Patterns Of Allelic Variation Research Articles

Documentation of extensive genetic diversity in the Ovar-DRB1 gene in native Turkish sheep

Indigenous breeds have a high level of genetic diversity that might contribute to develop animal breeds with desired traits such as disease resistance and high productivity. Major histocompatibility complex (MHC) is a key component of adaptive immune system and consists of highly polymorphic genes that take part in adaptive immune response and disease resistance. Exploring and understanding the effect of polymorphisms in MHC could be beneficial to future animal breeding strategies. In this study, we sequenced the highly polymorphic Exon2 of the ovine DRB1 gene using Sanger sequencing to explore the diversity of this gene in six indigenous Turkish sheep breeds and two crossbreeds. In total, 894 haplotypes from 447 sheep were investigated, and 69 different haplotypes including 27 novel ones were identified. Among the identified haplotypes there were common and breed specific haplotypes. There was a relatively high diversity of the alleles within indigenous breeds. Allelic diversity patterns were mostly associated with geographical differences. The results of this study highlight the genetic variation within indigenous breeds which has important implications for biodiversity and the adaptability of breeds to specific environments. There is value to further studies which include other genomic regions and traits, and these could guide breeding strategies.

Genetic diversity of native and introduced Phragmites (common reed) in Wisconsin.

Two subspecies of common reed (Phragmites australis; Poaceae) exist in northern North America: the native P. australis subsp. americanus and the introduced P. australis subsp. australis. There are numerous native populations in Wisconsin, in addition to the more recently established populations of the introduced subspecies. We studied populations of both P. australis subspecies across Wisconsin in order to characterize the genetic diversity of both subspecies and to investigate whether any instances of hybridization could be ascertained in Wisconsin. Using eight microsatellite markers, we found minimal overlap in the alleles that could be recovered from native and introduced plants, and we found no evidence to suggest hybridization between subspecies, even in localities where native and introduced plants grow in close proximity. Overall, we found greater genetic variation in plants of the introduced subspecies relative to the native subspecies, and we observed some geographic patterns of allelic diversity.

Clonality and genetic diversity in the eastern North American spring ephemeral Erythronium americanum Ker-Gawl. (American Trout Lily)1,2

As an adaptation to unpredictable light and pollination levels in the deciduous forest, clonality in early spring flowering plants enables continued population persistence but can have profound impacts on the population genetic structure. To gain a better understanding of clonal diversity in the common spring ephemeral herb Erythronium americanum (American Trout Lily) and its relationship to population size, five populations were examined using microsatellite markers. In general, levels of genetic diversity were low, with larger and more dense populations of E. americanum exhibiting less allelic diversity than smaller and less dense populations. Over half (68.56%) of sampled individuals overall were clones, with populations containing 2–21 clones and larger populations possessing the fewest number of clones. Several factors may contribute to these patterns, including geitonogamy, genetic drift, and intraspecific competition. The patterns of allelic diversity across the range indicate two distinct genetic clusters of E. americanum with northern and southern groups separated at the Ohio River valley.

Allelic diversity and selection at the MHC class I and class II in a bottlenecked bird of prey, the White-tailed Eagle

BackgroundGenes of the Major Histocompatibility Complex (MHC) are essential for adaptive immune response in vertebrates, as they encode receptors that recognize peptides derived from the processing of intracellular (MHC class I) and extracellular (MHC class II) pathogens. High MHC diversity in natural populations is primarily generated and maintained by pathogen-mediated diversifying and balancing selection. It is, however, debated whether selection at the MHC can counterbalance the effects of drift in bottlenecked populations. The aim of this study was to assess allelic diversity of MHC genes in a recently bottlenecked bird of prey, the White-tailed Eagle Haliaeetus albicilla, as well as to compare mechanisms that shaped the evolution of MHC class I and class II in this species.ResultsWe showed that significant levels of MHC diversity were retained in the core Central European (Polish) population of White-tailed Eagles. Ten MHC class I and 17 MHC class II alleles were recovered in total and individual birds showed high average MHC diversity (3.80 and 6.48 MHC class I and class II alleles per individual, respectively). Distribution of alleles within individuals provided evidence for the presence of at least three class I and five class II loci the White-tailed Eagle, which suggests recent duplication events. MHC class II showed greater sequence polymorphism than MHC class I and there was much stronger signature of diversifying selection acting on MHC class II than class I. Phylogenetic analysis provided evidence for trans-species similarity of class II, but not class I, sequences, which is likely consistent with stronger balancing selection at MHC class II.ConclusionsRelatively high MHC diversity retained in the White-tailed Eagles from northern Poland reinforces high conservation value of local eagle populations. At the same time, our study is the first to demonstrate contrasting patterns of allelic diversity and selection at MHC class I and class II in an accipitrid species, supporting the hypothesis that different mechanisms can shape evolutionary trajectories of MHC class I and class II genes.

Genetic diversity in natural populations of Hancornia speciosa Gomes: Implications for conservation of genetic resources

ABSTRACT The continuing fragmentation of forests has been a threat to the maintenance of genetic resources. Genetic diversity is fundamental to the survival of species in natural environments in the long term, as well as being the basis for genetic improvement. The objective of this study was to evaluate the genetic diversity in natural populations of Hancornia speciosa and to contribute to the development of conservation strategies. We sampled 105 individuals of H. speciosa, distributed in seven populations. The ISSR (Inter-Simple Sequence Repeat) markers provided 70 loci, of which 81% were polymorphic. The mean genetic diversity of Nei (h) was 0.19, and the Shannon index (I) was 0.27. The h and I diversity indices ranged respectively from 0.16 to 0.24 in the PAD (Parque das Dunas) population and from 0.21 to 0.29 in MAC (Macaíba) population. Resulting from a Bayesian analysis, the genotypes were divided into four groups (K = 4). The allelic diversity patterns observed indicated the occurrence of the genetic bottleneck in all populations, according to the stepwise mutation model (SMM). The infinite allele model (IAM) revealed an imbalance between mutation and genetic drift only in the PAD population. Genetic conservation strategies for H. speciosa should cover each genetic group that was differentially structured. We recommend in situ conservation and the creation of germplasm banks, especially with the PAD population which demonstrated the lower genetic diversity and decreased effective population size according to the two mutational models.

Patterns of Allelic Diversity in Spring Wheat Populations by SSR-Markers

Precise assessment of diversity in available breeding germplasm helps to preempt epidemics and abrupt environmental changes. Spring wheat germplasm consisting of 84 accessions including cultivars, breeding lines and landraces from various origins was scanned with 44 SSRs. For allele frequencies, allelic patterns, heterozygosity and polymorphism the selected population was divided in three subpopulations: (i) pre-green revolution (pre-1965), (ii) post-green revolution (post-1965), (iii) post-veery (post-2000). Alleles produced in pre-1965, post-1965 and post-2000 subpopulations were 115, 144 and 131, respectively. Mean PIC values for pre-1965, post-1965 and post-2000 subpopulations were 0.48, 0.52 and 051, respectively. Allelic patterns showed no locally common alleles in any of the subpopulation. The pre-1965 subpopulation had also no private allele, however, average number of private alleles decreased from post-1965 to post-2000 subpopulation. In case of effective alleles and Shannon’s information index trend was increasing from pre-1965 to post-1965 and then decreasing from post-1965 to post-2000. The decreasing trend alarms the reduced genetic diversity in wheat varieties developed after 2000. PCA and cluster analysis didn’t clearly differentiated subpopulations, though pre-1965 genotypes showed higher genetic distance from post-1965 and post-2000 subpopulations. The decreasing measures of genetic diversity in post-2000 wheat genotypes should be a concern for wheat breeders, therefore, all sources of broadening genetic diversity should be exploited.

Genetic Evidence for the Introduction of Rhagoletis pomonella (Diptera: Tephritidae) into the Northwestern United States.

The apple maggot fly, Rhagoletis pomonella Walsh (Diptera: Tephritidae), is a serious quarantine pest in the apple-growing regions of central Washington and Oregon. The fly is believed to have been introduced into the Pacific Northwest via the transport of larval-infested apples near Portland, Oregon, within the last 40 yr. However, R. pomonella also attacks native black hawthorn, Crataegus douglasii Lindley (Rosales: Rosaceae), and introduced ornamental hawthorn, Crataegus monogyna Jacquin, in the region. It is, therefore, possible that R. pomonella was not introduced but has always been present on black hawthorn. If true, then the fly may have independently shifted from hawthorn onto apple in the Pacific Northwest within the last 40 yr after apples were introduced. Here, we test the introduction hypothesis through a microsatellite genetic survey of 10 R. pomonella sites in Washington and 5 in the eastern United States, as well as a comparison to patterns of genetic variation between populations of Rhagoletis cingulata Loew and Rhagoletis indifferens Curran, two sister species of cherry-infesting flies known to be native to the eastern and western United States, respectively. We report results based on genetic distance networks, patterns of allelic variation, and estimated times of population divergence that are consistent with the introduction hypothesis for R. pomonella. The results have important implications for R. pomonella management, suggesting that black hawthorn-infesting flies near commercial apple-growing regions of central Washington may harbor sufficient variation to utilize apple as an alternate host, urging careful monitoring, and possible removal of hawthorn trees near orchards.

Negligence in the Atlantic forest, northern Brazil: a case study of an endangered orchid

Currently, many Brazilian orchids are threatened with extinction resulting from habitat loss and intense harvesting pressure stemming from their value as ornamental plants. Therefore, the genetic diversity in remaining populations is fundamental to the survival of these species in natural environments. In order to inform conservation strategies, this study evaluated the genetic diversity and structure of Cattleya granulosa populations. The sample consisted of 151 individuals from 12 populations in the Atlantic Forest, northeastern Brazil, evaluated using 91 ISSR markers. Genetic variability was assessed through molecular variance, diversity indexes, clusters of genotypes through Bayesian analysis, and tests for genetic bottlenecks. From all polymorphic loci, genetic diversity (HE) varied between 0.210 and 0.321 and the Shannon index ranged from 0.323 and 0.472. Significant genetic differentiation between populations (ΦST = 0.391; P < 0.0001) resulted in the division of the populations into five groups based on the log-likelihood Bayesian analysis. We found significant positive correlation between geographical and genetic distances between populations (r = 0.794; P = 0.017), indicating isolation by distance. Patterns of allelic diversity within populations suggest the occurrence of bottlenecks in most C. granulosa populations (n = 8). Therefore, in order to maintain the genetic diversity of the species, the conservation of spatially distant groups is necessary.

DIVERSIDADE GENÉTICA E SELEÇÃO DE INICIADORES ISSR EM UMA POPULAÇÃO NATURAL DE MANGABA (Hancornia speciosa Gomes) (APOCYNACEAE)

RESUMO O conhecimento da diversidade genética de espécies nativas é de grande valia quando se objetiva o melhoramento e a conservação de populações naturais. Neste sentido, o objetivodeste trabalho foi selecionar iniciadores ISSR (inter repetições de sequências simples) para Hancornia speciosa (Apocynaceae), assim como quantificar a variabilidade genética em uma população natural. Foramamostrados 15 indivíduos de uma população localizada em Natal-RN. Amostras de caule foram coletadas para a posterior extração do DNA. DNA. Para a seleção, 19 primers ISSR foram testados, dos quais seis foram eficientes, apresentando locos nítidos e em maior número (UBC 808; UBC 810; UBC 826; UBC 827; UBC 841 e UBC 842), totalizando 63 locos. Desses, apenas 30 (47,62%) apresentaram polimorfismo. O valor de PIC (conteúdo de informações polimórficas) para os primers selecionados atingiu a média de 0,37, variando de 0,26 a 0,44. A diversidade genética foi considerada baixa dentro da população, com o número de alelos observados (na =1,48), número de alelos efetivos (ne = 1,32), índice de diversidade de Nei (He = 0,18) e índice de Shannon (I = 0,26). Os padrões de diversidade alélica encontrados indicam a ocorrência de um gargalo populacional recente. A utilização de marcadores ISSR para Hancornia speciosa mostrou-se eficaz para a quantificação da diversidade genética dos indivíduos, servindo como aporte para estratégias e planos que visem à conservação e à manutenção da espécie.

Population genetic structure of the ectomycorrhizal fungus Laccaria sp . A resembles that of its host tree Nothofagus cunninghamii

The ectomycorrhizal fungus Laccaria sp. A is restricted to temperate rainforest of southeast Australia, associated with its host tree Nothofagus cunninghamii. Eight mitochondrial microsatellite markers were used to investigate the population genetic structure of L. sp. A across its distribution in Tasmania and Victoria. The highest allelic diversity was found in Tasmania, which appeared to contain a panmictic population, whereas the more fragmented Victorian populations were characterized by low allelic diversity and differentiation between east and west. There is evidence of glacial refugia in the west and the northeast of Tasmania, and in Victoria in the Otway Ranges and Central Highlands, with postglacial migration into the Strzelecki Ranges. Narrow host-specificity may have contributed to the presence of population structure in this fungus. Allelic diversity patterns in L. sp. A are largely congruent with diversity patterns already established in populations of its host, N. cunninghamii.

Molecular genetic characterization of Central European soybean breeding germplasm

AbstractSoybean is the most important oilseed and protein crop worldwide, but in Europe, the acreage is comparably low. Thus, Europe strongly depends on imports of soy products and consequently recent efforts aim at expanding the acreage of soybean, particularly in Central Europe. The aim of this study was, therefore, to assist these breeding efforts by characterizing Central European soybean germplasm, employing a genotyping‐by‐sequencing approach yielding 7741 genomewide distributed markers. Our analysis on genetic diversity and population structure revealed that the Central European lines are most closely related to Swiss and Canadian lines and somewhat more distant from the investigated Chinese and US lines. In addition, we analysed patterns of allelic diversity and the extent of linkage disequilibrium. Collectively, our results can assist breeding of Central European soybean and suggest that further progress can be made by crosses among adapted material but a long‐term success also requires introgression of alleles from non‐European germplasm to further broaden the genetic diversity and incorporate novel traits.

Drift rather than selection dominates MHC class II allelic diversity patterns at the biogeographical range scale in natterjack toads Bufo calamita.

Study of major histocompatibility complex (MHC) loci has gained great popularity in recent years, partly due to their function in protecting vertebrates from infections. This is of particular interest in amphibians on account of major threats many species face from emergent diseases such as chytridiomycosis. In this study we compare levels of diversity in an expressed MHC class II locus with neutral genetic diversity at microsatellite loci in natterjack toad (Bufo (Epidalea) calamita) populations across the whole of the species’ biogeographical range. Variation at both classes of loci was high in the glacial refugium areas (REF) and much lower in postglacial expansion areas (PGE), especially in range edge populations. Although there was clear evidence that the MHC locus was influenced by positive selection in the past, congruence with the neutral markers suggested that historical demographic events were the main force shaping MHC variation in the PGE area. Both neutral and adaptive genetic variation declined with distance from glacial refugia. Nevertheless, there were also some indications from differential isolation by distance and allele abundance patterns that weak effects of selection have been superimposed on the main drift effect in the PGE zone.

Geographic patterns of Got1, Got2, Got3 and Est2 enzyme-encoding genes allelic variation in Aegilops tauschii Coss.

Geographic patterns of Got1, Got2, Got3 and Est2 enzyme-encoding genes allelic variation were investigated among 322 accessions of Aegilops tauschii Coss., 161 accessions of ssp. tauschii and 161 accessions of ssp. strangulata, representing all the species area. It was found that: (1) in the two ecologically different subspecies, ssp. tauschii and ssp. strangulata, the patterns of allelic variation of the four genes differ greatly; (2) the same allozymes have originated several times independently in different Ae. tauschii local populations; (3) allelic variation of Got1, Got3 and Est2 in ssp. strangulata corresponds to climatic conditions. The data obtained reflected that Ae. tauschii has been inhabiting its area from ancient times; and allelic variation patterns of Got1, Got2, Got3, Est2 loci were mostly formed by natural selection. Further investigations of these loci with molecular genetic methods are prospective for understanding the peculiarities of Ae. tauschii evolution.

Retention of agronomically important variation in germplasm core collections: implications for allele mining

The primary targets of allele mining efforts are loci of agronomic importance. Agronomic loci typically exhibit patterns of allelic diversity that are consistent with a history of natural or artificial selection. Natural or artificial selection causes the distribution of genetic diversity at such loci to deviate substantially from the pattern found at neutral loci. The germplasm utilized for allele mining should contain maximum allelic variation at loci of interest, in the smallest possible number of samples. We show that the popular core collection assembly procedure "M" (marker allele richness), which leverages variation at neutral loci, performs worse than random assembly for retaining variation at a locus of agronomic importance in sugar beet (Beta vulgaris L. subsp. vulgaris) that is under selection. We present a corrected procedure ("M+") that outperforms M. An extensive coalescent simulation was performed to demonstrate more generally the retention of neutral versus selected allelic variation in core subsets assembled with M+. A negative correlation in level of allelic diversity between neutral and selected loci was observed in 42% of simulated data sets. When core collection assembly is guided by neutral marker loci, as is the current common practice, enhanced allelic variation at agronomically important loci should not necessarily be expected.

The structure of allelic diversity in the presence of purifying selection

In the absence of selection, the structure of equilibrium allelic diversity is described by the elegant sampling formula of Ewens. This formula has helped to shape our expectations of empirical patterns of molecular variation. Along with coalescent theory, it provides statistical techniques for rejecting the null model of neutrality. However, we still do not fully understand the statistics of the allelic diversity expected in the presence of natural selection. Earlier work has described the effects of strongly deleterious mutations linked to many neutral sites, and allelic variation in models where offspring fitness is unrelated to parental fitness, but it has proven difficult to understand allelic diversity in the presence of purifying selection at many linked sites. Here, we study the population genetics of infinitely many perfectly linked sites, some neutral and some deleterious. Our approach is based on studying the lineage structure within each class of individuals of similar fitness in the deleterious mutation-selection balance. Consistent with previous observations, we find that for moderate and weak selection pressures, the patterns of allelic diversity cannot be described by a neutral model for any choice of the effective population site. We compute precisely how purifying selection at many linked sites distorts the patterns of allelic diversity, by developing expressions for the likelihood of any configuration of allelic types in a sample analogous to the Ewens sampling formula.

The Genetic Mosaic ofIrisSeriesHexagonaein Florida: Inferences on the Holocene History of the Louisiana Irises and Anthropogenic Effects on Their Distribution

Iris series Hexagonae is a small, monophyletic complex of five species and associated hybrid populations, popularly known as the “Louisiana irises.” The Hexagonae alliance of Iris has been recognized as a textbook case of introgressive hybridization on the basis of numerous studies in Louisiana. In light of field observations of phenotypic heterogeneity among Florida populations, we investigated genetic variation within and among 37 populations of Iris series Hexagonae from Florida, seven from Louisiana, and one from Texas with 19 microsatellite loci. We wished to test the hypothesis that the Lake Wales Ridge and Polk Uplands of Florida, which remained above sea level maxima during the early Pleistocene, might be the sites of origin for all extant iris populations in Florida and possibly the series Hexagonae in its entirety. We also hypothesized that phenotypic heterogeneity observed in Florida iris would be reflected in patterns of allelic diversity. Finally, we wished to seek evidence of Native American influence on the present distribution of iris in Florida. Most populations display clonality, including three populations that are predominantly one or two clones. Levels of gene diversity are high and the populations are highly differentiated, although 19 exhibit significant homozygote excess. The majority of genetic variation occurs within populations. Genetic distance resolves three large clusters. One unites the southernmost populations with those from north-central Gulf coastal Florida. A second encompasses southwest Gulf coastal populations. If three compound repeat loci are dropped from the data, these two clusters are united. The third group joins populations derived from the Central Highlands of peninsular Florida, which are genetically isolated from all other peninsular Florida populations, with populations of Iris fulva, Iris brevicaulis, Iris giganticaerulea, and Iris hexagona s.s., supporting recognition of the latter two as distinct species. Model-based Bayesian clustering supports high population differentiation () and isolation of Central Highlands populations, and it resolves the terminal clusters of the genetic distance topology. Isolation by distance is significant yet weak because of genetic distance relationships that contradict biogeographic expectations. We propose that the Lake Wales Ridge and Polk Uplands, which constituted the Wicomico shoreline during an early Pleistocene interglacial inundation, functioned as refugia for series Hexagonae. Moreover, we suggest that Florida iris populations occupying high, dry habitats close to the Central Highlands ridges represent relicts of once-larger populations that adapted to the more xeric condition during the last glacial maximum. These populations have a distinctive floral phenotype and are related to species of Hexagonae iris outside of peninsular Florida. Excessively clonal populations may have been deliberately cultivated by Native Americans, which may also have influenced the connection between southern and north-central Florida populations, but there is insufficient evidence to validate this hypothesis. Many populations test positively for recent bottlenecks, which we attribute primarily to founder effects, given the low migration rates of the species and the high degree of population differentiation as well as the Holocene geological history of the Florida peninsula. We present evidence of peripatric divergence in series Hexagonae iris and suggest that this may function as a significant generator of species diversity in the group.

Allelic polymorphism of human FcγRIIA-H/R131 receptor in American tegumentary leishmaniasis

FcγRIIA binding to IgG subclasses with different levels of affinity is influenced by the polymorphism in the gene that encodes this receptor. The substitution of arginine (R) for histidine (H) in the 131 position defines three allelic patterns, H/H, R/R, and H/R, resulting in FcγRIIA-H/H131 affinity for IgG2 and higher affinity for IgG3 subclasses. Studies have shown the importance of genetic host factors in leishmaniasis and participation of FcγRs on the macrophage infection by amastigote forms and in the immune response to Leishmania sp. We analysed the influence of allelic diversity patterns of the receptor FcγRIIA on American tegumentary leishmaniasis (ATL). FcγRIIA-H/R131 polymorphism was determined by PCR followed by an allele-specific enzymatic digestion in 88 individuals with ATL and 98 healthy volunteer blood donors (control group). The genotypic and allelic distributions of FcγRIIA-H/R131 were similar among the studied groups as well in mild and severe clinical forms of ATL. Our results suggest no association between this allelic polymorphism and susceptibility or resistance to ATL, neither influencing the development of different clinical forms of this illness.

Divergent patterns of allelic diversity from similar origins: the case of oilseed rape (Brassica napusL.) in China and Australia

Oilseed rape (Brassica napus) in Australia and China have similar origins, with introductions from Europe, Canada, and Japan in the mid 20th century, and there has been some interchange of germplasm between China and Australia since that time. Allelic diversity of 72 B. napus genotypes representing contemporary germplasm in Australia and China, including samples from India, Europe, and Canada, was characterized by 55 polymorphic simple sequence repeat (SSR) markers spanning the entire B. napus genome. Hierarchical clustering and two-dimensional multidimensional scaling identified a Chinese group (China-1) that was separated from "mixed group" of Australian, Chinese (China-2), European, and Canadian lines. A small group from India was distinctly separated from all other B. napus genotypes. Chinese genotypes, especially in the China-1 group, have inherited unique alleles from interspecific crossing, primarily with B. rapa, and the China-2 group has many alleles in common with Australian genotypes. The concept of "private alleles" is introduced to describe both the greater genetic diversity and the genetic distinctiveness of Chinese germplasm, compared with Australian germplasm, after 50 years of breeding from similar origins.

Allelic diversity associated with aridity gradient in wild emmer wheat populations

The association between allelic diversity and ecogeographical variables was studied in natural populations of wild emmer wheat [Triticum turgidum ssp. dicoccoides (Körn.) Thell.], the tetraploid progenitor of cultivated wheat. Patterns of allelic diversity in 54 microsatellite loci were analyzed in a collection of 145 wild emmer wheat accessions representing 25 populations that were sampled across naturally occurring aridity gradient in Israel and surrounding regions. The obtained results revealed that 56% of the genetic variation resided among accessions within populations, while only 44% of the variation resided between populations. An unweighted pair-group method analysis (UPGMA) tree constructed based on the microsatellite allelic diversity divided the 25 populations into six major groups. Several groups were comprised of populations that were collected in ecologically similar but geographically remote habitats. Furthermore, genetic differentiation between populations was independent of the geographical distances. An interesting evolutionary phenomenon is highlighted by the unimodal relationship between allelic diversity and annual rainfall (r = 0.74, P < 0.0002), indicating higher allelic diversity in populations originated from habitats with intermediate environmental stress (i.e. rainfall 350-550 mm year(-1)). These results show for the first time that the 'intermediate-disturbance hypothesis', explaining biological diversity at the ecosystem level, also dominates the genetic diversity within a single species, the lowest hierarchical element of the biological diversity.

Genetic variation and structure in the expanding moss Pogonatum dentatum (Polytrichaceae) in its area of origin and in a recently colonized area

Genetic variation in the expanding moss species Pogonatum dentatum was studied using intersimple sequence repeat (ISSR) markers. The genetic consequences of range expansion were studied by comparing source populations in a mountain area with populations from a recently colonized lowland area in Sweden. Indices of genetic variation show slightly lower number of alleles per locus in the lowlands and a similar gene diversity in both areas. Three of four lowland populations had evidence of a recently passed bottleneck. Considerably higher haplotype diversity was found in the recently colonized lowlands compared to source populations in the mountains. Patterns of allelic diversity suggest that P. dentatum experiences loss of genetic variation through founder effects and genetic drift when expanding its distribution range. Higher haplotypic diversity, less linkage disequilibrium, and fewer compatible loci indicate that sexual recombination is relatively more important in the lowlands compared to the mountains. A likely explanation is higher success of establishment from spores in the lowlands, while clonal propagation predominates in the mountains. Less genetic differentiation among lowland populations indicates more gene flow in the lowland area, involving more spores and/or fragments moving among populations.