The genetic distances between the inhabitants of nine Mediterranean countries and the three major human races

Gene frequency data of several genetic markers in 9 Mediterranean countries as well as in the 3 major human races were collected from the literature. Using these data the genetic distances between the inhabitants of the 9 countries and the 3 human races were determined. Algerians are closer to Negroids, while the other Mediterraneans are closer to Caucasoids.     

Allozymic variation of the endangered killifishAphanius iberus and its application to conservation

Synopsis Genetic differentiation and patterns of variability in the endangered Iberian endemic,Aphanius iberus, were analyzed by allozyme electrophoresis as a valuable database for conservation purposes. Genetic variability values expressed as heterozygosity (H = 0.015–0.097) were close to the values found in other members of Cyprinodontidae (H = 0.012–0.123). Polymorphism values (P = 0.125–0.542) were higher than reported in the literature (P = 0.036–0.150) for Cyprinodontidae. Significant correlation existed between salinity values and genetic variability expressed as heterozygosity (r = – 0.76, p < 0.01) and polymorphism (r = – 0.60, p = 0.04). Low genetic variability values (H = 0.024–0.055, P = 0.125–0.292) were exhibited by populations which inhabit salty lagoons. The highest values were found in populations occurring in marshes and irrigation channels (H = 0.051–0.097, P = 0.250–0.542). Associations among genetic, geographic and ecological parameters were tested using a Mantel test indicated that most of the genetic distances were explained by geographic distances but not by ecological factors, suggesting that isolation by distance could be the main factor explaining the differentiation between sites. According to the genetic distances obtained, two mainA. iberus groups were discernible: the Atlantic and Mediterranean. Genetic distances between both groups (DRogers = 0.179–0.261) were higher than values between recognized species of other cyprinodontids (DRogers = 0.11–0.27). On the basis of genetic distances we have dated the fragmentation of both populations to the Upper Miocene-Pliocene when most of the Mediterranean sea dried up. Subsequently, gene flow between Mediterranean and Atlantic populations was interrupted. The results of our genetic analyses suggested the existence of five operational conservation units (OCUs) forA. iberus. These units are defined as a continuous area limited by geographical boundaries, and inhabited by one or more populations sharing the same genetic pattern.     

Historical biogeography in a linear system: genetic variation of sea rocket (Cakile maritima) and sea holly (Eryngium maritimum) along European coasts

The exclusively coastal Cakile maritima and Eryngium maritimum represent a linear biogeographical system. Genetic variation among 25 individuals of C. maritima and 16 individuals of E. maritimum, from the coasts of Europe, North Africa and the Canary Islands, was analysed using random amplified polymorphic DNAs (RAPDs) and intersimple sequence repeats (ISSRs). Genetic distances (Dice) were calculated and used to investigate the correlation between genetic and geographical distances, to construct Neighbour Joining (NJ) trees, and to compare mean genetic distances between areas within and across species. Genetic distances and geographical distances measured along the coast are well correlated in Cakile and Eryngium. This implies that dispersal in both species is largely along the coast. The NJ analyses resulted in the recognition of Atlantic and Mediterranean clusters in both Cakile and Eryngium. The genetic distance between these two clusters is much larger in Eryngium (0. 285) than in Cakile (0.037). Mean genetic distances are substantially higher in the Mediterranean than in the Atlantic clusters in both species, and higher in Cakile than in Eryngium particularly in the Atlantic cluster. It is argued that all similarities and differences between the two species can be explained with the presumed distribution of the two species in the Würm glacial as reconstructed from their extant temperature requirements, the distribution of ice cover, permafrost, and sea surface temperatures in that period, and indirect fossil evidence.     

Patterns of gene flow inferred from genetic distances in the Mediterranean region.

The analysis of population structure may lead to inferences about demographic phenomena. In particular, regions of sharp genetic differentiation suggest the existence of factors that impaired gene flow and increased the evolutionary role of genetic drift. Here, we present an analysis of a data set of 10 allele frequencies in 39 populations of the Mediterranean region. As a preliminary step, we describe spatial patterns of allele frequencies using spatial autocorrelation analysis. We then construct a network connecting localities and estimate genetic distances along the edges of the network. By applying specific algorithms, we locate on the map the areas of sharpest genetic differentiation, or genetic boundaries. The main boundaries separate the northern and the southern coasts, especially in their western portions; in addition, several localities appear genetically isolated. The comparatively high genetic differentiation across the western Mediterranean, where the sea distances between localities are shorter, strongly suggests that the sea distance by itself can hardly be regarded as a major isolating factor among these populations. On the contrary, the decrease in genetic resemblance between populations of the 2 coasts as one proceeds westward may reflect an increased genetic exchange in the eastern Mediterranean basin or independent human dispersal along the 2 coasts or both.     

Genetic diversity of X-chromosome in populations of aboriginal Siberian ethnic groups: linkage disequilibrium structure and haplotype philogeography of ZFX locus

The structure of gene pool of the Siberian aboriginal population has been described based on the data on polymorphism of ZFX gene located on X-chromosome. In ten populations under study 49 haplotypes have been determined, three of which are presented with high frequency. Comparing the obtained results with the available data from HapMap project unique "African" haplotypes were revealed, which occurred in Yoruba population with the frequency of 3-7% and were not found in other populations. A coefficient of genetic differentiation of the Siberian ethnic groups under study amounted to 0.0486. Correlation analysis involving Mantel test did not reveal any significant correlations between a matrix of genetic distances and the matrices of geographic, linguistic and anthropological differences, where a maximum coefficient was obtained at the comparison with the anthropological matrix. Phylogenetic analysis proved strong isolation of African population from the other investigated ethnic groups. The Siberian populations were subdivided into two separate clusters: the first one included Yakuts, Buryats and Kets, while the second cluster included Altaians, Tuvinians and Khanty. A principal component analysis enabled to combine the investigated populations in three groups, which clearly differed by a degree of manifestation of Caucasoid and Mongoloid components. The first group included Europe inhabitants and one of Khanty populations, the second one--populations of South Siberia and China inhabitants. Mongoloid populations of East Siberia, the Japanese and Kets were combined in the third group. The results of barrier analysis revealed similar structure of genetic differentiation in the Siberian population. Linkage disequilibrium structure was obtained for six ethnic groups of Siberia. A unified linkage block by ten SNP of ZFX gene was found in five of the presented ethnic groups (excluding Ket population).     

Population genetics of the Malm? polymorphism of coagulation factor IX.

The distribution of 1,198 Malm? alleles was examined in 822 men from 16 indigenous populations and 188 women from 7 of the ethnic groups. Subjects were from several European countries, the Mediterranean, East Asia, and the USA (Anglo- and African-Americans). The frequencies of the rarer (Malm? B) allele were approximately equal across Europe, the highest frequencies (0.36) being in the French and Anglo-Americans; no population was observed with clearly the highest frequency. They diminished slightly at moderate distances from Europe (Tunisia, Ethiopia) and greatly at longer distances (East Asia and West Africa). In Orientals, the frequencies ranged from 0.07 (East Indians) to 0.03 (the Chinese) and from 0.0 to 0.15 in African-Americans. Assuming selective neutrality, the data are consistent with the European origin of the 'B' allele when the population was small and outward spread.     

Environmental Heterogeneity Explains the Genetic Structure of Continental and Mediterranean Populations of Fraxinus angustifolia Vahl

Tree species with wide distributions often exhibit different levels of genetic structuring correlated to their environment. However, understanding how environmental heterogeneity influences genetic variation is difficult because the effects of gene flow, drift and selection are confounded. We investigated the genetic variation and its ecological correlates in a wind-pollinated Mediterranean tree species, Fraxinus angustifolia Vahl, within a recognised glacial refugium in Croatia. We sampled 11 populations from environmentally divergent habitats within the Continental and Mediterranean biogeographical regions. We combined genetic data analyses based on nuclear microsatellite loci, multivariate statistics on environmental data and ecological niche modelling (ENM). We identified a geographic structure with a high genetic diversity and low differentiation in the Continental region, which contrasted with the significantly lower genetic diversity and higher population divergence in the Mediterranean region. The positive and significant correlation between environmental and genetic distances after controlling for geographic distance suggests an important influence of ecological divergence of the sites in shaping genetic variation. The ENM provided support for niche differentiation between the populations from the Continental and Mediterranean regions, suggesting that contemporary populations may represent two divergent ecotypes. Ecotype differentiation was also supported by multivariate environmental and genetic distance analyses. Our results suggest that despite extensive gene flow in continental areas, long-term stability of heterogeneous environments have likely promoted genetic divergence of ashes in this region and can explain the present-day genetic variation patterns of these ancient populations.     

Geographic limits of a clonal population of wheat yellow rust in the Mediterranean region

Most plant pathogens present complex life cycles, in which the clonal reproduction may impede the delimitation of population entities. By studying the genetic structure of the wheat yellow rust caused by Puccinia striiformis f.sp. tritici (PST), we highlighted difficulties impeding population delimitation in highly clonal species. Despite the high dispersal potential of PST, southern France isolates were shown to be divergent from a northwestern European population. A 2-year survey was performed in the Mediterranean region to assess the geographic distribution of southern isolates: 453 isolates collected in 11 countries were genotyped using 15 simple sequence repeat markers. A subsample was analysed for virulence against 23 resistance genes. The dominant strain in the western Mediterranean region was further studied with amplified fragment length polymorphism markers to test for a geographic substructure. Both 'individual'- and 'population'-centred analyses of polymorphism markers revealed two south-specific groups: a predominant group, with a broad variability and a wide distribution in both western and eastern Mediterranean countries, and a minor group in the western Mediterranean. The east–west gradient of genetic diversity suggested gene flow from the Middle East with subsequent founder effects and genetic divergence, and demonstrated the local survival of a western Mediterranean population. The high frequency of the resistance gene Yr8 observed in cultivars from Tunisia and Algeria may contribute to maintain the north/south structure observed in France. In addition to migration and local adaptation, the dynamics of clonal lineage diversification and replacement should be considered to define population entities in strongly clonal species.     

Great bustard population structure in central Spain: concordant results from genetic analysis and dispersal study.

We found significant sex differences in the mtDNA genetic structure and dispersal patterns of great bustards in a population of 11 breeding groups, "leks", in central Spain. The analysis of genetic distances showed that the female population was divided into three groups of leks separated by ca. 50 km, whereas male haplotypes were randomly distributed among leks. Genetic distances among pairs of leks were positively correlated with geographical distances in females but not in males. While female haplotype distributions were homogeneous among leks at close distances, differences in male genetic structure were highly variable even between two close leks. These results from genetic analyses were concordant with those from a radiotracking study on natal dispersal. Natal dispersal distances were higher in males than in females. Also, the frequency of movement of a female between two leks was positively correlated with their genetic affinity and geographical proximity. In males, the frequency of movement was correlated with geographical proximity but not with genetic affinity. Males dispersed among genetically unrelated leks, contributing to keep nuclear genetic diversity in the population, whereas females tended to be philopatric. These results suggest that isolation-by-distance influences the distribution of maternal lineages at a regional level.     

Phylogeographical lineages in brown trout (Salmo trutta): investigating microgeographical differentiation between native populations from Northern Spain

SUMMARY 1. The large microgeographical differentiation revealed by allozyme studies in brown trout ( Salmo trutta) populations is one of the most striking features of this species. Additionally, allozymes showed great genetic differences between Atlantic and Mediterranean populations on a macrogeographical scale.
2. This study was carried out in order to assess whether the great differences observed between Atlantic and Mediterranean populations persisted where the two are geographically close (the 'microgeographical scale'). Sixteen populations of brown trout, S. trutta , were screened for genetic variation at 25 allozyme loci. The sampling sites, which occupied a relatively small geographical area, were distributed across Cantabrian (Atlantic) and Mediterranean drainages in Northern Spain.
3. The neighbour-joining tree, inferred from Nei's genetic distance, showed that brown trout populations clustered into two different groups. These groups corresponded to the Cantabrian and the Mediterranean groups of populations, although no clear geographical pattern emerged within each of the groups. This geographical pattern is basically caused by significant differences in the frequency distribution of the CK-A1 * locus, with a higher frequency of * 115 in Cantabrian samples (0.586 ± 0.091) while allele * 100 was more frequent in Mediterranean samples (0.931 ± 0.038). In addition, this study revealed alleles exclusive to the Mediterranean and Cantabrian populations, agreeing with previous findings.
4. Genetic differentiation between Cantabrian and Mediterranean regions (14.19%) was similar to that estimated in Spain at a larger scale (13%), showing that most of the differences between the regions can be observed even in a small geographical area.     

Spatial graphs highlight how multi-generational dispersal shapes landscape genetic patterns

Current approaches that compare spatial genetic structure of a given species and the dispersal of its mobile phase can detect a mismatch between both patterns mainly due to processes acting at different temporal scales. Genetic structure result from gene flow and other evolutionary and demographic processes over many generations, while dispersal predicted from the mobile phase often represents solely one generation on a single time-step. In this study, we present a spatial graph approach to landscape genetics that extends connectivity networks with a stepping-stone model to represent dispersal between suitable habitat patches over multiple generations. We illustrate the approach with the case of the striped red mullet Mullus surmuletus in the Mediterranean Sea. The genetic connectivity of M. surmuletus was not correlate with the estimated dispersal probability over one generation, but with the stepping-stone estimate of larval dispersal, revealing the temporal scale of connectivity across the Mediterranean Sea. Our results highlight the importance of considering multiple generations and different time scales when relating demographic and genetic connectivity. The spatial graph of genetic distances further untangles intra-population genetic structure revealing the Siculo-Tunisian Strait as an important corridor rather than a barrier for gene flow between the Western- and Eastern Mediterranean basins, and identifying Mediterranean islands as important stepping-stones for gene flow between continental populations. Our approach can be easily extended to other systems and environments.     

The distribution and molecular diversity of the Eastern Atlantic and Mediterranean chthamalids (Crustacea, Cirripedia)

The three chthamalids Chthamalus stellatus , C. montagui and Euraphia depressa are common inhabitants of the intertidal zone in the Eastern Atlantic, Mediterranean Sea and Black Sea. In this study, we investigated the occurrence of these barnacles in a wide range of their distribution. Population divergences of these two species have been inferred using three molecular markers — internal transcribed spacer (ITS), elongation factor 1α (EF-1α) and cytochrome oxidase subunit I (COI). ITS sequences of C. stellatus were identical throughout the species range, whereas ITS sequences of C. montagui indicated that the Black Sea and Mediterranean populations are isolated from the Atlantic population. The COI and EF-1α sequences were the most variable and informative. They indicated a high genetic divergence between Atlantic, Mediterranean and Black Sea populations for C. montagui . In addition significant genetic structure was found among the populations of C. stellatus based on EF-1α but not COI. Interestingly, our molecular dating analysis correlated the pattern of diversification in C. montagui to major geological changes that occurred in the Mediterranean during the end of the Messinian and Pleiocene periods. We suggest that palaeohistory shaped the divergences between Chthamalus populations that have probably been maintained by current hydrographic conditions. Finally, COI phylogenetic analysis placed the genus Euraphia within the Chthamalus clade, suggesting the need for a taxonomic revision of Euraphia . This study represents the most detailed phylogeographical analysis of intertidal Mediterranean species to date, and shows that geological events have strongly shaped the current diversity pattern of this fauna.     

Genetic position of Andalusians from Huelva in relation to other European and North African populations: a study based on GM and KM allotypes

An understanding of population relationships in the Mediterranean region is crucial to the reconstruction of recent human evolution. Andalusia, the most southern region of Spain, has been continuously and densely occupied since ancient times and has a rich history of contacts with many different Mediterranean populations. Thus, to understand the Mediterranean peopling process, investigators should analyze the population relationships between the Iberian peninsula and northern Africa based on an assessment of genetic diversity that takes Andalusia into consideration. The aim of this study was to address the extent of genetic variation in the Iberian peninsula between its geographic extremes (Huelva and the Basque area) and to explain the intensity of the phylogenetic relationships between Andalusians and other neighboring populations, such as those from North Africa. We present, for the first time, results on allotype markers (GM and KM) of human immunoglobulins in the Andalusian population from Huelva. The most frequent GM haplotypes in Andalusia correspond to those that are also the most common in Europe. A sub-Saharan haplotype was found at a relatively high frequency compared to other Iberian samples, and a North Asian marker did not reach polymorphic frequencies in the study sample. A hierarchical cluster analysis based on the first two principal components (94.1% of the total genetic variance) revealed an interesting geographic structure for the 49 populations selected from the literature. The Huelva sample showed a central position in the multivariate space--despite being geographically located at one of the extremes of the Mediterranean basin--and clustered with most Western European populations. Western Europe and Eastern Europe (the latter group paradoxically including Italy and the major islands of the western Mediterranean) were differentiated. North African populations were grouped in two clusters that did not separate either Arabs and Berbers or their present-day countries. Analysis of immunoglobulin allotype markers shows that gene flow among human populations should generally be interpreted in terms of complex patterns, with the observed frequencies being the consequence of the entire genetic and demographic history of the population. Single historical events rarely determine gene frequencies in large human populations. Analysis of the GM system has shown that the Andalusian population from Huelva, as a result of its complex history, is not simply an outstanding part of the Mediterranean world but rather the genetic center of gravity of that world.     

Archeology, population genetics and studies of human racial ancestry.

Using traditional classification, many people have tried to determine the place and time of origin of the major human races. Two basically conflicting views have prevailed, one asserting that races developed as local variations in a species evolving phyletically over much of the Old World for up to one million years, and the other view holding that present races are local variations on populations recently expanding into most areas and replacing the previous hominid inhabitants, this occurring since the time of the Neanderthals. Both views are based on the same archeological data. Gene frequency data and time-calibrated genetic distance measures have recently been applied to this problem. Here we show that the probable demographic nature of Pleistocene populations has obscured genetic distances to such an extent that they cannot be used to discriminate between the two viewpoints of racial origins. The racial classifications themselves are probably not useful in this context and obscure the question, so that we presently do not have a scientifically valid understanding of racial origins.     

Ancient mitochondrial DNA evidence for prehistoric population movement: the Numic expansion

The mitochondrial DNA of modern Native Americans has been shown to fall into one of at least five haplogroups (A, B, C, D, or X) whose frequencies differ among tribal groups. The frequencies of these five haplogroups in a collection of ancient individuals from Western Nevada dating to between approximately 350-9,200 years BP were determined. These data were used to test the hypothesis, supported by archaeological and linguistic data, that the current inhabitants of the Great Basin, the Numic speakers, are recent immigrants into the area who replaced the previous non-Numic inhabitants. The frequency distributions of haplogroups in the ancient and modern Native Americans differed significantly, suggesting that there is a genetic discontinuity between the ancient inhabitants and the modern Numic speakers, providing further support for the Recent Numic Expansion hypothesis. The distribution of mitochondrial haplogroups of the ancient inhabitants of the Great Basin is most similar to those of some of the modern Native American inhabitants of California.     

Genetic Diversity of ND5 mitochondrial patterns in Ceratitis capitata (Diptera: Tephritidae) populations from Tunisia

The Mediterranean fruit fly (medfly) Ceratitis capitata (Diptera: Tephritidae) is known to be one of the most destructive and economically important agricultural pests worldwide. Several previous research projects have investigated the genetic makeup of regional populations of this pest and the relationships of populations from different areas of the world, including countries from the Mediterranean region. However, previously, little information has been reported on populations from Tunisia, despite the fact that this pest occurs in several agriculturally sensitive areas of this country. In order to study the genetic diversity of medfly populations within Tunisia, specimens were collected from the Coastal, Northern, Central and Southern regions of the country. Results using mitochondrial ND5 sequences show the presence of distinct haplotypes. This data used to analyze the levels of genetic variability within and between populations from Tunisia as well as from other countries in the Mediterranean region (Morocco and Israel) and in the world (Seychelles and Hawaii). This study also leads to a better understanding of the origin of new infestations and the colonization processes involving this pest.     

Testing the "malaria hypothesis" for the case of Thailand: a genetic appraisal

This study provides statistical analyses of allele frequencies for populations of Thailand, with an attempt to trace the roles of differential malarial selection and genetic admixtures on the observed frequency variation of certain red cell genetic abnormalities (the two beta-globin variants--hemoglobin E and beta-thalassemia--and G-6PD deficiency), probably evolving under malarial endemicity. It is found that frequencies of hemoglobin E vary accordingly with those of G-6PD deficiency, and with diverse malarial ecology. The levels of genetic diversity are greater for hemoglobin E and G-6PD deficiency than for most other nonmalarial related genetic markers, suggesting the evolution of these two genetic abnormalities under differential selection. Results of the Mantel's statistical test for correspondence between distance matrices suggest distinctive patterns of allele frequency differentiation between malarial-related and nonmalarial-related genetic loci. Correlations between beta-globin and G-6PD genetic distances, as well as those between both sets of distances and the malarial distances, are statistically significant. On the other hand, a correlation between malarial distances and the genetic distances for nonmalarial-related genetic loci is not significant statistically. A correlation between the beta-globin genetic distances and the genetic distances for nonmalarial-related genetic loci is, however, statistically significant. The latter result could be attributed largely to the clustering of relatively high hemoglobin E frequencies among genetically closely related populations of northeastern Thailand, whose recent homeland was Laos. The consistently low frequencies of beta-thalassemia observed in most studied populations are explained as a result of the replacement of this genetic variant by hemoglobin E, under long-term malarial selection.     

Distribution pattern and genetic structure of Aedes zammitii (Diptera: Culicidae) along the Mediterranean and Aegean coasts of Turkey

The Mariae species complex, consisting of Aedes mariae, Aedes phoeniciae, and Aedes zammitii, has a limited distribution worldwide. All three species are found in rocky habitats on the coastal areas of Mediterranean countries. Aedes phoeniciae and Ae. zammitii are two members of the Mariae complex that exist in Turkey. The aim of this study was to determine the distribution pattern and genetic structure of Ae. zammitii along the Mediterranean and Aegean regions. For this purpose, larval and adult samples of Ae. zammitii were collected from 19 different rocky habitats along the coastal regions of Antalya, Mu?la, Ayd?n, ?zmir, Bal?kesir, and Çanakkale provinces. DNA isolation was performed primarily from collected samples, and mitochondrial NADH dehydrogenase 4 (ND4) gene was amplified by polymerase chain reaction. Based on ND4 sequence analyses, 21 haplotypes were detected along the distribution range of the species. Analyses of molecular variance (AMOVA) and spatial analyses of molecular variance (SAMOVA) indicated six groups, and most of the variation was among groups, demonstrating the population structuring at group level. Isolation by distance analyses (IBD) showed a correlation between geographic and genetic distances.     

Genetic diversity in inter‐simple sequence repeat profiles across natural populations of Indian pomegranate (Punica granatum L.)

Pomegranate (Punica granatum L.), in the monogeneric family Punicaceae, is found in Iran, Afghanistan, India and Mediterranean countries. Iran is considered to be its primary centre of origin. In India, pomegranate occurs naturally only in the Western Himalayan regions of Jammu and Kashmir, Himachal Pradesh and Uttarakhand States. However, there is no information about genetic variation in wild pomegranate at population level. In this paper, we describe genetic diversity across natural populations of Indian pomegranate based on inter‐simple sequence repeat (ISSR) markers. Forty‐nine accessions representing eight populations from two regions were analysed using ISSR. Seventeen ISSR primers resulted in 268 polymorphic bands, with 87.01% polymorphism throughout the accessions. Pair‐wise population genetic distances ranged from 0.05 to 0.45, with a mean of 0.25 between populations. amova and Nei’s genetic diversity analyses revealed higher genetic variation within populations than among populations. A higher genetic differentiation (G_ST) was observed between the spatially distant populations, indicating a low level of genetic exchange (Nm) among these populations. However, clustering of populations was not in accordance with their geographical affiliations in the tree. The results indicate that the ISSR method is sufficiently informative and powerful to assess genetic variability in pomegranate, and that patterns of genetic variability observed among populations of wild pomegranate from the Western Himalaya differ. Estimation of genetic variation reported here provides a significant insight for in situ conservation and exploitation of genetic resources for this economically important species as potential breeding material.     

Geographical patterns of genetic variation in rosemary (Rosmarinus officinalis) in the Mediterranean basin

Climate changes during the Quaternary had important effects on the evolution of European plant species. The distribution of genetic variability in rosemary, a strictly Mediterranean species of reputed Plio‐Quaternary origin for which the diversification centre is hypothesized to be located in the western part of the Mediterranean basin, was investigated across the species range by using plastid microsatellites [plastid simple sequence repeat (cpSSR)] markers. Seven out of the 17 primer pairs screened were polymorphic, with up to four alleles, yielding a total of 17 size variants combined into ten haplotypes. A permutation test to investigate for geographical structure showed no significant differences between R_ST and G_ST, indicating that the species lacks geographical structure. Low correlation between genetic and geographical distances was shown by the Mantel test. Bayesian analysis identified two coancestry groups of populations. The distribution of genetic diversity supports the hypothesized origin in the western Mediterranean basin, and with the demographic expansion test indicates three different routes of migration: a northern route expanding along the northern side of the Mediterranean and two southern routes, one from west to east through North Africa and reaching Cyrenaica, and a second to the south‐west of the Iberian Peninsula, from where it came back to the south–central areas. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013 , 171 , 700–712.