Berber Population Research Articles

Racial myth, colonial reform, and the invention of customary law in Morocco, 1912–1930

Throughout the period of French indirect rule in Morocco (1912–56), colonial scholars and administrators generated a mass of literature to explain the perceived differences between Arab and so-called Berber (Amazeeri) societies. In the minds of many colonial enthusiasts, objective ethnic difference translated into a series of clear social, cultural, and political traits that had divided the indigenous populations for centuries. Historians of the period have highlighted this ethnographic focus as one of the preeminent justifications for French reforms in Morocco. The reform and institutionalisation of customary Berber law, which provoked the most widespread urban protest in nearly two decades, has long been understood as an example of legislative overreaching: the vibrant popular reactions to the so-called Berber dahir of 1930 signalled to many contemporary actors and subsequent historians the veritable birth of the Moroccan nationalist movement, as urban populations protested the what they saw as the diminution of the sultan's jurisdiction and the attenuation of shari‘a in many rural areas. Implicit in this narrative is the idea that by 1930 the otherwise cautious and informed process of ethnically specific reform somehow went awry. This article seeks to demonstrate that the French legal reform program was, from its inception, founded upon vague, contested understandings of Berber social and political organisation that colonial administrators mobilised for very specific political ends. The so-called reform of Berber customary law was rather an invention of a system that French administrators used to remove the Berber population from the influence of the Arab makhzan, to marginalise religious law in favour of secular European values, and ultimately to fracture the network of social, cultural, and political relationships between the sultan and his people. Understanding the manipulative bases of legal reform sheds light on the relationship between the human sciences and colonial policy – specifically the complicated strength and malleability of scholarly mythmaking – and underscores the paradoxes of the French ‘civilising mission’.

Mitochondrial DNA and Y-chromosome microstructure in Tunisia

Mitochondrial DNA (mtDNA) and Y-chromosome variation has been studied in Bou Omrane and Bou Saâd, two Tunisian Berber populations. In spite of their close geographic proximity, genetic distances between them were high and significant with both uniparental markers. A global analysis, including all previously studied Tunisian samples, confirmed the existence of a high female and male population structure in this country. Analyses of molecular variance analysis evidenced that this differentiation was not attributable to ethnic differences. Mantel test showed that, in all cases, Y-chromosome haplotypic distances correlated poorly with geography, whereas after excluding the more isolated samples of Bou Omrane and Bou Saâd, the mtDNA pattern of variation is significantly correlated with geography. Congruently, the N(m) ratio of males versus females pointed to a significant excess of female migration rate across localities, which could be explained by patrilocality, a common marriage system in rural Tunisia. In addition, it has been observed that cultural isolation in rural communities promotes, by the effect of genetic drift, stronger loss of diversity and larger genetic differentiation levels than those observed in urban areas as deduced from comparisons of their respective mean genetic diversity and their respective mean genetic distances among populations. It is likely that the permanent exodus from rural to urban areas will have important repercussions in the future genetic structure of this country.

Endogenous Berber Forest Management and the Functional Shaping of Rural Forests in Southern Morocco: Implications for Shared Forest Management Options

On the basis of two case studies in rural Morocco, one in a mountainous area of the Central High Atlas and the other in the argan tree area of the southwest Atlantic coast, we show how local Berber populations have actively shaped their forest areas through endogenous management systems at different scales: 1) at the individual tree level by differential cutting or trimming which lead to specific conformations of the tree, 2) at the tree stand level, by determining the type, structure and level of resources, and 3) at the landscape level in which complementary patches of forest areas with particular functions are consciously organized within the overall territory. These practices are strongly linked with the overall socioeconomic organization of the local communities, and mix individual with common rights of access and uses. Forests are viewed as part of the domestic sphere of local livelihoods. Hence, they typically constitute what we refer to as rural or domestic forests since they integrate production and conservation with social, political and spiritual dimensions. These features are of importance for considering forester-local community relationships, and for developing alternative forest management policies.

The Berbers of the Arabs

Arab conquest of northwest Africa is a major event in the field ofNorth African Studies. For scholars, its most lasting effects were the Islamization and Arabization of the Berber populations. While historians often disagree about the chronology, character, and extent of these processes, they ail agree that they are essential to understanding the medieval period. In truth, the scholarly consensus is slightly more complicated than this more popular view. Specialists know that the category is of Ara bie origin and thus that the Arabs could not have conquered peoples called Barbar prior to the conquest.1 Furthermore, while Latin and Greek sources refer to some people in northwest Africa as (βάρβαροι and bar bari), the ideas associated with these wil be shown to be so dif ferent from the Arabie Barbar that it is difficult to confuse the two. Thus, in spite of the obvious linguistic similarity of the Greek, Latin, and Arabie terms, the Arabs did not apply the term Barbar to describe the exact same groups of peoples called barbarians in classical and late antique sources. So, who were the Barbar of the Arabs? Encapsulating a widely held view among specialists, Gabriel Camps explained, The Berbers of the Arabs are the Moors of the Romans.2 According to this understanding, instead of conquering the Berbers, the

Genetic differences among North African Berber and Arab-speaking populations revealed by Y-STR diversity

Y-chromosome STR polymorphisms are inherited in a haploid state which makes them a powerful tool for easy tracing of paternal lineage and for use in human population evolutionary studies. North-African Y chromosomal diversity has traditionally been studied in order to find genetic and geographic associations as well as to test how natural and cultural barriers have affected the degree of genetic flow not only within North Africa but also in a wider Mediterranean context.The degree of Berber/Arab genetic differentiation in the Moroccan population has been tested for a complete set of forensic markers as sixteen Y-chromosomal short tandem repeats (STRs) (DYS19, DYS385, DYS389I, DYS389II, DYS390, DYS391, DYS392, DYS393, DYS437, DYS438, DYS439, DYS448, DYS456, DYS458, DYS635 and GATA H4.1). The results suggest considerable population heterogeneity in North Africa.

Polymorphism Study of the Insulin Receptor Substrate IRS1 and IRS2 Genes Associated with Type 2 Diabetes in Ethnic Groups of Djerba Island

Aim: We aimed to evaluate the influence of the Gly972 Arg and Ala513Pro variants of the insulin receptor substrate1 gene (IRS1) and the Gly1057 Asp variant in IRS on type 2 diabetes mellitus (T2DM) in Arab and Berber men and women from Djerba Island, Tunisia. Methods: Genotypes, allelic and genotypic frequencies were studied. The amplified products were analyzed by Restriction Fragment Length Polymorphism (RFLP) methods by comparing T2DM with healthy controls from the same ethnicity. Results: No differences in genotype or allelic frequencies were found between T2DM and healthy controls in either Arab or Berber ethnic groups. Conclusion: The Gly972 Arg and Ala513Pro variants in IRS1 and Gly1057 Asp in IRS2 polymorphisms are not associated with T2DM in the Arab or Berber populations of Djerba Island

Ancient Local Evolution of African mtDNA Haplogroups in Tunisian Berber Populations

Our objective is to highlight the age of sub-Saharan gene flows in North Africa and particularly in Tunisia. Therefore we analyzed in a broad phylogeographic context sub-Saharan mtDNA haplogroups of Tunisian Berber populations considered representative of ancient settlement. More than 2,000 sequences were collected from the literature, and networks were constructed. The results show that the most ancient haplogroup is L3*, which would have been introduced to North Africa from eastern sub-Saharan populations around 20,000 years ago. Our results also point to a less ancient western sub-Saharan gene flow to Tunisia, including haplogroups L2a and L3b. This conclusion points to an ancient African gene flow to Tunisia before 20,000 BP. These findings parallel the more recent findings of both archaeology and linguistics on the prehistory of Africa. The present work suggests that sub-Saharan contributions to North Africa have experienced several complex population processes after the occupation of the region by anatomically modern humans. Our results reveal that Berber speakers have a foundational biogeographic root in Africa and that deep African lineages have continued to evolve in supra-Saharan Africa.

Ancient Local Evolution of African mtDNA Haplogroups in Tunisian Berber Populations

Our objective is to highlight the age of sub-Saharan gene flows in North Africa and particularly in Tunisia. Therefore we analyzed in a broad phylogeographic context sub-Saharan mtDNA haplogroups of Tunisian Berber populations considered representative of ancient settlement. More than 2,000 sequences were collected from the literature, and networks were constructed. The results show that the most ancient haplogroup is L3*, which would have been introduced to North Africa from eastern sub-Saharan populations around 20,000 years ago. Our results also point to a less ancient western sub-Saharan gene flow to Tunisia, including haplogroups L2a and L3b. This conclusion points to an ancient African gene flow to Tunisia before 20,000 BP. These findings parallel the more recent findings of both archaeology and linguistics on the prehistory of Africa. The present work suggests that sub-Saharan contributions to North Africa have experienced several complex population processes after the occupation of the region by anatomically modern humans. Our results reveal that Berber speakers have a foundational biogeographic root in Africa and that deep African lineages have continued to evolve in supra-Saharan Africa.

Assessing human genetic diversity in Tunisian Berber populations by Alu insertion polymorphisms

Background: The most recent Alu insertions reveal different series of characteristics such as stability that make them particularly suitable genetic markers for human biological studies.Aim: Six human-specific Alu insertion polymorphisms were typed in two Tunisian Berber populations with the aim of analysing the genetic diversity of these two communities and the genetic relationships between this region of North Africa and other populations.Subjects and methods: Forty-seven Berbers from Sejnane and 33 from Takrouna were sampled. Alu insertion polymorphism was analysed using PCR with loci specific primers.Results: A similar level of gene diversity was detected in Sejnane and Takrouna populations. PC results revealed genetic affinities between these two populations and some Eurasian populations (Germany, Genova and Syria). In contrast, there is a differentiation between these two Berber communities and North African and Iberian populations.Conclusion: The results of this study confirm the heterogeneity of Berbers in North Africa, which suggests their diverse origins. In the case of Sejnane and Takrouna populations, these results are in line with an ancient Euro Mediterranean background that has already been studied by archaeologists, particularly for the population of Sejnane.

The Andalusian population from Huelva reveals a high diversification of Y-DNA paternal lineages from haplogroup E: Identifying human male movements within the Mediterranean space

Background: Gene flow among human populations is generally interpreted in terms of complex patterns, with the observed gene frequencies being the consequence of the entire genetic and demographic histories of the population.Aims: This study performs a high-resolution analysis of the Y-chromosome haplogroup E in Western Andalusians (Huelva province). The genetic information presented here provides new insights into migration processes that took place throughout the Mediterranean space and tries to evaluate its impact on the current genetic composition of the most southwestern population of Spain.Subjects and methods: 167 unrelated males were previously typed for the presence/absence of the Y-chromosome Alu polymorphism (YAP). The group of YAP (+) Andalusians was genotyped for 16 Y-SNPs and also characterized for 16 Y-STR loci.Results: The distribution of E-M81 haplogroup, a Berber marker, was found at a frequency of 3% in our sample. The distribution of M81 frequencies in Iberia seems to be not concordant with the regions where Islamic rule was most intense and long-lasting. The study also showed that most of M78 derived allele (6.6%) led to the V13* subhaplogroup. We also found the most basal and rare paragroup M78* and others with V12 and V65 mutations. The lineage defined by M34 mutation, which is quite frequent in Jews, was detected as well.Conclusions: The haplogroup E among Western Andalusians revealed a complex admixture of genetic markers from the Mediterranean space, with interesting signatures of populations from the Middle East and the Balkan Peninsula and a surprisingly low influence by Berber populations compared to other areas of the Iberian Peninsula.

Temporal evolution of the ABO allele frequencies in the Canary Islands: the impact of the European colonization

The indigenous Canary Islands population suffered a strong cultural and genetic impact when they were colonized by Europeans in the fifteenth century. The molecular analysis of the ABO blood group gene on aboriginal and seventeenth to eighteenth century remains confirms the demographic history of the islands depicted by previous archaeological, anthropological, and genetic studies. ABO allele frequencies clearly related Canarian aborigines with North African Berber populations, its most probable source of origin, and is far related to Iberian and to the current population of the archipelago. The historical sample shows a congruent intermediate position testifying already a strong European influence that would go in augment since then to present times, affecting all the islands with the important exception of La Gomera.

Dhar Néma: from early agriculture to metallurgy in southeastern Mauritania

Dhar Néma, easternmost of the Tichitt escarpment chain, was neglected by archaeologists throughout the twentieth century. However, since 2000, two separate archaeological teams have conducted survey and test excavation work in the region. This article presents the final research results of one of these teams. Two sites in particular are closely examined: the settlement mound of Djiganyai with stratified intermittent occupation between c. 2000 and 800 cal. BC, and the multi-component site of Bou Khzama II, featuring structural elements, inhumations, and large scale iron-smelting remains. It is argued that Dhar Néma, although an ecologically favourable zone, was politically peripheral to the central Tichitt-Walata polity and comparatively sparsely settled. A period of relative regional abandonment, or widespread shifts in settlement, between 1400 and 1100 cal. BC is also noted. Regardless, a range of Pre-Tichitt and Tichitt Tradition settlements are documented for the region, including evidence for millet agriculture equivalent in age to that of the Dhar Tichitt region. It is suggested that the origins of Tichitt agriculture must now be sought in the third millennium BC. Concerning more recent periods, putative new data for first millennium BC iron metallurgy in the Dhar Néma region are presented and their technical aspects analysed. Finally, a revised hypothesis for the role of incoming Berber populations in the denouement of the Tichitt Tradition is considered, with cultural syncretism rather than replacement advocated.

The Complex and Diversified Mitochondrial Gene Pool of Berber Populations

The mitochondrial DNA variation of 295 Berber-speakers from Morocco (Asni, Bouhria and Figuig) and the Egyptian oasis of Siwa was evaluated by sequencing a portion of the control region (including HVS-I and part of HVS-II) and surveying haplogroup-specific coding region markers. Our findings show that the Berber mitochondrial pool is characterized by an overall high frequency of Western Eurasian haplogroups, a somehow lower frequency of sub-Saharan L lineages, and a significant (but differential) presence of North African haplogroups U6 and M1, thus occupying an intermediate position between European and sub-Saharan populations in PCA analysis. A clear and significant genetic differentiation between the Berbers from Maghreb and Egyptian Berbers was also observed. The first are related to European populations as shown by haplogroup H1 and V frequencies, whereas the latter share more affinities with East African and Nile Valley populations as indicated by the high frequency of M1 and the presence of L0a1, L3i, L4*, and L4b2 lineages. Moreover, haplogroup U6 was not observed in Siwa. We conclude that the origins and maternal diversity of Berber populations are old and complex, and these communities bear genetic characteristics resulting from various events of gene flow with surrounding and migrating populations.

Substructure of a Tunisian Berber Population as Inferred from 15 Autosomal Short Tandem Repeat Loci

Currently, language and cultural practices are the only criteria to distinguish between Berber autochthonous Tunisian populations. To evaluate these populations’ possible genetic structure and differentiation, we have analyzed 15 autosomal short tandem repeat loci (CSF1PO, D3S1358, D5S818, D7S820, D8S1179, D13S317, D16S539, D18S51, D21S11, FGA, TH01, TPOX, VWA, D2S1338, and D19S433) in three southern Tunisian Berber groups: Sened, Matmata, and Chenini-Douiret. The exact test of population differentiation based on allele frequencies at the 15 loci shows significant P values at 7 loci between Chenini-Douiret and both Sened and Matmata, whereas just 5 loci show significant P values between Sened and Matmata. Comparative analyses between the three Berber groups based on genetic distances show that P values for FST distances are significant between the three Berber groups. Population analysis performed using Structure shows a clear differentiation between these Berber groups, with strong genetic isolation of Chenini-Douiret. These results confirm at the autosomal level the high degree of heterogeneity of Tunisian Berber populations that had been previously reported for uniparental markers.

Substructure of a Tunisian Berber Population as Inferred from 15 Autosomal Short Tandem Repeat Loci

Currently, language and cultural practices are the only criteria to distinguish between Berber autochthonous Tunisian populations. To evaluate these populations' possible genetic structure and differentiation, we have analyzed 15 autosomal short tandem repeat loci (CSF1PO, D3S1358, D5S818, D7S820, D8S1179, D13S317, D16S539, D18S51, D21S11, FGA, TH01, TPOX, VWA, D2S1338, and D19S433) in three southern Tunisian Berber groups: Sened, Matmata, and Chenini-Douiret. The exact test of population differentiation based on allele frequencies at the 15 loci shows significant P values at 7 loci between Chenini-Douiret and both Sened and Matmata, whereas just 5 loci show significant P values between Sened and Matmata. Comparative analyses between the three Berber groups based on genetic distances show that P values for F(ST) distances are significant between the three Berber groups. Population analysis performed using Structure shows a clear differentiation between these Berber groups, with strong genetic isolation of Chenini-Douiret. These results confirm at the autosomal level the high degree of heterogeneity of Tunisian Berber populations that had been previously reported for uniparental markers.

Analytical Plug-In Method for Kernel Density Estimator Applied to Genetic Neutrality Study

The plug-in method enables optimization of the bandwidth of the kernel density estimator in order to estimate probability density functions (pdfs). Here, a faster procedure than that of the common plug-in method is proposed. The mean integrated square error (MISE) depends directly upon which is linked to the second-order derivative of the pdf. As we intend to introduce an analytical approximation of , the pdf is estimated only once, at the end of iterations. These two kinds of algorithm are tested on different random variables having distributions known for their difficult estimation. Finally, they are applied to genetic data in order to provide a better characterisation in the mean of neutrality of Tunisian Berber populations.

Identification of the CCR5-Δ32 HIV resistance allele and new mutations of the CCR5 gene in different Tunisian populations

Polymorphisms in some chemokine receptor genes are associated with susceptibility to and progression of human immunodeficiency virus-1 (HIV-1) infection. Most mutations detected in the CC-chemokine receptor 5 (CCR5) gene are specific to different populations. In this study, we focused on polymorphisms of the CCR5 coding region in three healthy populations from Tunisia, corresponding to a cosmopolitan population from Tunis, and two isolated Berber populations. In addition to the CCR5-Delta32 deletion, eleven single nucleotide polymorphisms were detected. Some of these point mutations were associated with the same genotype and even the same haplotype. The (L55Q-C101X), I124, V131F, T143N, A159V, I237, T239A and G301R alleles have not been described previously, whereas the CCR5-Delta32, L55Q, A335V and Y339F variants have already been reported in the literature. The distribution and frequency of these variants were different among the three groups studied, a result in agreement with the mosaic genetic structure of the Tunisian population. To determine whether these alleles affect HIV-1 transmission, we compared allele frequencies between healthy and HIV-1 infected individuals from Tunis. The frequency of the CCR5-Delta32 variant was significantly different between the two groups, leading us to conclude that this mutation might confer protection against HIV infection in Tunisian populations.

North African Berber and Arab Influences in the Western Mediterranean Revealed by Y-Chromosome DNA Haplotypes

We have analyzed Y-chromosome diversity in the western Mediterranean area, examining p49a,f TaqI haplotype V and subhaplotypes Vb (Berber) and Va (Arab). A total of 2,196 unrelated DNA samples, belonging to 22 populations from North Africa and the southern Mediterranean coast of occidental Europe, have been typed. Subhaplotype Vb, predominant in a Berber population of Morocco (63.5%), was also found at high frequencies in southern Portugal (35.9%) and Andalusia (25.4%). The Arab subhaplotype Va, predominant in Algeria (53.9%) and Tunisia (50.6%), was also found at a relatively high frequency in Sicily (23.1%) and Naples (16.4%); its highest frequency in Iberia was in northern Portugal (22.8%) and Andalusia (15.5%). In Iberia there is a gradient of decreasing frequencies in latitude for both subhaplotypes Va and Vb, related to eight centuries of Muslim domination (8th to 15th centuries) in southern Iberia.

Population genetic data of 15 tetrameric short tandem repeats (STRs) in Berbers from Morocco

The allele frequency distribution of 15 short tandem repeats (STR) loci contained in the AmpF lSTR Identifiler™ PCR Amplification Kit (Applied Biosystems), was determined in two Berber populations from Asni and Bouhria, in Central and Eastern Morocco, respectively. A total of 209 individuals were typed. No deviations from the Hardy–Weinberg equilibrium were observed for Asni at the 15 STRs loci whereas for the Bouhria samples, two loci (D5S818 and TH01) showed significant departures from Hardy–Weinberg expectations (after Bonferroni's correction). All loci are highly polymorphic and population differentiation tests showed that the Moroccan samples from Asni and Bouhria have significant differences in 4 out of 15 loci (D21S11, D7S820, D16S539 and TPOX). The aim of the study was to obtain accurate allele frequencies relevant for forensic applications. Comparative analyses between our population data and other population samples gathered from the literature are also presented.

MICA Polymorphism in a Population From North Morocco, Metalsa Berbers, Using Sequence-Based Typing

The MICA gene encodes a family of nonclassical major histocompatibility complex class I molecules. Data on MICA polymorphism in different populations are still limited. In the present study, MICA allele frequencies (af) were assessed in 82 unrelated healthy individuals from a Moroccan Berber population named Metalsa (ME) by means of sequence-based typing of exons 2, 3, 4, and 5. In consideration of the linkage disequilibrium existing between MICA and human leukocyte antigen (HLA) class I alleles, MICA/HLA-B, MICA/HLA-Cw, and MICA/HLA-A haplotype frequencies (hf) were estimated. A wide allelic distribution including 16 different MICA alleles was found in ME. The most common MICA alleles were MICA*00801 (af = 0.268), *004 (0.232), *00902 (0.140), *00901 (0.085), and *00901 (0.073). The most common MICA/HLA-B haplotypes were MICA*004-B*4403 and MICA*009-B*50 (hf = 0.113 for both these haplotypes). Some known MICA and HLA-B associations were confirmed in this population. Noteworthy was the high frequency of MICA*009 (af = 0.226); the high frequency of B*50 found in ME (af = 0.114) permitted us to evidence the associations of MICA*00902 with B*5001 (hf = 0.068) or *5002 (hf = 0.045), whereas MICA*00901 was mainly associated with B*5101 (hf = 0.038), which corresponds to the previously described association MICA*009/A6-HLA-B*51. This study extends the previous knowledge on MICA polymorphism to a North African white population and may have implications for disease associations and transplantation.