a central rationale, while the language affiliation of the Fula is seen as a non-issue. The language aspect of the Fula, is however, quite a central issue in any argument about their origins; it's what makes them, well, "Fula"
the name of their language phylum. Linguistic examination of this language places it in the "Atlantic" cluster or branch of the Niger-Congo super-phylum. This means that Fula clusters with "sibling" languages that belong in the said "Atlantic" cluster before it clusters with any group outside of this cluster, including other Niger-Congo sub-phylums. On a more macro level, the Atlantic branch being a sub-phylum of the Niger-Congo super-phylum means that Fula more closely shares "genetic" relationship with other Niger-Congo languages before it does any language group outside of the Niger-Congo phylum. The Niger-Congo branch is unquestionably most diverse in western Africa than it is in either central, eastern Africa or southern Africa. This means that Fula, as a Niger-Congo sub-phylum, more than likely evolved in the area where the Niger-Congo phylum underwent the most diversification from a common "proto-Niger-Congo" ancestor. The Niger-Congo phylum is essentially absent in northeastern Africa, and so, an isolated case like the Fula presence in Sudan presents itself as an "outlier" in a case study of language distribution for that general region. This means, in order for one to diminish the importance of Fula being a Niger-Congo language and the associated implication of its western African origins, one must argue that it doesn't belong in the Niger-Congo super-phylum, and that the "Atlantic" cluster as whole doesn't belong in this super-phylum, not to mention whether that cluster is even deemed to exist at all. To this end, recall that Mulcare et al. (2004) make note of research, that could not ignore the very close relationship between the various Fula dialects and that of Tukulor. It would be quite a neat trick to demonstrate that such relationship doesn't exist.

The following perhaps provides a good indication of the shaky premise of an effort to demonstrate very close, if not "sibling", familial relationship between Fula and Ancient Egyptian, merely through a highly idealistic selection and cherry-picking of terms that supposedly give the "appearance" of phonological similarities, as opposed to an in depth objective and materialistic examination of the language structures or broad lexical index of the languages under study:

...here's some linguistic data comparing Fulani /Mdu Ntr...

I - mi / ni
you - on / un
we - en / un
they - be / bu (people)
to be bad - bonude / boone
bad - boni / bon
death; to die - maayde;maayude / maati, moute, moout, mouti
last year - rawanen / snouf; ronpe
eye - yitere / eirti
blessing - barka / baraka
cow - nagge / naga
father - baaba / baba
...
the evidence is inexhaustible..

Here, all one needs to point out about the faulty logic behind the comparison, is to simply reference the variety and highly contrasting elements present in a single lexicon's set. For instance,...

Let's take say, "you" in Ancient Egyptian [language].

A number of words can mean "you" in Ancient Egyptian: e.g. tw (masculine, dep.), ntk (masculine, indep.), tn (feminine or plural, dep.), ntt (feminine, indep.) or nttn (plural, indep.); not to leave out affixed counterparts like
'-k' (masculine), '-t' (feminine), or '-tn' (plural).

The Ancient Egyptian word forms for "you" as just shown, are fairly well-known legitimate equivalents for the English "you", but are yet, markedly different from the "word" selected in that "Fulani-Ancient Egyptian" lexical comparison list. The word forms are also clearly structured in a definite grammatical format. This presents a linguistic situation that is more complex than that presented in the so-called "Fulani/Mdu ntr" list cited above.

As another example, let's take "I".

Here too, a number of derivatives come to attention:

'Wi/wy' (masculine, dep.), 'ink' (masculine, indep.), and affixed
'-i'.

Similarly, a Mdu Ntr term for "cow", as an obvious alternative to "naga" used the cited list above, is "[j]H.t", which itself is the de-neutered feminine version of
"[j]H", which is the equivalent of "ox".

Similar treatment can be applied to several of the other terms listed.

These examples serve to show that as a good rule of thumb, in any lexical comparisons between two distinct languages, designated words must be examined as to whether they are merely a derived form of a more basal term that is not subject to grammatical modification, whether there is more than one word for an idea or object and whether any such set of words were invented several times independently with respect to one another or whether they actually converge to a singular common ancestor, and in relation to this last point, in the event that one is dealing with word forms that were independently invented, raise the question as to which word form from a set with a variety represents the most basic manifestation as the denotation for an idea or name of an object.

Now of course, in lieu of the noticeably scarce evidence of linguistic basis for an eastern African origin for this group, i.e. the Fula (Fulani), ideologues of this idealistic hypothesis turn to certain cultural items that suggest "cultural affinity"; these are items like pectorals, artistic renditions that bespeak certain cultural respect for cattle or cow, gourd ornamentations, headrest designs, scepters and so forth, as discussed here: Examples of Cultural Similarities between those in the Nile Valley and those in other areas of Africa

These however, need not necessarily be suggestive of mass demic diffusion from the Nile Valley; rather, as hinted in that link, these may very well simply be relics of distant common cultural origins. One notable point of origin for such cultural convergence, is the Sahara; the once fertile Sahara has certainly proven to be one of the main areas of vibrant interaction between diverse autochthonous African populations in prehistory, that could have easily facilitated further cultural similarities between groups spread along its length. Cultural parallelism between western and eastern African, and possibly including southern African groups, can certainly be observable in just about any western African group used as a test subject, and hence, not something that the Fulani could be specialized or isolated for. If one were to read too much into this as the aforementioned ideologues do, then essentially entire western Africa would be envisioned as originating en mass from the Nile Valley
a highly questionable, if not unlikely, prospect.

The molecular genetic evidence side of things, closely follows that of language, as described above. All Fula samples undertaken are united by markers typical of "western African" ancestry. This means they are characterized by markers that diversified in western Africa.

mtDNA...

Despite the large size of the contemporary nomadic Fulani population (roughly 13 million people), the genetic diversity and degree of differentiation of Fulanis compared to other sub-Saharan populations remain unknown. We sampled four Fulani nomad populations (n = 186) in three countries of sub-Saharan Africa (Chad, Cameroon, and Burkina Faso) and analyzed sequences of the first hypervariable segment of the mitochondrial DNA. Most of the haplotypes belong to haplogroups of West African origin, such as L1b, L3b, L3d, L2b, L2c, and L2d (79.6% in total), which are all well represented in each of the four geographically separated samples. The haplogroups of Western Eurasian origin, such as J1b, U5, H, and V, were also detected but in rather low frequencies (8.1% in total).
- Cerny et al. (2006), mtDNA of Fulani Nomads and Their Genetic Relationships to Neighboring Sedentary Populations

Given what was readily available online on the study, including the mention of the 79.6% markers of West African origin, and the other 8.1% suggestive of coastal northwest African ancestry, it would appear that the remaining 12.3% of the sample were either "Pan-African" or "non-West African-specific" African markers. Not certain about this, because the actual tables of specific clusters of major markers were inaccessible at the time of examination, but if U6 was invoked in the markers of "Western Eurasian" origin, as the authors seem to have done elsewhere in their report, then that 8.1% contribution would actually be trimmed down further, considering that U6 is actually an autochthonous African marker, as discussed here: U6: A standalone clade?


As in African hunter-gatherers (Pygmies and Khoisan) and some populations from central Tunisia (Kesra and Zriba), three of the Fulani nomad samples do not reveal significant negative values of Fu's selective neutrality test. The multidimensional scaling of F^sub ST^ genetic distances of related sub-Saharan populations and the analysis of molecular variance (AMOVA) show clear and close relationships between all pairs of the four Fulani nomad samples, irrespective of their geographic origin. The only group of nomadic Fulani that manifests some similarities with geographically related agricultural populations (from Guinea-Bissau and Nigeria) comes from Tcheboua in northern Cameroon. -
Cerny et al. (2006), mtDNA of Fulani Nomads and Their Genetic Relationships to Neighboring Sedentary Populations

The message related here, goes back to the aforementioned "unifying" feature of Fula gene pools, primarily by definite markers of western African origin. Some may take it for granted that when the authors above state that "79.6%" of their Fula samples, taken together, constitute western African-specific markers, that this should simply be understood as, well, the obvious primary parental role of western African common recent ancestors to the contemporary Fula, while all else should be seen as "[foreign] admixture" to the autochthonous gene pool of the contemporary Fula, which as just mentioned, is west African in origin. However, as obvious as this is, there are individuals who are simply incapable of picking up on this straightforward logic, and instead, go onto profess ancestry elsewhere based on flimsy "admixture" elements
(supposedly "exotic" ancestry, vis-a-vis western Africa) invoked in the sample gene pool. Hence, one comes across ideologuesdepending on what ideological isle they belongwork hard at building shaky mythic origin theories around the small elements of any "admixture" component of the overall gene pool; for instance, one ideological side will swear that any fairly small occurrence of so-called "Eurasian" markers in the Fula sample indicates their ultimate "non-African", or more specifically, "western Eurasian" origin, while the other side will swear that "relic" markers, usually deemed to be persistent elements of distant or ancient common eastern African origin, indicates an east African ethnogenesis of the Fulain other words, the Fula are "in reality" or originally an autochthonous eastern African ethnic group instead of a western African one; their ethnogenesis as Fula or Fulani supposedly took place entirely in eastern Africa, but their present predominantly west African distribution is merely a function of their traditionally nomadic lifestyle, having recently migrated from east to west. The former ideologues for example, explain the predominant western African gene pool of the Fula, in terms of the Hamitic or "Hamitic-like" hypothesis, wherein the Fula, as originally a small "western Eurasian" group, undertook considerable genetic exchange with local western Africans, the supposed subjects of their final "back-migratory" destination. As for the latter ideologues, simply substitute the "western Eurasian" bit, and you get a similar situation, so that one gets, the Fula as "originally east African" in ethnogensis, considerably "admixed" with local west Africans on their westward migratory route along the Sahara. As the mtDNA examination shows [e.g. like the study cited above] however, such logic is just hogwash. All the "characteristic" markers of the Fula are of western African origin. To provide a demonstration for the message of this last entire passage, let's consider the following extract on the "admixture" test part of the aforementioned study:

Admixture Analysis. The question of the maternal origin of the Fulani was investigated using Admix 2.0. This software was developed to estimate the contribution of parental populations to the population under study. Data were entered according to group C from the AMOVA (i.e., the Mandenka were not considered; see Table 3); the mutation rate was 0.00005 per year. Admixture coefficients ([m.sub.Y]) were calculated from allele frequencies without taking into account the molecular divergence between the alleles. Because the exact period of Fulani migration cannot be estimated from archeological studies, we selected 4,000 years rather intuitively as the time to the possible admixture event. Bootstrap estimates of the coefficients and their standard deviations were calculated using 1,000 replicates. The results are summarized in Table 4 and show that none of the selected groups made a dominant contribution to the mitochondrial pool of the sampled Fulani. The only weakly prominent parental populations are from the Nile valley.
- Cerny et al. (2006), mtDNA of Fulani Nomads and Their Genetic Relationships to Neighboring Sedentary Populations

The piece above has for example, been used to argue the idea that the authors are concluding a "Nile Valley" origin of the Fula. This piece, for instance:
question of the maternal origin of the Fulani was investigated, is emphasized to state that this segment of the mtDNA analysis was the only section of the examination that deals with the question of Fula origins; the fact that the just-mentioned "79.6 %" component bespeaks just that, i.e. the maternal origin of the Fula, is carelessly thrown out the window. Meanwhile, the following piece: to estimate the contribution of parental populations to the population under study, is used in tandem with the aforementioned one, to add an exclamation point to the skewed idea about the authors' supposed conclusion that places Nile Valley populations as "the" parental population of the Fula. Naturally, the advocates of this obviously skewed/distorted interpretation of the study in question did not avail themselves the opportunity to fully examine either the study in its entirety or the specific extract in question, and thereof, understand them. For instance, common sense intimates that "any" genetic contribution to Fula gene pool, as the case is for any biological group, whether it happens to be one of post-ethnogenesis of the Fula as the well-defined ethnic group they are, or prior, has to be "parental"; hence, even the so-called little elements of "western Eurasian origin" [i.e. markers], are "parental" contributions. After all, any and all genes can only be transmitted from a "parent(s)". This however, should not be interpreted as "THE" parental component, i.e. the "proto-Fulani". The extract above is certainly not saying or implying that Nile Valley populations are "THE" parental populations of the Fula, but that is nonetheless how the extract is used in the wrong hands. In fact, the piece says, "The results are summarized in Table 4 and show that none of the selected groups made a dominant contribution to the mitochondrial pool of the sampled Fulani. The only weakly prominent parental populations are from the Nile valley, saying the populations that figure relatively 'prominently' in a parental role, do so "weakly".

When pressed to explain how populations that are supposed to be "THE" parental "proto-Fulani" population of the Fula only "weakly"—rather than strongly—figure in said potential "parental" role, the rationale put forward as an answer, is that the said ancestry is so "ancient", that the "parental" link became "weak" with time. Meanwhile, it is worth noting that while the extract claims that none of the selected groups made a dominant contribution to the mitochondrial pool of the sampled Fulani, it is NOT saying that said "selected groups" did not make any contribution at all. This is where the term "weakly" comes into the picture, because what it is implying, is that even though the said "selected groups" may well have contributed to the Fula gene pool, relatively speaking, it just didn't appear to be at the magnitude as that suggested by the "allele" sharing observed between the said Nile Valley samples and the Fula samples. By the same token, neither did the Nile Valley samples show any significant degree of "parental" role, but one of fairly "weak" invocation at that. As one reads further on in the study at hand, one learns just how tenuous this link is, as evidenced by the genetic distance or F^ST analysis. Contrary to the aforementioned rationale put forward, 'time frame' is not being offered by the extract as the primary responsible factor for this observation. In fact, the extract notes, recap: Admixture coefficients ([m.sub.Y]) were calculated from allele frequencies without taking into account the molecular divergence between the alleles. What does this mean? Well, if time frame were offered as a rationale for the said "weak" invocation of parental role of the Nile Valley groups, then one would have to attribute such to "molecular" divergences accumulated over time, which as noted, was not considered in the "admixture" test, which in turn actually only detected select "allele" sharing between test samples. Furthermore, because the "molecular divergence" was not considered, the test was not able to detect or account for the more immediate familial ties between distinctive alleles under test, which would have otherwise shown closer relationship between the Fula and other western African sample groups, with whom they share predominantly typical west African ancestry, before they did with groups outside western Africa. For example, if two different alleles linked to the same common recent ancestor were only a single nucleotide away from one another, the "admixture" test here would not have detected it; the "admixture" test only takes into account the "shared" alleles and their incidence (frequency) in respective samples. Hence forth, even though the Fula maternal gene pool was predominantly or primarily western African, as meted out by the results of the study itself, the direct "parental" role of other western African groups to the Fula could not be discerned from the "admixture" test, because the typical west African markers that characterized the Fula samples belonged to clusters that were "internally" [from a within-cluster molecular level] distinct from those of many of the aforementioned western African groups, again even though they shared the same "west African" common recent ancestry (like in the 79.6% component mentioned).

This brings us to the F^ST distance test mentioned in the last passage:

In the introduction we mentioned several hypotheses for the origin of the (nomadic) Fulani. One well-known hypothesis is that the Fulani come from the Nile valley (e.g., Tauxier 1937). Analysis of F^sub ST^ distances, however, shows no close relationship between the sampled Fulani and the analyzed Nilotic populations. Admixture analysis, however, does not exclude the possible parental role of the Nilotic populations because the admixture coefficient for these populations is high. It is necessary to state that the conclusiveness of this finding is rather low. Further geographic sampling, particularly from Niger and other parts of the Sudanic belt of Africa, is needed to acquire a deeper insight into the genetic structure of the nomadic people of the African Sahel. - Cerny et al. (2006), mtDNA of Fulani Nomads and Their Genetic Relationships to Neighboring Sedentary Populations

The aforementioned notion, that the previous extract was supposedly the only section of the study that examined the "origin" of the Fula, is further falsified by this extract of the study. In fact, this is the passage that directly confronts the "Nile Valley origin" hypothesis; to repeat:
In the introduction we mentioned several hypotheses for the origin of the (nomadic) Fulani. One well-known hypothesis is that the Fulani come from the Nile valley (e.g., Tauxier 1937), concluding that the F^ST distancesthe focus of the present passage of this blog topic does not support such a hypothesis: Analysis of F^sub ST^ distances, however, shows no close relationship between the sampled Fulani and the analyzed Nilotic populations.

The study follows this line with reminding the reader that the "admixture" test section of their study, however, "
does not exclude the possible parental role of the Nilotic populations because the admixture coefficient for these populations is high", which recalling from above, did not rule out a very tenuous possibility of a "parental" role by certain Nile Valley groups, which were subjects of the study. This seeming contradiction between the F^ST distance and the "admixture" test is worth examining. The F^ST distance measures the genetic distances in pair-wise comparisons between the populations under study, taking into account both the variation between two populations and variation within respective test sample populations; this means the more two test populations are similar in their gene pool, the smaller the F^ST distance is likely to be. In this case, there is no close genetic distance between the Fula and the Nile Valley group, and indeed, the predominantly west African origin of the Fula should serve as the immediate telltale sign of this; whereas, those of the Nile Valley groups are likely to be dominated by typical eastern African markers. The "admixture" test on the other hand, did not rule out but also could not definitely prove immediate parental role of the Nile Valley groups; such a tenuous link is likely to be observed in just about any western African group as was the case for the nomadic Fula samples, because "relic" east African ancestry is scattered along the length of western Africa, and indeed elsewhere on the continent. The conventional understanding is that the major clades of the African L0, L1, L2, and L3 *ultimately* originated in eastern Africa, but that these clades dispersed to other areas of the continent, where they would eventually expand and undergo localized internal diversification, ushering in region-specific sub-branches. Hence, it is expected to see relics of distantthe keywords hereeastern African ancestry in the Fula, but then, this could just as apply to just about any other west African ethnic group. Deducing an east African "proto-Fula" from such retained "low-incidence" distant ancestry is therefore irrational, nor does it really specifically tie them to just the Nile Valley, but eastern Africa in general. See, from the study in question for example:

From the phylogenetic point of view the mtDNA sequences from sub-Saharan Africa have been classified into L-type haplogroups (Chen et al. 1995, 2000; Watson et al. 1996, 1997; Rando et al. 1998; Bandelt et al. 2001; Pereira et al. 2001; Torroni et al. 2001; Brehm et al. 2002). About 30 sub-Saharan L-type haplogroups have been identified, and their ethnic or geographic origins and coalescence times have recently been summarized (Salas et al. 2002, 2004; Kivisild et al. 2004; Rosa et al. 2004). It seems that the main diversifications originated in East Africa but that the West African regions also contributed to the recent, wide mtDNA diversity
- Cerny et al. (2006), mtDNA of Fulani Nomads and Their Genetic Relationships to Neighboring Sedentary Populations

This
piece is simply repeating the oft said line about L-type mtDNA haplogroups likely having originated in eastern Africa, which is supposed to apply for *all* western Africans and indeed the rest of humanity, but that the ensuing "diversification" of haplotypes, like the ones the Fula are primarily known for, are of western African origin. Additionally, this west African diversification forms part of the "wide mtDNA diversity" involving L-type haplogroups. As funny as it may seem, this extract itself has erroneously been interpreted by certain individuals to mean that Fula lineages, specifically, are of "mainly east African origin" and that west African regions only recently contributed to this gene pool. The fact that the 79.6% figure of west-African specific [i.e. "sub-Saharan"] component had already been cited by the study, apparently failed to serve as a dead give-away to these individuals. Speaking of which, the study lays out the major components as follows:
  • The most numerous haplogroups are L3b (determined by the motif 16124, 16223, 16278, and 16362) and L3d (determined by the motif 16124 and 16223); these two haplogroups could not be distinguished from each other when only the HVS-I sequence was available. However, because both share the same geographic origin in West Africa, the HVS-II motif and RFLP polymorphisms were not further studied for a more specific molecular determination. Together, the L3b and L3d haplogroups contain 62 sequences with 12 haplotypes.
  • The second well-diversified haplogroup found in the Fulani nomads sample is L1b (50 sequences, 10 haplotypes), with the determining HVS-I motif 16126, 16187, 16189, 16223, 16264, 16270, 16278, and 16311, the origin of which also lies in West Africa (Salas et al. 2002, 2004).
  • All the other haplogroups identified are represented by conspicuously lower numbers; the most numerous among them, comparatively speaking, are L2b and L2c, which are also of West African origin.
- Cerny et al. (2006), mtDNA of Fulani Nomads and Their Genetic Relationships to Neighboring Sedentary Populations

One would think that with such a straightforward layout of the major components of Fula maternal gene pool, that there would be very little room for misinterpretation; alas, that hasn't stopped certain folks from confusing themselves over this simple information. It says right there, in plain English, that each of the said major components are of western African origin.

To give a brief synopsis of the message of the study at hand, it should be reiterated right off the bat, that the study has reconfirmed what other studies have found, and addition to certain "sedentary vs. nomadic" gene pool differentiations between Fula communities:

  • The Fula maternal gene pool is primarily or predominantly western African in origin. It therefore logically follows, that the main diversification of the Fula maternal gene pool originated in western Africa.
  • Even though these are typical west African-specific markers, the internal variation of their "clusters" suggest that the nomadic sections of the Fula, in contrast to sedentary Fula, generally have not gained much contribution from surrounding sedentary west African groups. What does this then mean?
It means that the Fula must certainly have an autochthonous western African origin, since not onlyas noted aboveare their characteristic maternal markers predominantly or primarily west African-specific, but also the internal variations of the clusters of these markers show no indication of any dominant genetic contribution from the Fula's neighboring or surrounding sedentary western African groups, except for that isolated case of the Mandinka [a sedentary western African group] which shows close matches with the nomadic Fula samples, thereby urging the authors to leave the Mandinka out of other case studies, like the "admixture" test.

Recalling...

On the other hand, the sampled Fulani groups differ from all other neighboring settled populations, including the Fulbe (Fulani) reported by Watson et al. (1997); the only exception to this is the Fulani sample from Tcheboua (Cameroon), which is not differentiated from three populations of the Central Sahel (the Hausa, Fulbe, and Yoruba).
- Cerny et al. (2006), mtDNA of Fulani Nomads and Their Genetic Relationships to Neighboring Sedentary Populations

If the neighboring west African groups have not contributed much to the nomadic Fula gene pool, and yet, it is still predominantly west-African specific, this ought to tell any clear-headed person that the Fula must have an autochthonous west African origin, and underwent ethnogenesis which entailed a considerable degree of endogamous or within-ethnic group inbreeding, thereby contributing to accumulation of genetic variation that became largely internalized within that ethnic group. It is for this reason, the nomadic Fula show a good degree differentiation from other west African groups when subjected to the "admixture" test, even though like the latter, their gene pool is primarily west African-specific.

  • About 79.6%, not counting U6, are of west African specific markers; where does this leave the remainder, which is not in that 8.1% figure?
As noted earlier, those markers [some 12.3%] would have comprised "pan-African" markers, which west African groups generally or basically share with eastern African groups, or else markers acting as 'relics' of eastern African ancestry, which were historically picked up along the Saharan corridor. It should be reiterated that markers acting as relics of distant east African origin are scattered, if not fragmentary in some cases, along the length of western Africa, and throughout the rest of the continent.

As a little side note: The basic eastern African markers scattered across western Africa are likely to sport relatively lower internal variation than their western African-specific counterparts, simply because the latter seems to have undergone major expansions and accompanying internal variation that accumulated over time, thereby considerably altering the original basic structure of the markers that provided the basis for the initial expansion events. In a few words: west African-specific markers are those that underwent extensive localized diversification in western Africa, while those markers suggestive of most common recent ancestry in eastern Africa, in most cases, are acting as relics of the original markers that provided the basis for the ones that have come to characterize typical west African-specific ancestry. Using such relics of east African ancestry to conjure up some "recent east African origin" stories is irrational, because as just noted, such ancestry is scattered across western Africa and elsewhere on the continent, and could be observed in just about any west African ethnic group.

  • Throughout the passages above, quick references to differences between nomadic and sedentary Fula have been made, notwithstanding genetic unity between them, including the level of genetic contribution from non-Fula groups neighboring the Fula, wherein nomadic Fula seemed to have received much less gene flow from neighboring sedentary populations than the sedentary Fula; well, here's another example of differentiation between the nomadic groups and the sedentary ones:
Some differences of the maternal gene pool between the nomadic and sedentary Fulani populations are also apparent from research conducted in physiological characteristics, for example, lactase persistence; the incidence of this trait among sedentary Fulani is far lower than among their nomadic counterparts (Holden and Mace 2002). - Cerny et al. (2006), mtDNA of Fulani Nomads and Their Genetic Relationships to Neighboring Sedentary Populations
  • The authors of the study above make a quick but notable reference to Y-DNA marker examinations of the past:
Y-chromosome data of 22 African populations, including the Fulani from Burkina Faso and northern Cameroon, were analyzed by Cruciani et al. (2002). The main result of Cruciani's study is that different populations from northern Cameroon (Fali, Ouldeme, Daba, and some mixed samples) reveal traces of backmigration from Asia to Africa because of a high proportion of haplotype 117. However, the Fulani sample from northern Cameroon considered by Cruciani and colleagues shows a rather low frequency of this haplotype, and the Fulani, which have a high frequency of haplotype 43, are situated as outliers. Cruciani et al. (2002) also showed that the Fulani from Burkina Faso have reduced diversity, because only two Y-chromosome haplotypes were observed in their sample. - Cerny et al. (2006), mtDNA of Fulani Nomads and Their Genetic Relationships to Neighboring Sedentary Populations

It is worth noting that haplotype 117 is the same marker that was the subject of R1*-M173 chromosomes in northern Cameroon earlier on this site; see:
R1*-M173 in Africa [a topic that is regularly updated]. Haplotype 43 corresponds to E1-M33 chromosomes, a marker that seems to figure regularly in Fula samples. Indeed, Cruciani et al.'s (2002) northern Cameroonian Fula sample featured E1-M33 as the predominant marker, and while haplotype 117 (R1*-M173) occur in much lower frequencies than this marker, the latter were still detected in noticeable frequencies. It is important to note that R1*-M173 incidences was not limited to the Fula sample, but was featured in other northern Cameroonian groups, most of them non-Bantu speaking groups. Naturally, this means that the Fula picked R1*-M173 markers up along their west-to-central African migratory path or via the Chadic region; it wasn't an autochthonous Fula marker as the case seems to be in terms of E1-M33, E3a and possibly E2-M75. However, it appears that northern Cameroonian Fula groups did not stay put in the vicinity of central Africa and/or Chad, but that they headed further eastward, and wound up in Sudan. Indicators of this migratory trajectory is evidenced from the fact that the Sudanese Fula predominantly featured R1*-M173 [Hassan et al. 2008], the markers first implicated in Cruciani et al.'s (2002) Cameroonian Fula sample, and yet still retained their E1-M33 ancestry, which is considered to be a typical west African-specific paternal marker.
  • Despite this predominant R1* representation, there's something still unmistakable about the Sudanese Fula sample: they retain their western African ancestry, as indicated per E1-M33, which was absent in all autochthonous Sudanese groups, save for other western African groups now in Sudan like the Fula [see the authors' references to the Senegalese Wolof, and their Sudanese Hausa sample, for example].
The elevated frequency of R1*-M173 in the Sudanese Fula implies that they drew from just a section of a pre-existing west African [starting from Cameroon] Fula population that had considerable frequencies, though not predominant, of R1*. This would ensure that the original diversity pattern attained in the parental Fula group would have been lost in the divergent nomadic Fula group that headed eastward. Thus, if Hassan et al.'s (2008) Fula sample are any indication, then it is reasonable to assume that random genetic drift might well have elevated the R1* component of the male gene pool, while it worked the other way on the west African-specific markers like E1-M33, which the Sudanese group still retains. An indicator of this reduced genetic diversity can be seen from the fact that northern Cameroonian Fulbe/Fula featured Hg A3b2, Hg E1-M33, Hg K2 and Hg R1*-M173, while the Sudanese group only featured Hg E-M33, Hg R1*-M173, and Hg E3b1. As one can see, putting Hg E3b1 aside, the Sudanese Fula gene pool appears to carry diversity which is a subset of that of their northern Cameroonian brethren. It is easy to understand how Hg E3b1 figured in the Sudanese Fula, while Cruciani et al.'s (2002) Fula sample showed no signs of it; the Sudanese Fula naturally picked up meaningful Hg E3b1 ancestry from neighboring autochthonous Sudanese ethnic groups, likely fellow nomadic pastoralists characterized by these markers. Hg E3b1 after all, appears to be a prominent marker in autochthonous Sudanese gene pool. All that side, the Sudanese Fula bear R1*-M173 and E1-M33 markers just as their northern Cameroonian Brethren do. This is what one would expect, if they derived from a segment of northern Cameroonian Fula or like group in the vicinity of that area as a subset of that segment, thereby only featuring only a portion of the diversity of their parental population.

It's insightful to note that Cruciani et al.'s (2002) Burkina Faso Fula sample too featured noticeable frequencies of Hg E1-M33; this marker appears to be a regular occurrence in Fulbe samples, regardless of how divergent they may be with respect to one another; here though, it assumes the role of the next dominant marker, after Hg E3a, which appears to generally be predominantly represented in many western African Fula samples, further west of Cameroon. Rosa et al.'s (2007) Fulbe/Fula sample too were predominantly featured by E3a markers (~75%), and yes, it too was noticeably featured by E1-M33 (~ 7%). As Rosa et al.'s (2007) sample reaffirms, E3b markers are scattered across the western Sahel and "Sub-Saharan" areas in low incidences, and it appears to be performing just that function in the Guinean [Bissau] sample, wherein *all* the groups represented were tested postive for the marker, usually in low but noticeable incidences. In the case of the Guinea Bissau Fula, overall frequency of E3b-M35 comprised about 12% of the sample, the bulk of which was E3b1/E-M78 (~10%), with the remainder being the paraphyletic E3b*. Indeed, as Cerny et al. (2006) note, Cruciani et al.'s (2002) Burkina Faso Fulani sample was characterized by a relatively reduced paternal lineage diversity, with only Hgs E1-M33 and E3a being featured therein. By contrast, we see greater diverstiy in the Guinean Fula sample, featuring Hgs E1-M33, E2-M75 (~1.7%), E3a, E3b, A1 (~ 1.7%) and R1b (~1.7%). The nomadic tradition of the Fula apparently comes across in the Fula samples taken from different areas; recurring markers like the aforementioned E1 suggest continuity across the Fula spectrum, but whereas the northern Cameroonian Fula featured Hg A3b2, the Guinean sample featured the Hg A1 derivative, the most common A-M91 marker in western Africa. Hg A3b2 was featured in the Guinean sample, but in the Balanta sample. Likewise the Cameroonian Fula sample featured Hg K2 and the paraphyletic Hg R1*, which were not represented in the Guinean Fulani sample, but as just shown, a derivative of these markers, R1b, managed to get into the Fula gene pool, but in noticeably low frequency (~1.7%). The different patterns observed here, again appear to bespeak the traditionally nomadic pastoralist lifestyle of the Fula, wherein a portion of the gene pool in the areas they spend any meaningful time, they managed to pick up a piece of the gene pools therein. The other differences, as already noted in earlier passages, are likely a manifestation of whether a Fula group is sedentary or nomadic. This notwithstanding, it's worth reiterating that continuity is observed in the paternal gene pool. The telltale sign of this unifying feature unequivocally re-affirms their west African origin and ethnogenesis in this region as an ethnic group. Henceforth, it is no accident that dissidents of this fact, are unable to account for why, even the Sudanese Fula, the *only* attestable eastern African community of the Fula [pending demonstration of tangible material otherwise] , retain relics of their western African ancestry in the form of E1-M33 markers, while this marker appears to be absent in the traditional autochthonous ethnic groups of Sudan and northeastern Africa in general. They surely must have been in western Africa at some point in their bio-history, so as to have picked up ancestry typical of western African areas. Furthermore, the fact that the Sudanese Fula retain their "Atlantic" language subphylum of the Niger-Congo phylum is glossed over by these same ideological dissidents, even as they fail to account for the fact, and even as this cultural trait again bespeaks their ultimate western African origin. Little wonder a question such as this, "Can I get an *elaborate* grammatical and syntax correspondence between Fula and Ancient Egyptian?, is greeted with graveyard silence.

Even simple visual aids such as this, demonstrating the obvious "outlier" nature of Fula language presence in Sudan, the only eastern African territory in all of eastern Africa implicated in its distribution, proves to be quite challenging for the grasping capacity of ideologues mentioned time and again over the course of this topic:
insertmapu800.gif
The last several passages discussed above will, and should, help any perspective reader answer the following questions, as it pertains to Cerny et al.'s (2006) examined extract:

The question of the maternal origin of the Fulani was investigated using Admix 2.0. This software was developed to estimate the contribution of parental populations to the population under study. Data were entered according to group C from the AMOVA (i.e., the Mandenka were not considered; see Table 3); the mutation rate was 0.00005 per year. Admixture coefficients ([m.sub.Y]) were calculated from allele frequencies without taking into account the molecular divergence between the alleles. Because the exact period of Fulani migration cannot be estimated from archeological studies , we selected 4,000 years rather intuitively as the time to the possible admixture event. Bootstrap estimates of the coefficients and their standard deviations were calculated using 1,000 replicates. The results are summarized in Table 4 and show that none of the selected groups made a dominant contribution to the mitochondrial pool of the sampled Fulani. The only weakly prominent parental populations are from the Nile valley . - Cerny et al. (2006), mtDNA of Fulani Nomads and Their Genetic Relationships to Neighboring Sedentary Populations

Why despite the Fula lineages being overwhelmingly west African in origin, as the authors' results cite, do they say...

1)"none of the selected groups made a dominant contribution to the mitochondrial pool of the sampled Fulani"?

2)The only weakly prominent parental populations are from the Nile valley ?

3)Why..."the Mandenka were not considered"?

4)And what is the implication of this:

"Admixture coefficients ([m.sub.Y]) were calculated from allele frequencies without taking into account the molecular divergence between the alleles."

As typical as the Fula are, i.e. genetically and culturally, in their west African character, if some other western African group were the subject of discussion and had these very same traits, there would be no question about it, that NO controversy whatsoever would so much as be generated about them; to put it simply, the evidence is overwhelmingly one-sided: the west African origin!
______________________________________________________________

Miscellaneous notes:

This pertains to excerpts taken from: The Genetic Structure and History of Africans and African Americans

Sampling entailed...

We studied 121 African populations, four African American populations, and 60 non-African populations for patterns of variation at 1327 nuclear microsatellite and insertion/deletion markers. We identified 14 ancestral population clusters in Africa that correlate with self-described ethnicity and shared cultural and/or linguistic properties.

With noticeable mention of...Fulani (Nigeria and Cameroon), the Baggara Arabs (Cameroon), the Koma (Nigeria), and Beja (Sudan)

Moving onto excerpts of interest...

Mitochondrial DNA analysis indicates that Fulani have lineages of predominantly West African origin and that they cluster together and close to the Mandenka population from Senegal [S93].

By contrast, Y chromosome analyses of Fulani
sampled in the Sudan indicates shared ancestry with Nilo-Saharan and Afro- Asiatic speaking populations [S89].

The line about the mtDNA is essentially a reversion back to what has been cited in the main post above, as observed by Cerny et al. (2006). The line about the Y chromosome analysis however, is quite misleading; one only needs to refer to Hassan et al. (2008) [see: Y-Chromosome Variation Among Sudanese] to see how: The Sudanese Fula sample here, as noted in the main post above, predominantly featured Hg R1*-M173 markers in the uniparental paternal gene pool. This marker is in fact generally quite rare in "Afro-Asiatic" speaking populations; rather, its highest incidences has thus far been reported in Niger-Congo, Nilo-Saharan and "Afro-Asiatic" (Chadic) speaking groups in northern Cameroon, with smaller occurrences in the rest of central Africa [see Luis et al. 2004]. Nilo-Saharan groups in Sudan itself, it should be noted, did NOT share this ancestry with the Sudanese-based Fulbe at all. This reverts back to what had already put forward in the main post, about the implication being that these Sudanese Fula derived from a section of the northern Cameroonian Fula, wherein the Hg R1*-M173 is featured in a considerable but not the predominant frequency.

The only marker that the Sudanese Fula shared with Nilo-Saharan and Afrasan ("Afro-Asiatic") speaking Sudanese groups was Hg E3b1 (M78). Common sense however, intimates that this aspect of the paternal gene pool was largely picked up from neighboring populations, only after the Fula situated themselves in Sudan. It makes sense, since after all, E3b1 is one of the prominently featured markers found in the Sudanese sample.

To recap, there's something still unmistakable about the Sudanese Fula sample: they retain western African ancestry, as signified by Hg E1-M33, which was absent in all autochthonous Sudanese groups; the only other groups where this marker was implicated in Hassan et al.'s (2008) work, were, well, the well-established west African groups like the Hausa and Wolof. When pressed on to explain away this anomalous feature of the Sudanese Fula's paternal gene pool, dissidents of the west African origin noticeably become mute on the matter.

These results raise the possibility of differential patterns of male and female gene flow into this population.

Whatever may be "differential" about male and female gene flow patterns in Fula sample, no doubt influenced by matters like life style - recalling on nomadic vs sedentary, both the maternal and paternal gene pools communicate the same thing: west African origin!

Our analysis, using genome-wide nuclear markers and STRUCTURE, indicates that the Fulani have distinctive ancestry [fuchsia] at K = 14 in the global analysis [Figs. 3,4] and at K = 9 -14 in the Africa analysis [Fig. S13]. The Fulani cluster with the Chadic and Central Sudanic speaking populations at K <13> They also cluster near the Chadic and Central Sudanic speaking populations in the NJ tree based on population genetic distances [Fig. 1].

Fula samples taken from either central Africa or Sudan are expected to cluster with groups in central Africa-Chad, as the Fula would have arrived in Sudan via a central African and/or Chadic corridor, having experienced a bottleneck [see main blog post]. The central African Fula in turn would have arrived ultimately from Atlantic-coastal regions of western Africa, where their ethnogenesis as the Fula-"proper" likely occurred.

The pattern of the Sudanese Fula's Y-DNA composition is for instance, undoubtedly different from their western African brethren, but continuity still exists via recurring typical western African markers that unite all Fula groups, regardless of where they spread to; e.g. primarily Hgs E1 and E3a, and a lesser extent, Hg E2. Either of these clades are telltale signs of so-and-so Fula groups' western African ancestry.


In the global STRUCTURE analysis, the Fulani show low to moderate levels of European/Middle Eastern ancestry [blue], consistent with mtDNA and Y chromosome [S89] analyses, as well as the presence at low frequency of the -13910 mutation associated with lactose tolerance in Europeans in this population [S94].

Again quite misleading. Yes, understandably, "low" levels of shared ancestry attributable to potential "European" ancestry, might be correlated with "low level" incidences (~1.7 %) of R1b [pertaining the Rosa et al.'s (2007) Guinean sample] or relics of this ancestry attained from coastal northwestern populations [Imazighen] seen on the mtDNA side [Cerny et al. 2006], as cited in the main post. However, in the case of Fula samples, from northern Cameroon through to Sudan, a good degree of presumed shared "European/Middle Eastern ancestry [blue]" may very well actually be linked to common "distant" ancestry, as the paraphyletic R*-M207 and R1*-M173 markers suggest; these latter two are essentially rare to absent in "Middle Eastern" and "European" populations, which are mainly characterized by more downstream markers, especially European populations, who have no upstream markers for R1. Now, because R1b and R1a markers are still 'molecularly' linked to the upstream R*, any group bearing the latter or else any intemediary nodes between the upstream R* and the downstream R1a and R1b will "appear" to share ancestry with Europeans and/or possibly "Middle Easterners", which could misleadingly be misread as ancestry from "European/Middle East". Given the features of northern Cameroonian and Sudanese Fula sample uniparental male gene pool, it is a safe bet that this is the underlying issue at work, especially when the red flag of "moderate" levels is evoked; the Fula uniparental gene pool specifics , whether maternal or paternal, simply do not bear out "moderate" ancestry from "Middle East" or "Europe".

Recalling a previous blog post here
, R1*-M173 in Africa, we have:

Interestingly, having re-visited Wood et al. (2005), it should be pointed out that paraphyletic clade of R*-M207 was detected amongst some "Afro-Asiatic" African groups, along with the paraphyletic clade R1*-M173 [it is worth noting that authors implicate the Egyptian sample here as something other than that of Semitic speakers (Arabic)], while some Niger-Congo groups though in small frequencies [pooled] — tested positive for the paraphyletic R1b*, lacking the established downstream R1b markers. Henceforth, R*-M207, lacking downstream mutations have been identified in African groups via this study; and yes, the basic nodes of all Hg R's downstream clades had been accounted for, which means that R*, as predicted above, is NOT relegated to the Indian sub-continent. All in all, this suggests that African Hg R pool is more diverse than many seem to think.

As for the matter concerning lactose tolerance, it has been pointed out here before, how this has nothing to do with European ancestry. Recap, see:
More on R1*-M173 bearers

In summation, the piece essentially rehashes the well-understood facts of primary west African ancestry of the Fula, both in terms of genealogy and language, along with genealogical "admixture" elements picked up over the course of migratory events, but suffers from matters, as laid out in detail in the main blog post, pertaining to idealistic urges for mystification where none is necessary or warranted.

*Watch for updates.

A spin-off post of this topic has recently been posted here: Reviewing: The Genetic Structure and History of Africans and African Americans
_____________________________________________________________

Miscellaneous reading...

Perceptions of Sudanese Fulani inside Sudan itself may well be instructive, particularly to those who might harbor preconceived notions about local perceptions without having set foot in the region; to aid in this, here a few extracts taken from the net:

November 1, 2008 — Omer al-Bashir, the embattled president of Sudan, has been reported to have made a statement in a Sudanese newspaper al-Ayaam two months ago that the Hausa Fulani people are not Sudanese and that they should not vote in the 2009 general elections stipulated by the Comprehensive Peace Agreement (CPA)! This sparked off a vigorous protest in the Hausa Fulani communities all over the Sudan especially in the city of al-Gadhariff in Eastern Sudan. During the protest, in which people from the Hausa neighborhood organized a peaceful march led by a group of tribal leaders and officials in a popular demonstration to hand a memorandum of protest to the Head Quarter of the Gadhariff state to express their disapproval of President al-Bashir’s reported offensive statement that the Sudanese tribes of Hausa, Fulani, Bornu and Tama groups as non-Sudanese and they not have the right to participate in the upcoming elections in Sudan. Government of Sudan forces backed by heavy weapons attacked the unarmed civilians and chased them into their neighborhoods using tear gas and live ammunition, which resulted in the martyrdom of four people who were killed in cold blood and hundreds were wounded and among the seriously injured were children and women and local hospital had difficulty coping with the casualties. Accordingly, the NCP regime added insult to injury. The irresponsible, imprudent and a racist statement by the president al-Bashir and the killing of peacefully demonstrating civilians are deplorable.

We all know that the Hausa Fulani people are hard working, devout Muslims of high moral code and patriotic Sudanese citizens who are widely distributed all over the country for centuries. They have contributed immensely to the development projects. It is unfair and an unjust accusation on the part of a president of a nation to label his fellow compatriots as aliens. Though it is appalling, but not surprising as al-Bashir is not a stranger to controversy with respect to random and misjudged statements of his personal views. He is renowned for the vulgarity of the language he tends to use without giving any attention to the consequences it may lead to. Al-Bashir’s blunders have tarnished the Sudanese reputation and the once good name of Sudan. Similar reckless distasteful obnoxious assertions were made by Abdel-Rahim Mohamed Hussein, Sudan’s Minister of Defence, who was reported to have stated in a press conference at the Sudanese Embassy in Riyadh, Saudi Arabia, on Thursday March the 13th 2008 at 11:45pm in an answer to the question whether it was true that the Government of Sudan (GOS) was planning to settle Five Million Egyptians mainly from Upper Egypt, the Nile valley south of Cairo, in the Northern Region ”Shamaliya” by saying: " If we assume the argument that the government seeks to resettle 5 million Egyptians in northern region why not? What is good of 8 million people in Darfur, all of whom are Africans came from abroad. Isn’t it better for you those who come from Egypt or those Africans who have come from West Africa?! "... - By Mahmoud A. Suleiman, The Hausa - Fulani are Genuine Sudanese, Mr. President!

Read further here

The following are from photo-journalist Vit Hassan in Sudan, who has also contributed to National Geographic...

Image 1
Woman from a Umm Bororo tribe wearing traditional clothing..
Umm Bororo are nomads with cattles by origin from Nigeria...

Taken in Eastern Sudan,close to Ethiopia.

Image 2
Dance of girls from a Umm Bororo tribe.(No cropping)

-This is for Globaltrotter-

Taken in Eastern Sudan,close to Ethiopia.

Image 3

Image 4
Woman from a Falata Uda tribe.
Falata are divided to 13 main branches:Uda,Voila,Djafun,Dooga,Nara,
Malle,Dindima,Adamawa,Danedji,Gamba,Sisilbi,Fuuta,Boodi..

They are nomads with cattles.
She wearing traditional colorful clothes of a Falata Uda tribe...
By way-"boodum" means in their language-'beautiful'...

Image 5
Another boy with a traditional hair style of a Umm Bororo tribe...

Image 6
2 girls from a Umm Bororo tribe coming back to home from market.
They keeping dishes with kind of white joghurt...

Taken in Eastern Sudan,close to Ethiopia.

[Note that the Sudanese Fula retain usage of large gourd pots or bowls, very common in western Africa, especially in areas near the Atlantic Ocean]

Image 7
Girl from a Falata Gamba tribe.
She has small tribal marks ("shluh" )in the shape of sun.This means-she is from Falata Gamba tribe..

Taken in Eastern Sudan,close to Ethiopia.

Image 8
Woman from a Falata Uda tribe wearing traditional colorful clothing of her tribe...
They are divided to 13 main branches..
Women of Falata 'Uda' are by their features close to some tribes in Chad..

Taken in Eastern Sudan.

Image 9
Girl from a Umm Bororo tribe.
They are nomads with cattles,by origin from countries in Western Africa.

They are divided to many main branches and every branch have a different marks("Shluh")on their faces..She has also unique black tatoo on her lips...

Taken in Eastern Sudan,close to Ethiopia.

Image 10
Boy from a Umm Bororo tribe.
They are by [b]origin from countries in Western Africa[/b].
Unmarried boy has a sign of his possition long hairs...

Taken in Eastern Sudan,close to Ethiopia.

Image 11
Girl from a Umm Bororo tribe

Image 12
Man from a Falata tribe selling "Hidjab"

...otherwise known as "jujus" in some quarters; again, a common sight in western Africa, where one comes across fortune "prescribers" [for lack of a better term in English; it is supposedly passed off as their 'trade, expertise, specialty or turf'] of certain "Quranic" lines enclosed in leather 'casings' [via sewing through leather].

Source: Photos and accompanying notes [in blue] by Vit Hassan - taken from here: Link

More seen here, another site: Link 2


Elsewhere, we have the following pages taken from Sudanese anthropologist Rogaia Mustafa Abusharaf's work - Female Circumcision 2007:

 -

 -

Source: Female Circumcision 2007 (click on the images for higher resolution)

If these extracts are any indicators, the perception concerning the Fulani's ultimate arrival from western Africa appears to be a widespread one even in Sudan itself, by both right-wing elements and liberal ones.