Population genetic structure and distribution of introduced American mink (<Emphasis Type="Italic">Mustela vison</Emphasis>) in Spain,based on microsatellite variation

The population genetic structure of an invasive species in Spain, the American mink (Mustela vison), was investigated using microsatellite DNA markers. This semi-aquatic carnivore, originating from North America, was imported into Europe for fur farming since the beginning of the 20th century. Due to massive escapes, farm damages, deliberate releases and/or accidents, feral mink populations were established in the aquatic ecosystems of many European countries, including Spain. We genotyped 155 American mink originating from the Spanish regions Basque Country, Catalonia, Castilla-Leon, Aragon, Valencia and Galicia using 10 polymorphic microsatellite loci to highlight population genetic structure, distribution and dispersal. M. vison populations in Spain appear differentiated and not yet connected by gene flow. Bayesian clustering analyses and spatial analyses of molecular variance detected four inferred clusters, overall coinciding with the sampled geographical localities. Preliminary testing shows moderate to large estimated effective population sizes. Molecular analyses result useful to provide baseline data for further research on the evolution of invasive mink populations, as well as support local management strategies and indirectly benefit the conservation of threatened species in Spain, such as the endangered European mink (Mustela lutreola), and the polecat (Mustela putorius), which share the habitat with the American mink. This paper is dedicated to the memory of Xavier Domingo-Roura.     

Multiple introductions and no loss of genetic diversity: invasion history of Japanese Rose, Rosa rugosa, in Europe

The shrub Rosa rugosa (Japanese Rose), native to East Asia, is considered one of the most troublesome invasive plant species in natural or semi-natural habitats of northern Europe and has proven very difficult to control. We aimed at disentangling the species’ invasion history in Europe, including determining the number of introductions and their geographic origin, and at investigating whether populations in the introduced and native ranges differ in genetic diversity, structure and degree of differentiation. We found that introduced (n = 16) and native (n = 16) populations had similar levels of genetic diversity at seven nuclear SSR (microsatellite) loci. European populations lack isolation by distance and are less genetically differentiated than are populations in East Asia. Multiple and at least three independent colonization events, one of which was particularly successful, gave rise to current R. rugosa populations in Europe. The geographic distribution patterns of these three genetic clusters could not be explained by natural dispersal alone, indicating that human mediated secondary dispersal is driving the expansion in Europe. One cluster representing three of the European populations was most likely derived from NW Japan, whereas the origin of the remaining thirteen populations could not clearly be resolved. The introduction and expansion in Europe occurred with no significant loss of genetic diversity. We conclude that high propagule pressure at the primary establishment phase is the most parsimonious explanation for this pattern. A potential for long distance seed dispersal, coastal habitat connectivity and an outcrossing breeding system are factors likely to have enabled populations of R. rugosa to avoid detrimental effects of genetic bottlenecks and will further increase the species’ range size and abundance in Europe. We recommend that human-mediated dispersal should be prevented in order to halt the continued expansion.     

Genetic biogeography of the rate “copper moss”,Scopelophila cataractae (Pottiaceae)

Scopelophila cataractae, one of the so-called copper mosses, has a broad geographic distribution that includes North, Central, and South America, Europe, and Asia, but is rare throughout its range. A genetic analysis of 32 populations from the United States, Europe, and Asia based on 15 putative allozyme loci indicates that levels of genetic diversity vary among geographic regions. Six European populations are fixed for the same alleles at all 15 loci, consistent with the hypothesis thatS. cataractae is a recent immigrant in that region. The species is more diverse in the U.S., where it appears to be native. Five populations collected on copper-enriched soils around shrines and temples in Tokyo are genetically monomorphic, but Asian populations from another Japanese site, India, and Nepal are exceptionally diverse in terms of numbers of alleles and multilocus haplotypes, total gene diversity (H_T), and in the degree of differentiation among populations (measured as Nei'sI andD). Long-distance dispersal has probably played an important role in the geographic history ofS. cataractae, but the species appears to be native in both the New and Old Worlds. Gene flow between plants disjunct on different continents is insufficient to explain the lack of geographically correlated morphological and genetic differentiation inS. cataractae.     

Testing hypotheses on the dispersal and evolutionary history of freshwater mussels (Mollusca: Bivalvia: Unionidae)

The relationship between dispersal and differentiation of the European freshwater mussel Unio pictorum (Linnaeus, 1758) was studied with molecular genetic methods. Forty‐two populations from France, Italy and central Europe were analysed. Genetic relationships were assessed from the geographical distribution of allele frequencies at 17 enzyme loci. Neighbouring groups of populations show small to moderate mean genetic distances (0.020 < D_mean < 0.263). With a few exceptions the genetic affinities of the populations are the closest within the same drainage basin. In central Europe and Northern Italy genetic differences between drainage systems are relatively large. Populations from north‐eastern Italy are genetically similar to Danubian populations. Mussels from the islands of Corsica and Sardinia are more closely related to populations from the Italian peninsula than to French populations from the Rhône drainage system. Genetic relationships within U. pictorum from central Europe reflect palaeogeographical relationships between river systems during the Pliocene and Pleistocene. Literature data on two North American unionid species and one European fish species show the same relationship between genetic diversity and the history of drainage systems, although the correlations are less strong. In France and Italy this correspondence is much less evident. Population dynamic processes and human activities leading to populational bottlenecks might have obscured it.     

Threatened freshwater pearl mussel <Emphasis Type="Italic">Margaritifera margaritifera</Emphasis> L. in NW Spain: low and very structured genetic variation in southern peripheral populations assessed using microsatellite markers

A genetic analysis of freshwater pearl mussel Margaritifera margaritifera populations from NW Spain, a peripheral area of its European distribution, was carried out using microsatellite markers. These populations were formerly reported as genetically differentiated on the basis of growth and longevity studies. Ten loci previously characterized in populations from central Europe were used to comparatively analyze the genetic variability at the southern edge of the species’ range. Iberian pearl mussel populations showed very low genetic variability and significant high genetic differentiation. Half of the total genetic diversity observed appeared to be distributed between populations, which suggested a highly structured adaptive potential in pearl mussel at the southern peripheral distribution of the species. Population distinctiveness was evidenced by assignment tests, which revealed a high accuracy of individual assignments to their population of origin. All data suggested low effective population size and major effects of genetic drift on population genetic structure. In order to avoid further loss of genetic variation in biologically distinctive populations from NW Spain, prioritization of genetic resources of this species is required for conservation and management.     

Different dispersal histories of lineages of the earthworm <Emphasis Type="Italic">Aporrectodea caliginosa</Emphasis> (Lumbricidae,Annelida) in the Palearctic

Earthworms are among the most abundant and ecologically important invasive species, and are therefore a good object for studying genetic processes in invasive populations. Aporrectodea caliginosa is one of the most widespread invasive earthworms in the temperate zone. It is believed to have dispersed from Europe to all continents except Antarctica. It is known that A. caliginosa consists of three genetic lineages, and genetic diversity is high both among and between them. We attempted to use that high genetic diversity to study A. caliginosa dispersal in the Palearctic based on a sample of 40 localities ranging from eastern Europe to the Russian Far East, and to compare our data to other studies on this species in western Europe and North America. Two genetic lineages were found in the studied sample. Only negligible decrease in genetic diversity was observed for the lineage 2 of A. caliginosa from West Europe to the Far East, suggesting multiple human-mediated introductions. In contrast, lineage 3 is abundant in West Europe and Belarus, but is absent from the East European Plain, the Urals, and the Far East. However, it is present in West Siberia, where it has greatly reduced genetic diversity, indicating long-distance dispersal accompanied by a bottleneck event. Thus, although these two lineages of A. caliginosa are morphologically indistinguishable, they have dramatic differences in their distributions and dispersal histories.     

Variation of colony types in the antFormica cinerea

Summary Formica cinerea is a rare ant species in northern Europe where it occurs in few isolated populations. Estimates of genetic relatedness among worker nestmates revealed very different colonial structures. Relatedness was g = 0.81 in one population, and g = –0.03 and = 0.01 in two others. These results indicate that some populations of the species have mainly monogynous colonies (perhaps with monandrous queens), whereas others consist of polygynous and possibly polydomous colonies. Genetic differentiation of closely located populations suggests restricted dispersal.     

Repeated jumps from Northwest Africa to the European continent: The case of peripheral populations of an annual plant

Peripheral populations (i.e., those occurring on the edge of a species’ distribution range) can have different origins and genetic characteristics, and they may be critical for the conservation of genetic diversity. We investigated European peripheral populations of Scrophularia arguta, a widespread, annual plant distributed from Arabia to Northwest Africa and Macaronesia. Only two small disjunct population groups of this species occur in Europe, specifically in West‐Central and Southeast Iberia. To disclose the origin of these populations and determine their importance for the conservation of S. arguta genetic diversity, we analyzed DNA sequences from two nuclear and two plastid regions and amplified fragment length polymorphism markers in populations sampled mainly across the western distribution range of the species, and modeled the species distribution under present and late Quaternary conditions. The analyses revealed the presence of three distinct lineages of S. arguta in Europe, as a result of multiple colonization waves at different times in the Quaternary. Two of these lineages, occurring in Southeast Iberia, are the result of more or less recent dispersal from Northwest Africa. In contrast, West‐Central Iberian populations are strongly differentiated from the remaining range of S. arguta and can be considered as peripheral relict populations. Our study is the first to demonstrate the occurrence of at least three colonizations of the European continent from Africa by a native plant species. The diverse histories and genetic makeup of the resulting populations confirm the importance of peripheral populations, and particularly of ancient relict populations, for the conservation of global genetic diversity in widespread species.     

Diffusion of the Nearctic leafhopper Scaphoideus titanus Ball in Europe: a consequence of human trading activity

Scaphoideus titanus Ball is a Nearctic leafhopper that was introduced for the first time in Europe probably at the beginning of the 20th century. In Europe, this species is a specialist on cultivated grapevines and is of great economic importance as the vector of Flavescence dorée (FD), a Grapevine Yellows disease caused by Candidatus Phytoplasma vitis. The Random Amplified Polymorphic DNA (RAPD) technique was employed to obtain genetic information about the diffusion and the structure of S. titanus populations. Two American and 14 European populations were analysed. A total of 188 reproducible bands, obtained from three arbitrary primers, were considered to assess the amount and the pattern of genetic variation within and among leafhopper populations. American populations showed high levels of intra-population polymorphism and dissimilarity and appeared to be the most isolated of all the tested samples. The results confirm the historical role of American samples as the sources for the more recently founded European populations. RAPD analyses revealed a weak genetic structure of European samples that could probably be explained invoking the human role in their diffusion. The non-natural spreading of S. titanus across Europe is in fact attributable to the exchange of grapevine canes and grafts carrying eggs that the insect laid under the bark to overwinter.     

Pathways of spread of the introduced ascidian Styela clava (Tunicata) in Northern Europe, as revealed by microsatellite markers

Styela clava, a solitary ascidian native to the NW Pacific, has become a conspicuous member of fouling communities in NW European waters. As its natural dispersal appears to be limited, the wide distribution of S. clava along coasts within its introduced range may be attributed to secondary spread assisted by human activities. Here, we used six microsatellite loci to examine the genetic diversity and extent of gene flow among S. clava populations in its European introduced range. Samples were collected from 21 populations within Europe (N = 808), 4 populations within the USA and two populations within the native range (Japan). Large variation in genetic diversity was observed among the European populations but were not explained either by the geographic distance from the first introduction area (i.e. Plymouth, UK) nor by the time elapsed since the introduction. No founder effect was observed in the introduced populations, except possibly in Puget Sound (USA). At least two different introductions occurred in Europe, identified as distinct genetic clusters: northern Danish populations (resembling one Japanese population), and the rest of Europe; a sample from Shoreham (England) possibly represents a third introduction. In North America, the population from the Atlantic was genetically similar to the majority of European populations, suggesting a European origin for populations on this seaboard, while populations from the Pacific coast were genetically similar to the same Japanese population as the Danish populations.     

Genetic and morphological divergence among Gravel Bank Grasshoppers, <Emphasis Type="BoldItalic">Chorthippus pullus</Emphasis> (Acrididae), from contrasting environments

Gravel Bank Grasshopper (Chorthippus pullus) populations inhabit two contrasting environments, pebbly gravel banks with scarce vegetation cover in mountainous areas along the Alps and lowland grasslands dominated by Common Heather (Calluna vulgaris). Heath populations of C. pullus have been rediscovered only recently, and show a distribution scattered across Central Europe. The wings are reduced in this species; thus, it has low potential for long-distance dispersal. We used sequence data on a newly developed non-coding nuclear marker from three gravel-bank and four heath populations to test whether grasshoppers from the two environments represent distinct lineages. Gravel-bank populations were studied in southern Germany (Bavaria), heath populations in eastern Germany (Brandenburg and Saxony) and Ukraine. We compared those genetic data with an analysis of variation in a suite of morphometric traits. Finally, we combined genetic and morphometric data to reconstruct a plausible scenario for the ecological shift observed in C. pullus. Our newly developed marker did not sort populations from contrasting environments in two monophyletic lineages. Nevertheless, we found a general lack of gene flow between the gravel-bank and heath populations. There was pronounced variation among populations in morphometric traits. That variation was partially partitioned by habitat type, and populations from the same habitat tended to be more similar than those from different habitats. Our data suggest that heath populations originated through northward expansion from multiple southern European refugia, and that the gravel-bank populations represent one of these sources. Patterns of genetic and morphometric divergence suggest that gravel-bank and heath populations may be in the process of incipient speciation.     

Genetic biogeography of the rare “copper moss,” Mielichhoferia elongata (Bryaceae)

Mielichhoferia elongata, one of the so-called “copper mosses,” has a broad but highly disjunctive geographic distribution and is rare throughout its range. A genetic analysis of 30 populations based on a survey of 21 allozyme loci reveals the following. 1) Total gene diversity at the specific level is high (0.41). 2) Within-population diversity is low, and over 90% of all genetic variation is among rather than within populations (mean G_ST = 0.93). 3) There is little differentiation in allele frequencies between North American and European populations. 4) Populations consist of one to six multilocus genotypes; 13 of the populations appear to consist of a single clone. 5) Colorado populations contain a tremendous reservoir of genetic variation (88% of all alleles found in the species in North America and Europe occur in one or more Colorado populations). 6) Populations in the eastern and western United States, and in Europe, contain subsets of the allelic diversity found in Colorado. The genetic structure of M. elongata suggests repeated dispersal and founding of populations.     

Genetic diversity of the relict moss <Emphasis Type="Italic">Rhytidium rugosum</Emphasis> (Hypnales) in Europe inferred from the ITS region (nrDNA)

Post-glacial survival, potential migration routes, genetic diversity and phylogeography of the boreal moss species Rhytidium rugosum have been studied. This species is considered to be one of glacial relics of the wide but scattered Holarctic range. According to molecular data sampling from the selected European, American and Asian populations high genetic diversity of this species is present, even if this species is mostly sterile and produced sex organs extremely rarely and spread mostly asexually. Analysing the internal transcribed spacer (ITS) of the nuclear ribosomal DNA, it can be concluded that the populations of this species survived glaciations in various places in Europe and settled and re-settled present range space in various times from various refuges.     

Historical human activities reshape evolutionary trajectories across both native and introduced ranges

The same vectors that introduce species to new ranges could move them among native populations, but how human‐mediated dispersal impacts native ranges has been difficult to address because human‐mediated dispersal and natural dispersal can simultaneously shape patterns of gene flow. Here, we disentangle human‐mediated dispersal from natural dispersal by exploiting a system where the primary vector was once extensive but has since ceased. From 10th to 19th Centuries, ships in the North Atlantic exchanged sediments dredged from the intertidal for ballast, which ended when seawater ballast tanks were adopted. We investigate genetic patterns from RADseq‐derived SNPs in the amphipod Corophium volutator (n = 121; 4,870 SNPs) and the annelid Hediste diversicolor (n = 78; 3,820 SNPs), which were introduced from Europe to North America, have limited natural dispersal capabilities, are abundant in intertidal sediments, but not commonly found in modern water ballast tanks. We detect similar levels of genetic subdivision among introduced North American populations and among native European populations. Phylogenetic networks and clustering analyses reveal population structure between sites, a high degree of phylogenetic reticulation within ranges, and phylogenetic splits between European and North American populations. These patterns are inconsistent with phylogeographic structure expected to arise from natural dispersal alone, suggesting human activity eroded ancestral phylogeographic structure between native populations, but was insufficient to overcome divergent processes between naturalized populations and their sources. Our results suggest human activity may alter species' evolutionary trajectories on a broad geographic scale via regional homogenization and global diversification, in some cases precluding historical inference from genetic data.     

Holocene re‐colonisation,central–marginal distribution and habitat specialisation shape population genetic patterns within an Atlantic European grass species

Corynephorus canescens (L.) P.Beauv. is an outbreeding, short‐lived and wind‐dispersed grass species, highly specialised on scattered and disturbance‐dependent habitats of open sandy sites. Its distribution ranges from the Iberian Peninsula over Atlantic regions of Western and Central Europe, but excludes the two other classical European glacial refuge regions on the Apennine and Balkan Peninsulas. To investigate genetic patterns of this uncommon combination of ecological and biogeographic species characteristics, we analysed AFLP variation among 49 populations throughout the European distribution range, expecting (i) patterns of SW European glacial refugia and post‐glacial expansion to the NE; (ii) decreasing genetic diversity from central to marginal populations; and (iii) interacting effects of high gene flow and disturbance‐driven genetic drift. Decreasing genetic diversity from SW to NE and distinct gene pool clustering imply refugia on the Iberian Peninsula and in western France, from where range expansion originated towards the NE. High genetic diversity within and moderate genetic differentiation among populations, and a significant pattern of isolation‐by‐distance indicate a gene flow drift equilibrium within C. canescens, probably due to its restriction to scattered and dynamic habitats and limited dispersal distances. These features, as well as the re‐colonisation history, were found to affect genetic diversity gradients from central to marginal populations. Our study emphasises the need for including the specific ecology into analyses of species (re–)colonisation histories and range centre–margin analyses. To account for discontinuous distributions, new indices of marginality were tested for their suitability in studies of centre–periphery gradients.     

Southern isolation and northern long‐distance dispersal shaped the phylogeography of the widespread,but highly disjunct,European high mountain plant Artemisia eriantha (Asteraceae)

We investigated the range dynamics of Artemisia eriantha, a widespread, but rare, mountain plant with a highly disjunct distribution in the European Alpine System. We focused on testing the roles of vicariance and long‐distance dispersal in shaping the current distribution of the species. To this end, we collected AFLP and plastid DNA sequence data for 17 populations covering the entire distributional range of the species. Strong phylogeographical structure was found in both datasets. AFLP data suggested that almost all populations were genetically strongly differentiated, with 58% of the overall genetic variation partitioned among populations. Bayesian clustering identified five groups of populations: Balkans, Pyrenees, Central Apennines, one southwestern Alpine population and a Widespread cluster (eastern Pyrenees, Alps, Carpathians). Major groups were supported by neighbor‐joining and NeighbourNet analyses. Fourteen plastid haplotypes were found constituting five strongly distinct lineages: Alps plus Pyrenees, Apennines, Balkans, southern Carpathians, and a Widespread group (eastern Pyrenees, northern Carpathians, Mt. Olympus). Plastid DNA data suggested that A. eriantha colonized the European Alpine System in a westward direction. Although, in southern Europe, vicariant differentiation among the Iberian, Italian and Balkan Peninsulas predominated, thus highlighting their importance as glacial refugia for alpine species, in temperate mountain ranges, long‐distance dispersal prevailed. This study emphasizes that currently highly disjunct distributions can be shaped by both vicariance and long‐distance dispersal, although their relative importance may be geographically structured along, for instance, latitude, as in A. eriantha. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 214–226.     

Mitochondrial and nuclear DNA phylogeography of Crassostrea angulata, the Portuguese oyster endangered in Europe

The respective status of the Portuguese oyster, Crassostrea angulata, and the Pacific oyster, Crassostrea gigas, has long been a matter of controversy. Morphological and physiological similarities, homogeneity of allozyme allelic frequencies between populations of the two taxa and the demonstration of hybridization lead most authors to suggest that they should be regrouped within the same species. The risk of introgression and the present expansion of C. gigas aquaculture in Europe raises the question of the need for preservation of C. angulata in Europe, as only a few populations remain. We studied European and Asian populations of C. gigas and C. angulata using microsatellite and mitochondrial DNA markers to estimate their genetic diversity and differentiation. The analysis of genetic distances and the distribution of allelic and haplotype frequencies revealed significant genetic differences between taxa, showing two clusters: (1) C. gigas French and Japanese populations and (2) C. angulata Portuguese and Taiwanese populations. The Asian origin of the Crassostrea angulata taxa is therefore confirmed. Unlike previous studies based on allozymes, significant nuclear genome differences were noted between C. angulata and C. gigas. Despite the presumed history of the introduction of C. angulata into Southern Europe, these populations did not show any significant reduction of variability compared to Taiwanese populations. Any conservation plans for European C. angulata populations should take its non-native origin into account. They represent a valuable genetic resources for European breeding program.     

Reduced genetic diversity and low effective size in peripheral northern European catfish Silurus glanis populations

Using 10 polymorphic microsatellites and 1251 individual samples (some dating back to the early 1980s), genetic structure and effective population size in all native and introduced Swedish populations of the European wels catfish or Silurus glanis were studied. Levels of genetic variability and phylogeographic relationships were compared with data from a previous study of populations in other parts of Europe. The genetically distinct Swedish populations displayed comparably low levels of genetic variability and according to one-sample estimates based on linkage disequilibrium and sib ship-reconstruction, current local effective population sizes were lower than minimum levels recommended for short-term genetic conservation. In line with a previous suggestion of postglacial colonisation from a single refugium, all Swedish populations were assembled on a common branch in a star-shaped dendrogram together with other European populations. Two distinct subpopulations were detected in upper and lower habitats of River Emån, indicating that even minor dispersal barriers may restrict gene flow for wels in running waters. Genetic assignment of specimens encountered in the brackish Baltic Sea and in lakes where the species does not occur naturally indicated presence of long-distance sea dispersal and confirmed unauthorised translocations, respectively.     

Demographic bottlenecks and low gene flow in remnant populations of the critically endangered <Emphasis Type="Italic">Berchemiella wilsonii</Emphasis> var<Emphasis Type="Italic">. pubipetiolata</Emphasis> (Rhamnaceae) inferred from microsatellite markers

Berchemiella wilsonii var. pubipetiolata (Rhamnaceae) is an endangered plant with only four remnant populations in eastern China. Population genetic information is essential for understanding population history and formulating conservation strategies for this species. Thirteen microsatellite loci were used to investigate genetic variation and population structure of the four remnant populations. Moderate levels of expected heterozygosity (H _E = 0.466–0.543) and low allelic diversity (A = 3.1–3.6 and A _R = 2.2–2.4, respectively) were observed within populations. Bottleneck tests found three out of four populations to deviate from mutation-drift equilibrium under the two-phase model (TPM), suggesting a recent population decline, which is congruent with known demographic history. The evolutionary history of the species seems dominated by genetic drift rather than gene flow. Low historical gene flow was inferred from several different approaches and N _m ranged from 0.582 by the private allele method to 0.783 by the coalescent method. Contemporary gene flow was also found to be even lower for only one first generation migrant was detected with individual-based assignment analysis. Restricted pollen and seed dispersal as well as a recent decline in population size associated with habitat fragmentation may have contributed to low levels of historical and contemporary gene flow, and resulted in a high genetic differentiation. Under this scenario, Berchemiella wilsonii var. pubipetiolata populations are expected to display more pronounced population genetic structure in the future as a result of increased inbreeding and genetic drift.     

The fourth paradigm pattern of post-glacial range expansion of European terrestrial species: the phylogeography of the Marbled White butterfly (Satyrinae, Lepidoptera)

Aim Three paradigm patterns of post‐glacial dispersal are known for terrestrial species in Europe. However, the possibility of a fourth arises with the Italian and the Balkan lineages expanding to central Europe and the Iberian one being trapped by the Pyrenees. We test this hypothesis by analysing the molecular biogeography of the Marbled White butterfly. Location Twelve populations distributed over a major part of the European range of Melanargia galathea and M. lachesis. Methods We studied 18 allozyme loci of 403 individuals from 12 populations. Butterflies were sampled in the field, frozen in liquid nitrogen and stored under these conditions until analysis. We used cellulose acetate plates for allozyme electrophoresis. Results We detected three major genetic lineages within the M. galathea/lachesis complex. The M. lachesis sample from the southern Pyrenees was strongly genetically differentiated from M. galathea (F_CT: 0.312). Melanargia galathea splits into two major genetic lineages (F_CT: 0.115), which both were found in post‐glacially invaded regions. The further differentiation within these lineages was comparably low (F_SC: 0.028). The genetic diversity within populations was high compared with other butterfly species. Main conclusions Our findings support the existence of a fourth pattern with only the Iberian lineage not contributing considerably to the post‐glacial colonization of central Europe. Preliminary studies in other butterfly species of dry grasslands support the importance of this pattern possibly representing a fourth paradigm. The high genetic diversity within populations might be one reason for the recently observed expansions at the northern distribution limits.