Northwestern song sparrow populations show genetic effects of sequential colonization

Two genetic consequences are often considered evidence of a founder effect: substantial loss in genetic diversity and rapid divergence between source and founder populations. Single-step founder events have been studied for these effects, but with mixed results, causing continued controversy over the role of founder events in divergence. Experiments of serial bottlenecks have shown losses of diversity, increased divergence, and rapid behavioural changes possibly leading to reproductive isolation between source and final populations. The few studies conducted on natural, sequentially founded systems show some evidence of these effects. We examined a natural vertebrate system of sequential colonization among northwestern song sparrows (Melospiza melodia). This system has an effectively linear distribution, it was probably colonized within the last 10,000 years, there are morphological and behavioural differences among populations, and the westernmost populations occur in atypical habitats for the species. Eight microsatellite loci from eight populations in Alaska and British Columbia (n = 205) showed stepwise loss of genetic diversity, genetic evidence for strong population bottlenecks, and increased population divergence. The endpoint population on Attu Island has extremely low diversity (H(E) = 0.18). Our study shows that sequential bottlenecks or founder events can have powerful genetic effects in reducing diversity, possibly leading to rapid evolutionary divergence.     

Serial founder effects and genetic differentiation during worldwide range expansion of monarch butterflies

Range expansions can result in founder effects, increasing genetic differentiation between expanding populations and reducing genetic diversity along the expansion front. However, few studies have addressed these effects in long-distance migratory species, for which high dispersal ability might counter the effects of genetic drift. Monarchs (Danaus plexippus) are best known for undertaking a long-distance annual migration in North America, but have also dispersed around the world to form populations that do not migrate or travel only short distances. Here, we used microsatellite markers to assess genetic differentiation among 18 monarch populations and to determine worldwide colonization routes. Our results indicate that North American monarch populations connected by land show limited differentiation, probably because of the monarch''s ability to migrate long distances. Conversely, we found high genetic differentiation between populations separated by large bodies of water. Moreover, we show evidence for serial founder effects across the Pacific, suggesting stepwise dispersal from a North American origin. These findings demonstrate that genetic drift played a major role in shaping allele frequencies and created genetic differentiation among newly formed populations. Thus, range expansion can give rise to genetic differentiation and declines in genetic diversity, even in highly mobile species.     

Maintaining genetic diversity and population panmixia through dispersal and not gene flow in a holocyclic heteroecious aphid species

Heteroecious holocyclic aphids exhibit both sexual and asexual reproduction and alternate among primary and secondary hosts. Most of these aphids can feed on several related hosts, and invasions to new habitats may limit the number of suitable hosts. For example, the aphid specialist Aphis glycines survives only on the primary host buckthorn (Rhamnus spp.) and the secondary host soybean (Glycine max) in North America where it is invasive. Owing to this specialization and sparse primary host distribution, host colonization events could be localized and involve founder effects, impacting genetic diversity, population structure and adaptation. We characterized changes in the genetic diversity and structure across time among A. glycines populations. Populations were sampled from secondary hosts twice in the same geographical location: once after secondary colonization (early season), and again immediately before primary host colonization (late season). We tested for evidence of founder effects and genetic isolation in early season populations, and whether or not late-season dispersal restored genetic diversity and reduced fragmentation. A total of 24 single-nucleotide polymorphisms and 6 microsatellites were used for population genetic statistics. We found significantly lower levels of genotypic diversity and more genetic isolation among early season collections, indicating secondary host colonization occurred locally and involved founder effects. Pairwise F(ST) decreased from 0.046 to 0.017 in early and late collections, respectively, and while genetic relatedness significantly decreased with geographical distance in early season collections, no spatial structure was observed in late-season collections. Thus, late-season dispersal counteracts the secondary host colonization through homogenization and increases genetic diversity before primary host colonization.     

Colonization of Daphnia magna in a newly created pond: founder effects and secondary immigrants

In habitats recently colonized by cyclical parthenogens, founder events lead to genetic differences between populations that do not erode quickly despite ongoing dispersal. By comparing the genetic composition during initial colonization with that of the diapausing egg bank at a local scale, we here present the relative contribution of the founding clones to the build-up of genetic diversity and differentiation of a newly established cladoceran population. We monitored the population genetic structure of Daphnia magna in one newly created pond as well as the diapausing egg banks of four water bodies in the neighbouring area. Our population was founded by four individuals. After the first growing season, the largest contribution to the sexually produced resting egg bank came from only two clones. Descendants of initially rare clones and potentially also additional immigrant clones profited from outbreeding vigour and increased their frequency during the first few years after colonization. Beyond this, no further significant changes in genetic structure were observed in the egg bank. At this point, priority effects became fully operational and led to sustained population genetic differentiation from nearby ponds. Our results support that colonization dynamics strongly influence within and among population genetic variation and evolutionary potential of populations.     

Patterns of genetic diversity reveal multiple introductions and recurrent founder effects during range expansion in invasive populations of Geranium carolinianum (Geraniaceae)

Genetic diversity, and thus the adaptive potential of invasive populations, is largely based on three factors: patterns of genetic diversity in the species'' native range, the number and location of introductions and the number of founding individuals per introduction. Specifically, reductions in genetic diversity (‘founder effects'') should be stronger for species with low within-population diversity in their native range and few introductions of few individuals to the invasive range. We test these predictions with Geranium carolinianum, a winter annual herb native to North America and invasive in China. We measure the extent of founder effects using allozymes and microsatellites, and ask whether this is consistent with its colonization history and patterns of diversity in the native range. In the native range, genetic diversity is higher and structure is lower than expected based on life history traits. In China, our results provide evidence for multiple introductions near Nanjing, Jiangsu province, with subsequent range expansion to the west and south. Patterns of genetic diversity across China reveal weak founder effects that are driven largely by low-diversity populations at the expansion front, away from the introduction location. This suggests that reduced diversity in China has resulted from successive founder events during range expansion, and that the loss of genetic diversity in the Nanjing area was mitigated by multiple introductions from diverse source populations. This has implications for the future of G. carolinianum in China, as continued gene flow among populations should eventually increase genetic diversity within the more recently founded populations.     

Dispersal and group formation dynamics in a rare and endangered temperate forest bat (Nyctalus lasiopterus,Chiroptera: Vespertilionidae)

For elusive mammals like bats, colonization of new areas and colony formation are poorly understood, as is their relationship with the genetic structure of populations. Understanding dispersal and group formation behaviors is critical not only for a better comprehension of mammalian social dynamics, but also for guiding conservation efforts of rare and endangered species. Using nuclear and mitochondrial markers, we studied patterns of genetic diversity and differentiation among and within breeding colonies of giant noctule bats (Nyctalus lasiopterus), their relation to a new colony still in formation, and the impact of this ongoing process on the regionwide genetic makeup. Nuclear differentiation among colonies was relatively low and mostly nonsignificant. Mitochondrial variation followed this pattern, contrasting with findings for other temperate bat species. Our results suggest that this may indicate a recent population expansion. On average, female giant noctules were not more closely related to other colony members than to foreign individuals. This was also true for members of the newly forming colony and those of another, older group sampled shortly after its formation, suggesting that contrary to findings for other temperate bats, giant noctule colonies are not founded by relatives. However, mother–daughter pairs were found in the same populations more often than expected under random dispersal. Given this indication of philopatry, the lack of mitochondrial differentiation among most colonies in the region is probably due to the combination of a recent population expansion and group formation events.     

Isolation by distance and gene flow in the Eurasian badger (Meles meles) at both a local and broad scale

Eurasian badgers, Meles meles, have been shown to possess limited genetic population structure within Europe; however, field studies have detected high levels of philopatry, which are expected to increase population structure. Population structure will be a consequence of both contemporary dispersal and historical processes, each of which is expected to be evident at a different scale. Therefore, to gain a greater understanding of gene flow in the badger, we examined microsatellite diversity both among and within badger populations, focusing on populations from the British Isles and western Europe. We found that while populations differed in their allelic diversity, the British Isles displayed a similar degree of diversity to the rest of western Europe. The lower genetic diversity occurring in Ireland, Norway and Scotland was more likely to have resulted from founder effects rather than contemporary population density. While there was significant population structure (F ST = 0.19), divergence among populations was generally well explained by geographic distance (P < 0.0001) across the entire range studied of more than 3000 km. Transient effects from the Pleistocene appear to have been replaced by a strong pattern of genetic isolation by distance across western Europe, suggestive of colonization from a single refugium. Analysis of individuals within British populations through Mantel tests and spatial autocorrelation demonstrated that there was significant local population structure across 3-30 km, confirming that dispersal is indeed restricted. The isolation by distance observed among badger populations across western Europe is likely to be a consequence of this restricted local dispersal.     

High genetic diversity in a remote island population system: sans sex

It has been proposed that long-distance dispersal of mosses to the Hawaiian Islands rarely occurs and that the Hawaiian population of the allopolyploid peat moss Sphagnum palustre probably resulted from a single dispersal event. Here, we used microsatellites to investigate whether the Hawaiian population of the dioicous S. palustre had a single founder and to compare its genetic diversity to that found in populations of S. palustre in other regions. The genetic diversity of the Hawaiian population is comparable to that of larger population systems. Several lines of evidence, including a lack of sporophytes and an apparently restricted natural distribution, suggest that sexual reproduction is absent in the Hawaiian plants. In addition, all samples of Hawaiian S. palustre share a genetic trait rare in other populations. Time to most recent ancestor (TMRCA) analysis indicates that the Hawaiian population was probably founded 49-51 kyr ago. It appears that all Hawaiian plants of S. palustre descend from a single founder via vegetative propagation. The long-term viability of this clonal population coupled with the development of significant genetic diversity suggests that vegetative propagation in a moss does not necessarily preclude evolutionary success in the long term.     

Genetic Diversity and the Reintroduction of Meadow Species

Abstract: Restoration of formerly nutrient‐poor and species‐rich grasslands generally leads to an increase in species diversity. However, species without a persistent seed bank and with poor dispersal ability often do not re‐establish spontaneously. Here, reintroduction is an option. If existing populations are comparable in their genetic composition, any population will do. This is not the case if populations have local adaptations. Unfortunately, whether populations are adapted locally is not easily determined, in contrast to assessing differentiation using neutral genetic markers. We used AFLP to study genetic diversity of Cirsium dissectum and Succisa pratensis within and among several Junco‐Molinion plant communities in the Netherlands (up to 200 km apart) that were potential source populations, and followed the reintroduction using seeds from these populations. Also, vegetative growth phase characteristics of three populations of C. dissectum were analyzed under controlled conditions. Most of the genetic variation in these cross‐fertilizing species was found within populations. Small but significant genetic differences in band frequencies were found among populations (F_st 0.100 ‐ 0.135). The first generation of reintroduced plants contained less polymorphic bands than the source populations. The genetic differences caused by reintroduction using a limited number of seeds (founder effects) were significant in all except one case (F_st 0.012 ‐ 0.101 between source and corresponding reintroduced population), but the magnitude was smaller than the source population differentiation. In assignment tests, reintroduced populations resembled their source population more than any other population, but all populations contained sizeable proportions of plants that were assigned to most similar plants from other populations, indicating that the populations are only marginally distinct. Calculations show that reintroduction from more than one source population introduces significantly more polymorphic bands into the new population, capitalizing on the existence of band frequency differences among populations.     

Patterns of genetic diversity in colonizing plant species: Nassauvia lagascae var. lanata (Asteraceae: Mutisieae) on Volcan Lonquimay, Chile

The effect of colonization on the distribution of genetic diversity within and among populations in relation to species characteristics remains an open empirical question. The objective of this study was to contrast genetic diversity within and among established and colonizing populations of Nassauvia lagascae var. lanata on Volcán Lonquimay (Araucanía Region, Chile), which erupted on 25 December 1988, and relate genetic diversity to biological characteristics of the populations. We analyzed a total of 240 individuals from 15 populations distributed along the Andes Cordillera using AFLP and obtained a total of 307 AFLP bands, of which 97.7% are polymorphic. Values of population differentiation (F(ST)) did not differ significantly among established and colonizing populations, but colonizing populations did have reduced levels of genetic divergence (as indicated by private and rare bands) and genetic variation (e.g., Shannon index). We conclude that a founder effect through limited numbers of founding propagules derived from nearby source populations has not yet been compensated for by subsequent population growth and migration. Low rates of secondary dispersal via running water, kin-structure within populations, and slow population growth seem to contribute to the slow recovery of genetic diversity.     

Shifting dispersal modes at an expanding species’ range margin

While it is generally recognized that noncontiguous (long‐distance) dispersal of small numbers of individuals is important for range expansion over large geographic areas, it is often assumed that colonization on more local scales proceeds by population expansion and diffusion dispersal (larger numbers of individuals colonizing adjacent sites). There are few empirical studies of dispersal modes at the front of expanding ranges, and very little information is available on dispersal dynamics at smaller geographic scales where we expect contiguous (diffusion) dispersal to be prevalent. We used highly polymorphic genetic markers to characterize dispersal modes at a local geographic scale for populations at the edge of the range of a newly invasive grass species (Brachypodium sylvaticum) that is undergoing rapid range expansion in the Pacific Northwest of North America. Comparisons of Bayesian clustering of populations, patterns of genetic diversity, and gametic disequilibrium indicate that new populations are colonized ahead of the invasion front by noncontiguous dispersal from source populations, with admixture occurring as populations age. This pattern of noncontiguous colonization was maintained even at a local scale. Absence of evidence for dispersal among adjacent pioneer sites at the edge of the expanding range of this species suggests that pioneer populations undergo an establishment phase during which they do not contribute emigrants for colonization of neighbouring sites. Our data indicate that dispersal modes change as the invasion matures: initial colonization processes appear to be dominated by noncontiguous dispersal from only a few sources, while contiguous dispersal may play a greater role once populations become established.     

'Sax-sess'-- genetics of primary succession in a pioneer species on two parallel glacier forelands

The primary succession on glacier forelands is characterized by a sequence of early and late successional species, but whether there is also a chronosequence at the intraspecific, genetic level is a matter of debate. Two opposing hypotheses differ in their prediction of genetic diversity in colonizing populations due to founder effects and postcolonization gene immigration. The development of genetic diversity in the pioneer Saxifraga aizoides was investigated along a successional gradient on two parallel glacier forelands, in order to test whether populations from older successional stages were less genetically diverse than populations from younger successional stages, and to locate the sources of the propagules that originally colonized new glacier foreland. Genetic diversity was determined with amplified fragment length polymorphisms, and potential sources of colonizing propagules were assessed via assignment tests. Our results indicate considerable postcolonization gene flow among populations on glacier forelands, since population differentiation was low and genetic diversity within populations was significantly higher. Molecular diversity and differentiation of populations did not develop linearly. Dispersal events within the glacier foreland, from the adjacent valley slopes, and from parallel glacier valleys were identified. In summary, it seems that the colonization of glacier forelands in the European Alps is highly dynamic and stochastic.     

The founding mothers: the genetic structure of newly established Daphnia populations

Colonization dynamics may strongly influence within and among population genetic variation and evolutionary potential of populations. We here analyze the genetic structure during the first three years of 12 cyclical parthenogenetic Daphnia populations in newly created pond habitats. One to three genotypes were observed to colonize the populations, indicating a limited number of founders. Pronounced changes in genetic structure were associated with hatching of sexual dormant eggs after a period of absence of the newly founded populations from the active community. Despite rapid colonization, genetic differentiation among populations was fairly high with limited decay over time, suggesting long-lasting founder effects. After initial colonization, no new alleles were observed in any of the populations, and our analyses suggests that this reflects reduced establishment success of later arrivals. Rare alleles increased in frequency, which likely is the result of inbreeding depression in selfed offspring of initially abundant clones, providing a fitness advantage to the sexual offspring of initially rare clones.     

Invasion genetics of vendace (Coregonus albula (L.)) in the Inari‐Pasvik watercourse: revealing the origin and expansion pattern of a rapid colonization event

Species invasions can have wide‐ranging biological and socio‐economic effects and are generally unwanted by legislation. Identification of the source population as well as the ecology and genetics of both the invader population and the receiving community is of crucial importance. The rapid invasion of a small coregonid fish vendace (Coregonus albula) in a major northern European subarctic watercourse has resulted in a labile ecological situation in the receiving community. The ecological impact of the invasion has been thoroughly documented, but the genetics of the invasion remains to be explored. We analyzed the genetic diversity and divergence patterns among the two possible source populations from southern Finnish Lapland and three colonists populations within the Inari‐Pasvik watercourse using ten microsatellite loci in order to (i) identify the most likely source of the invasion, (ii) reveal the dispersal pattern and genetic structure of the secondary expansion, and (iii) to investigate whether the initial introduction and the secondary expansion were associated with founder effects. We revealed that repeated translocation of vendace from Lake Sinettäjärvi into a tributary lake of L. Inari in 1964–1966 is the most plausible source for the invasion. Both the initial introduction and the secondary expansion were found not to be associated with significant founder effects. The secondary expansion followed a stepping stone pattern and the source and colonist populations of this expansion have undergone rapid genetic divergence within a period of 15–35 years (ca. 8–17 generations). The rapid divergence may be contributed to lack of gene flow among the source and colonist populations due to the extensive hydroelectric damming in the watercourse. Multiple introductions and substantial genetic variation in combination with the boom‐and‐bust population development of the species thus likely counteracted the founder effects as well as fueled the rapid establishment and expansion of this species within the Inari‐Pasvik watercourse.     

Looking back to go forward: genetics informs future management of captive and reintroduced populations of the black-footed rock-wallaby <Emphasis Type="Italic">Petrogale lateralis</Emphasis>

Active management is essential to the survival of many threatened species globally. Captive breeding programmes can play an important role in facilitating the supplementation, translocation and reintroduction of wild populations. However, understanding the genetic dynamics within and among wild and captive populations is crucial to the planning and implementation of ex situ management, as adaptive potential is, in part, driven by genetic diversity. Here, we use 14 microsatellite loci and mitochondrial Control Region sequence to examine the population genetics of both wild populations and captive colonies of the endangered warru (the MacDonnell Ranges race of the black-footed rock-wallaby Petrogale lateralis) in central Australia, to understand how historical evolutionary processes have shaped current diversity and ensure effective ex situ management. Whilst microsatellite data reveal significant contemporary differentiation amongst remnant warru populations, evidence of contemporary dispersal and relatively weak isolation by distance, as well as a lack of phylogeographic structure suggests historical connectivity. Genetic diversity within current captive populations is lower than in the wild source populations. Based on our genetic data and ecological observations, we predict outbreeding depression is unlikely and hence make the recommendation that captive populations be managed as one genetic group. This will increase genetic diversity within the captive population and as a result increase the adaptive potential of reintroduced populations. We also identify a new site in the Musgrave Ranges which contains unique alleles but also connectivity with a population 6 km away. This novel genetic diversity could be used as a future source for supplementation.     

Origin and phylogeography of African savannah elephants (<Emphasis Type="Italic">Loxodonta africana</Emphasis>) in Kruger and nearby parks in southern Africa

African savannah elephants (Loxodonta africana) occur in fragmented and isolated populations across southern Africa. Transfrontier conservation efforts aim at preventing the negative effects of population fragmentation by maintaining and restoring linkages between protected areas. We sought to identify genetic linkages by comparing the elephants in Kruger National Park (South Africa) to populations in nearby countries (Botswana, Mozambique, Zambia and Zimbabwe). We used a 446 base pair mitochondrial DNA (mtDNA) control region fragment (141 individuals) and 9 nuclear DNA (nDNA) microsatellite markers (69 individuals) to investigate phylogenetic relationships and gene flow among elephant populations. The mtDNA and nDNA phylogeographic patterns were incongruent, with mtDNA patterns likely reflecting the effects of ancient female migrations, with patterns persisting due to female philopatry, and nDNA patterns likely reflecting male-mediated dispersal. Kruger elephant heterozygosity and differentiation were examined, and were not consistent with genetic isolation, a depleted gene pool or a strong founder effect. Mitochondrial DNA geographic patterns suggested that the Kruger population was founded by elephants from areas both north and south of Kruger, or has been augmented through migration from more than one geographic source. We discuss our findings in light of the need for conservation initiatives that aim at maintaining or restoring connectivity among populations. Such initiatives may provide a sustainable, self-regulating management approach for elephants in southern Africa while maintaining genetic diversity within and gene flow between Kruger and nearby regions.     

Genetic composition of the Jadro softmouth trout following translocation into a new habitat

Genetic founder effects of a historical translocation (1965; app. 12 generations ago) of endangered softmouth trout from the only remaining indigenous population to an adjacent uninhabited river were investigated. A comparison of 15 microsatellites from contemporary samples in both the source and re-established populations indicated a very low level of genetic diversity within the two populations. Furthermore, considerable differences in estimates of variability and effective population sizes were detected between populations, but no recent bottlenecks were evident. Our data suggest that the re-established population has lost variation due to genetic drift associated with founder effects following the translocation. Accordingly, in relation to management, we suggest that the re-established population could serve as a secondary source of individuals to buffer possible extinction due to demographic events. Finally, conservation initiatives to enhance the census population size and increase levels of variability in the re-established population are proposed. Electronic Supplementary Material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Survival of cold-adapted species in isolated mountains: the population genetics of the Sudeten ringlet,Erebia sudetica sudetica,in the Jeseník Mts., Czech Republic

Relic populations of cold-adapted species, trapped in isolated mountain pockets within the temperate zone, are predicted to suffer considerably due to ongoing climate warming. The butterfly Erebia sudetica sudetica is an example restricted to the Eastern Sudety Mts. Here, the butterfly forms permanent populations on subalpine tall-herb grasslands, but also occupies woodland clearings and hay meadows at lower altitudes. We assume differences among the genetic diversities of the populations due to differences in the temporal continuity of these habitats. Therefore, 17 allozyme loci were analysed for 276 individuals from 13 different localities (six tall-herb stands, two meadows, five forest clearings) in the Jeseník Mts. with a maximum distance of 20 km among them. We obtained a significantly higher genetic diversity for the subalpine populations than for the forest clearing populations. The genetic differentiation among the forest clearing populations was higher than among the subalpine ones. They also showed a significant isolation-by-distance system. These findings support the idea that the lower-elevation populations might have been founded by more than one dispersal event from the subalpine sites, but also secondary colonisations and gene flow in the forest belt. Due to founder effects and possibly further subsequent bottlenecks, these forest clearing populations did not harbour the entire genetic diversity of the taxon. Therefore, conservation actions should focus on the subalpine tall-herb formation.     

Genetics of the early stages of invasion of the Lessepsian rabbitfish Siganus luridus

Information on the initial stages of dispersal and settlement are of great interest in understanding the dynamics of biological invasions and in designing management responses. A newly settled population of the Lessepsian rabbitfish migrant Siganus luridus, that arrived in Linosa Island (Sicily Strait) in 2000, offered a unique opportunity to examine the genetic variability of the early phase of invasion and the starting point to test genetic variation within and between colonist and source populations.Demographics and dynamic aspects of S. luridus in the Mediterranean were evaluated by using phylogeographic and demographic (coalescent) methods based on DNA sequences of the mitochondrial control region. Sequences from 95 S. luridus, 25 Siganus rivulatus, and one of Siganus (Lo) vulpinus and S. doliatus were used. Samples were collected in one locality in the Red Sea (Eilat) and three localities in the Mediterranean (Israel, Greece and Linosa, Italy). Data showed (for the first time in a Lessepsian migrant) a lowering of the genetic diversity of the invading population (Mediterranean) (haplotype diversity 0.879, nucleotide diversity 0.592) compared to the parental one (Red Sea) (haplotype diversity 0.978, nucleotide diversity 0.958).Within the Mediterranean populations, there was no pattern of regional separation and mitochondrial diversity appeared to be preserved during the Linosa colonization, with no traces of founder events. Such evidence agrees with the idea that Lessepsian migration involves many individuals from its earliest stages.     

Genetic variability and phylogeographical patterns of a nonindigenous species invasion: a comparison of exotic vs. native zebra and quagga mussel populations

There have been few investigations of the number of founding sources and amount of genetic variability that lead to a successful nonindigenous species invasion, although genetic diversity is believed to play a central role. In the present study, population genetic structure, diversity and divergence patterns were analysed for the zebra mussel Dreissena polymorpha [n=280 samples and 63 putative randomly amplified polymorphic DNA (RAPDs) gene loci] and the quagga mussel D. bugensis (n=136 and 52 loci) from 10 nonindigenous North American and six Eurasian sampling sites, representing their present‐day ranges. Results showed that exotic populations of zebra and quagga mussels had surprisingly high genetic variability, similar to those in the Eurasian populations, suggesting large numbers of founding individuals and consistent with the hypothesis of multiple colonizations. Patterns of genetic relationships indicate that the North American populations of D. polymorpha likely were founded by multiple source populations from north‐western and northcentral Europe, but not from southcentral or eastern Europe. Sampling areas within North America also were significantly divergent, having levels of gene flow and migration about twice those separating long‐established Eurasian populations. Samples of D. bugensis in Lakes Erie and Ontario were significantly different, with the former being more closely related to a native population from the Dnieper River, Ukraine. No evidence for a founder effect was discerned for either species.