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.     

Alu insertions in the Iberian Peninsula and north west Africa--genetic boundaries or melting pot?

The Western Mediterranean Basin joins a set of ethnically different populations as Iberians and Basques in the North shore and Berbers and Arab-speakers in the South one. In spite of this differentiation, they have maintained historical contacts since ancient times. The existence of a possible common genetic background (specially for Berbers and Iberians) together with the genetic impact of the Islamic occupation of the Iberian Peninsula during 7 centuries are some of the intriguing anthropological questions that have been studied in this area using several classical and DNA markers. The aim of this work is to present the results on a survey of polymorphic Alu elements in 10 human populations of the Western Mediterranean. Recent Alu subfamilies include a significant number of polymorphic Alu insertions in humans. The polymorphic Alu elements are neutral genetic markers of identical descent with known ancestral states. This fact turns Alu insertions into useful markers for the study of human population genetics. A total number of 14 Alu insertions were analyzed in 5 Iberian populations, 3 Berber groups from North-Western Africa, an Arab-speaker population from Morocco and a sub-Saharan ethnic group from Ivory Coast. The results of this study allow the genetic characterization of Berber populations, which show a certain degree of differentiation from Arab-speaking groups of the same geographic area. Furthermore, a closer genetic distance between South Spain and Moroccan Berbers as compared with other Spanish samples supports a major genetic influx consistent with some (but not all) previous genetic studies on populations from the two shores of the Gibraltar Straits.     

Ethnic communities in Israel: The genetic blood markers of the Moroccan Jews

One hundred and ninety-six Moroccan Jews now settled in Israel were typed for 7 blood groups, 12 red cell enzymes and 2 plasma protein systems. Their blood group picture is in agreement with results previously obtained on different samples of Moroccan Jews: rather high B in ABO, somewhat elevated frequencies of cDE and cDe in Rh and K in Kell. Differences in various blood markers exist between them and other North African Jewish communities. This fact, together with data on disease distribution and HLA frequencies, supports our assumption that Jews in the North African diaspora lived as small secluded isolates even within the same geographical zones. Comparisons with meager data on the neighboring non-Jewish populations do not disclose any resemblance to either Arab or Berber inhabitants of Morocco.     

Genetic signals of ancient decline in Aleppo pine populations at the species' southwestern margins in the Mediterranean Basin

Microsatellite markers were used to characterize the structure of genetic diversity in natural Moroccan Aleppo pine (Pinus halepensis Mill.) populations, the most southwesterly marginal populations of the species in the Mediterranean Basin. Twenty-two natural populations and one artificial population, located in four regions covering most of the natural range of P. halepensis in the country, were sampled. Across this range, towards the south and west (and towards high altitudes) the populations become increasingly discrete and discontinuous. The nuclear microsatellite marker analysis suggests that a large proportion of the Aleppo pines in Morocco have derived from a single genetic lineage, represented by a central group of 11 of the examined populations located in the High and Middle Atlas Mountains. In addition, two smaller groups, represented by the marginal southwestern High Atlas populations, and three still smaller north / northeastern groups of populations located in the Rif and northeast Middle Atlas Mountains, could be genetically distinguished. Further, coalescence analysis of historical demographic population patterns suggests that ancient bottlenecks occurred in all of the natural populations. However, the population differentiation and genetic diversity levels we found were good (F(st) =15.47), presumably because of the species' good potential for long-distance dispersal of seeds and high invasive capacity, which appear to have maintained a state of stable near-equilibrium, meta-population dynamics since ancient times.     

Female gene pools of Berber and Arab neighboring communities in central Tunisia: microstructure of mtDNA variation in North Africa

North African populations are considered genetically closer to Eurasians than to sub-Saharans. However, they display a considerably high mtDNA heterogeneity among them, namely in the frequencies of the U6, East African, and sub-Saharan haplogroups. In this study, we describe and compare the female gene pools of two neighboring Tunisian populations, Kesra (Berber) and Zriba (non-Berber), which have contrasting historical backgrounds. Both populations presented lower diversity values than those observed for other North African populations, and they were the only populations not showing significant negative Fu's F(S) values. Kesra displayed a much higher proportion of typical sub-Saharan haplotypes (49%, including 4.2% of M1 haplogroup) than Zriba (8%). With respect to U6 sequences, frequencies were low (2% in Kesra and 8% in Zriba), and all belonged to the subhaplogroup U6a. An analysis of these data in the context of North Africa reveals that the emerging picture is complex, because Zriba would match the profile of a Berber Moroccan population, whereas Kesra, which shows twice the frequency of sub-Saharan lineages normally observed in northern coastal populations, would match a western Saharan population except for the low U6 frequency. The North African patchy mtDNA landscape has no parallel in other regions of the world and increasing the number of sampled populations has not been accompanied by any substantial increase in our understanding of its phylogeography. Available data up to now rely on sampling small, scattered populations, although they are carefully characterized in terms of their ethnic, linguistic, and historical backgrounds. It is therefore doubtful that this picture truly represents the complex historical demography of the region rather than being just the result of the type of samplings performed so far.     

Genetic structure of Pinus pinaster Ait. populations in Morocco revealed by nuclear microsatellites

Pinus pinaster is one of the most popular conifers used for reforestation in Morocco and represents an economically and ecologically important species for the region. In this study, nuclear microsatellites (ncSSRs) are used to compare genetic structure and diversity estimates of natural populations of Moroccan maritime pine. Samples were collected among 10 natural populations distributed in three biogeographically different regions, the Rif Mountain, the Middle and the High Atlas. Forty-five nuclear alleles at seven variable loci were found with a mean of 6.4 alleles per locus. A number of private alleles (17.1%) were shown in populations from Rif and Middle Atlas. Moreover, in Morocco, P. pinaster showed a lower genetic diversity than in other parts of its geographic range. Significant departures from Hardy–Weinberg equilibrium with excess homozygosity are observed indicating a high level of mating inside populations. Genetic diversity was structured with high variability among populations (Fst = 12%). Results show a correlation between genetic and geographic distances with an R-squared of 0.436. Two clusters were found using STRUCTURE, whereas three main clusters can be distinguished based on genetic distances of phylogenetic tree. Genetic relationships among maritime pine populations in Morocco appear to be related to historical, ecological as well as anthropogenic factors, suggesting the need for conservation strategies at the population level.     

Renal cell cancer in Israel: Sex and ethnic differences in incidence and mortality, 1980–2004

Background: The causes of renal cell cancer (RCC) remain largely unexplained. While the incidence is generally higher in men than in women, little has been reported on ethnic differences. We examine trends in RCC incidence and mortality rates among Israeli Arab and Jewish populations and compared with the rates in other countries. Methods: Age-adjusted RCC incidence and mortality rates in Israel, during 1980–2004, were calculated by sex and population group, using the National Cancer Registry. They were compared with the United States based on the Surveillance Epidemiology and End Results [SEER] program and the IARC database for international comparisons. Results: While RCC incidence rates in Israel are similar to the United States and the European average, the rates are significantly higher among Israeli Jews than Arabs. Men are affected more than women. Incidence rates over the last 24 years have increased among all men and Jewish women, but not among Arab women. Among men, the incidence rate ratio for Jews to Arabs declined from 3.96 in 1980–1982 to 2.34 in 2001–2004, whereas for women there was no change. The mortality rates were higher among Jews than Arab and among men than women. There were no significant change in the mortality rates and rate ratios. Conclusions: Our findings demonstrate marked ethnic differences in RCC in Israel. The lower incidence among Arabs stands in contrast to the higher prevalence of potential risk factors for RCC in this population group. Genetic factors, diet and other lifestyle factors could play protective roles.     

Population biology of two helminth parasites of flatfishes from the Atlantic coast of Morocco

On the Atlantic Moroccan coast, two species of pleuronectiform fish coexist, wedge sole ( Citharus linguatula ) a temperate species, and spotted flounder ( Dicohgoglossa cuneata ) a tropical one. These two species are the definitive host for Bothriocephalus andresi and Acanthocephaloïdes propinquus , respectively. A negative binomial fits the dispersion of B. andresi in the population of C. linguatula . The fish become parasitized between their first and third year, while still immature, with the highest abundances in the third year. After the fish become ichthyopagous, the level of infection drops drastically. We postulate that copepods are the sole source of infection. The presence of B. andresi in both Atlantic and Mediterranean populations of C. linguatula shows that the parasite follows the host during its migration along the Mediterranean coasts.
The prevalence of A. propinquus remains above 50% as a consequence of the large numbers of amphipods included in the diet of D. cuneata , regardless of the age of the fish or the season. The amphipods, Phtisica marina and Pariambus typicus are an important food and represent potential intermediate hosts for A. propinquus . The population dynamics of A. propinquus on the Atlantic Moroccan coasts are similar to those observed in parasite populations of Gobius niger in the Gulf of Lion (French Mediterranean). The observation of B. andresi and A. propinquus in both Moroccan Atlantic and Mediterranean fishes highlights the problem of the evolution of these parasite populations during the colonization of the Mediterranean by the hosts.     

Population demographic structure of the city of Ulan Ude: Ethnic composition and age at marriage

The marriage structure of the population of Ulan Ude, the capital of Buryat Republic, which is characterized by a mixed ethnic composition, has been studied. Differences between Ulan Ude districts in a number of genetic demographic parameters have been found. In ethnic terms, the city district populations differ from one another in the number of ethnic groups and the proportions of the largest groups (Russians and Buryats). In the total Ulan Ude population, the proportions of Russian men and women among persons contracting marriages are 71.44 and 73.79%, respectively; those of Buryat men and women are 23.36 and 22.79%, respectively. The total proportion of all other ethnic groups among both men and women contracting marriages is no more than 5.2%. The mean ages at first marriage in both indigenous and immigrant populations of Ulan Ude are relatively old; these values are 25.9 and 24.72 years for Buryat men and women and 24.86 and 22.69 for non-indigenous men and women, respectively.     

Linkage disequilibrium mapping places the gene causing familial Mediterranean fever close to D16S246.

This report presents refined genetic mapping data for the gene causing familial Mediterranean fever (FMF), a recessively inherited disorder of inflammation. We sampled 65 Jewish, Armenian, and Arab families and typed them for eight markers from chromosome 16p. Using a new algorithm that permits multipoint calculations for a dense map of markers in consanguineous families, we obtained a maximal LOD score of 49.2 at a location 1.6 cM centromeric to D16S246. A specific haplotype at D16S283-D16S94-D16S246 was found in 76% of Moroccan and 32% of non-Moroccan Jewish carrier chromosomes, but this haplotype was not overrepresented in Armenian or Arab FMF carriers. Moreover, the 2.5-kb allele at D16S246 was significantly associated with FMF in Moroccan and non-Moroccan Jews but not in Armenians or Arabs. Since the Moroccan Jewish community represents a relatively recently established and genetically isolated founder population, we analyzed the Moroccan linkage-disequilibrium data by using Luria-Delbrück formulas and simulations based on a Poisson branching process. These methods place the FMF susceptibility gene within 0.305 cM of D16S246 (2-LOD-unit range 0.02-0.64 cM).     

Genetic structure of Tunisian ethnic groups revealed by paternal lineages

Tunisia has experienced a variety of human migrations that have modeled the myriad cultural groups inhabiting the area. Both Arabic and Berber-speaking populations live in Tunisia. Berbers are commonly considered as in situ descendants of peoples who settled roughly in Palaeolithic times, and posterior demographic events such as the arrival of the Neolithic, the Arab migrations, and the expulsion of the "Moors" from Spain, had a strong cultural influence. Nonetheless, the genetic structure and the population relationships of the ethnic groups living in Tunisia have been poorly assessed. In order to gain insight into the paternal genetic landscape and population structure, more than 40 Y-chromosome single nucleotide polymorphisms and 17 short tandem repeats were analyzed in five Tunisian ethnic groups (three Berber-speaking isolates, one Andalusian, and one Cosmopolitan Arab). The most common lineage was the North African haplogroup E-M81 (71%), being fixed in two Berber samples (Chenini-Douiret and Jradou), suggesting isolation and genetic drift. Differential levels of paternal gene flow from the Near East were detected in the Tunisian samples (J-M267 lineage over 30%); however, no major sub-Saharan African or European influence was found. This result contrasts with the high amount of sub-Saharan and Eurasian maternal lineages previously described in Tunisia. Overall, our results reveal a certain genetic inter-population diversity, especially among Berber groups, and sexual asymmetry, paternal lineages being mostly of autochthonous origin. In addition, Andalusians, who are supposed to be migrants from southern Spain, do not exhibit any substantial contribution of European lineages, suggesting a North African origin for this ethnic group.     

Population structure of Moroccan water frogs: genetic cohesion despite a fragmented distribution

This study analyses the allozymic variation of 20 presumptive loci in eight populations of Rana saharica from Morocco. Populations were collected from the very different climatic zones of this country: the Rif area, the Atlas mountains and the desert. Moroccan water-frog populations are genetically well differentiated from the geographically closed Algerian populations. Thus, to check if such a differentiation process is taking place within Moroccan water frogs, we attempted to analyse the genetic structure and patterns of gene flow of Moroccan populations, by means of estimates of Hardy-Weinberg equilibrium, F-statistics and indirect measures of gene flow. F_st(0.250) and F_is(0.254) values were similar, which means that both intra and interpopulation differentiation contribute equally to the amount of genetic divergence revealed. F_is values indicated some degree of structure within ponds, which is possibly related to the homing behaviour of some amphibians. On the other hand, F_st and genetic distances between populations were not very high. Despite the low levels of gene flow estimated, together with the homing behaviour revealed and the spatially discontinuous distribution, it was found that genetic differentiation among populations was not as high as expected. The likelihood of genetic homogeneity being the consequence of continuous population extinction and recolonization events is discussed.     

Sex ratio and sexual dimorphism in mountain dioecious thuriferous juniper (Juniperus thurifera L., Cupressaceae)

Thuriferous juniper ( Juniperus thurifera L), a dioecious bush or tree is only found in isolated parts of the western Mediterranean: France, Spain, Algeria and Morocco. These mountain juniper stands are seriously endangered in Morocco as a result of intensive wood removal, and in Europe as a result of recolonization of stands by pines or oaks. Field studies were conducted to investigate sex ratio and sexual dimorphism, never previously examined, in eight different populations in the Atlas mountains and, for comparison, in one of two populations in the French Pyrenees. The sex ratio was female-biased for six of the eight Moroccan stands and especially for the oldest populations. The Pyrenean population showed a similar female-biased ratio. This particular sex ratio is possibly linked to cost of reproduction, paid by both males and females. Sex ratios can also be linked to population dynamics. Males begin to flower slightly younger than females, which explains their apparent dominance in young populations in Morocco or in a recolonization zone with young trees in the Pyrenees. Studies concerning sexual dimorphism in the western High Atlas sites showed no significant difference in phytomass between males and females. Females appear to be significantly taller but with a lower radial growth. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 138 , 237–244.     

Range-wide phylogeography of Juniperus thurifera L., a presumptive keystone species of western Mediterranean vegetation during cold stages of the Pleistocene

We investigate the range-wide population structure and phylogeography of thuriferous juniper (Juniperus thurifera L.), a species with a highly disjunct distribution in the western Mediterranean. We genotyped a total of 327 individuals from 20 populations using amplified fragment length polymorphisms (AFLP). Different analyses such as principal co-ordinate analysis (PCoA), nonmetric multidimensional scaling of F(ST) distances among populations, unweighted pair group method with arithmetic mean (UPGMA), and Bayesian clustering revealed that the Strait of Gibraltar acted as an efficient barrier against gene flow between the Moroccan and European populations for a very long time, and consequently support that the Moroccan populations should be recognised as a distinct subspecies (J. thurifera L. subsp. africana (Maire) Romo and Boratyńsky). The Algerian population was genetically more closely related to the European than to the Moroccan ones, probably due to dispersal events from Europe to Algeria. With respect to the mainland European populations, our data are not conclusive to reject any of the two following hypotheses: (1) the Iberian Peninsula was subdivided into different gene pools, and was the source for the colonisation of the Pyrenees and the Alps; and (2) the pattern we see today is partly the result of immigration into the Iberian Peninsula, e.g. from the Alps. Finally, the Corsican population was closely related genetically to two northern Iberian populations most probably due to relatively recent long-distance dispersal.     

Analysis of digital patterns in the Basque valley of Deba: multivariate comparison with other populations

Eighty-six females and 79 males from the Basque valley of Deba were analyzed with respect to their digital patterns using correspondence analysis. We found that there was a significant sexual difference for only one type of pattern, tented arch. Also, this population was compared with other Basque valley populations and with other Spanish populations. We found great variability among Basque subpopulations, despite their supposed common origin. Dermatoglyphic distances for some traits can be interpreted as genetic distances because there is high heritability of these traits. The results of the comparison between the Deba valley population and other Spanish populations showed that the Deba population is markedly distant from the other populations.     

Recent dynamics of the wet pastures at Oukaimeden plateau (High Atlas mountains, Morroco)

Wetlands are a very important universal heritage; in the Moroccan High Atlas mountains the geomorphology and the slopes evolution determine the existence of the numerous projecting ledges of pozzines on plateaus and oozing. This is particularly the case in Oukaimeden site located at 75 km south of Marrakesh. These wet grasslands have been developed on small surface areas supported by soils water accumulation; and close to the sources and on some banks constitute wetlands of high floristic richness, which makes them very coveted pastures by local populations. These wet pastures are highly sensitive to climatic factors. Recent climatic changes, the anthropic and pastoral pressures and the global environmental changes, affect these zones at the level of their structure (areas, soil characteristics…) and their ecological functions. In the Oukaimeden plateau, the use of aerial photography made it possible to carry out a detailed follow up of the hydraulic adjustments (dam construction in the 1970s) as well as the setting up of new adjustments (tracks, paths, drain channel) on the structure and functioning of these grasslands.     

Genetic Risk Score of NOS Gene Variants Associated with Myocardial Infarction Correlates with Coronary Incidence across Europe

Coronary artery disease (CAD) mortality and morbidity is present in the European continent in a four-fold gradient across populations, from the South (Spain and France) with the lowest CAD mortality, towards the North (Finland and UK). This observed gradient has not been fully explained by classical or single genetic risk factors, resulting in some cases in the so called Southern European or Mediterranean paradox. Here we approached population genetic risk estimates using genetic risk scores (GRS) constructed with single nucleotide polymorphisms (SNP) from nitric oxide synthases (NOS) genes. These SNPs appeared to be associated with myocardial infarction (MI) in 2165 cases and 2153 controls. The GRSs were computed in 34 general European populations. Although the contribution of these GRS was lower than 1% between cases and controls, the mean GRS per population was positively correlated with coronary incidence explaining 65–85% of the variation among populations (67% in women and 86% in men). This large contribution to CAD incidence variation among populations might be a result of colinearity with several other common genetic and environmental factors. These results are not consistent with the cardiovascular Mediterranean paradox for genetics and support a CAD genetic architecture mainly based on combinations of common genetic polymorphisms. Population genetic risk scores is a promising approach in public health interventions to develop lifestyle programs and prevent intermediate risk factors in certain subpopulations with specific genetic predisposition.     

Phylogeography of North African Atlas cedar (Cedrus atlantica, Pinaceae): Combined molecular and fossil data reveal a complex Quaternary history

Northwest Africa is a major hotspot of plant biodiversity, but very little is known about the Quaternary range dynamics of plant species in this region. Here we investigate the range-wide population structure and phylogeography of Atlas cedar (Cedrus atlantica), an emblematic forest tree endemic to Morocco and Algeria. We genotyped 261 individuals from 11 populations using AFLP markers. Data were analyzed using both conventional F(ST)-based techniques and Bayesian clustering. Overall population differentiation was high (F(ST) = 0.25). Two major groups of populations were identified, one distributed through the Rif and Middle Atlas mountains in Morocco and the other through the Algerian Tell Atlas and Aurès mountains as well as the Middle Atlas. Combined molecular and fossil data indicate that C. atlantica survived the Last Glacial Maximum in at least three disjunct refugia along the coast of the Mediterranean Sea, whereas the Middle Atlas, today the core of the species range, has been colonized relatively recently (<10000 yr BP). The colonization history of individual populations has left clear imprints in their present-day diversity, which may vary greatly even between nearby stands. Our study illustrates how integrating different data sources and analytical approaches can help elucidate complex range dynamics that would otherwise remain undeciphered.