Allozyme Variation and Phylogenetic Relationships in Picea jezoensis (Pinaceae) Populations of the Russian Far East

Genetic variation and differentiation of 12 populations of Picea jezoensis from the Russian Far East were studied using 20 allozyme loci. The mean number of alleles per locus was 2.63, the percent of polymorphic loci was 88.1%, the observed heterozygosity was 0.181, and the mean value of expected heterozygosity amounted to 0.189. The values of expected heterozygosity of the northern and central mainland populations were higher than in the southern part of the natural range. A significant bias of Hardy–Weinberg heterozygosity to equilibrium heterozygosity (H_eq) suggests that most of the mainland populations have recently experienced a severe expansion in population size while populations from Kamchatka Peninsula have undergone a reduction in population size. Unbiased Nei’s genetic distance values were low within and between the mainland and Sakhalin Island populations (D_N=0.008). The largest values (D_N=0.063) were found between the mainland/Sakhalin and Kamchatka Peninsula populations. Based on genetic distance, P. jezoensis and P. kamtschatkensis could be considered as distinct taxa, but P. ajanensis, P. microsperma, and P. komarovii do not warrant taxonomic recognition.     

Genetic variation and population structure ofCarex breviculmis (Cyperaceae) in Korea

We determined the genetic diversity and population structures ofCarex breviculmis (Cyperaceae) populations in Korea, using genetic variations at 23 allozyme loci.C. breviculmis is a long-lived herbaceous species that is widely distributed in eastern Asia. A high level of genetic variation was found in 15 populations. Twelve enzymes revealed 23 loci, of which 11 were polymorphic (47.8%). Genetic diversity at the speciesand population levels were 0.174 and 0.146, respectively. Total genetic diversity (H_T = 0.363) and within-population genetic diversity (H_s = 0.346) were high, whereas the extent of the population divergence was relatively low (G_ST = 0.063). Deviation from random mating (Fis) within the 15 populations was 0.206. An indirect estimate of the number of migrants per generation(Nm = 3.69) indicated that gene flow was extensive among Korean populations of this species. Analysis of fixation indices revealed a substantial heterozygote deficiency in some populations and at some loci. Genetic identity between popu-lations was high, exceeding 0.956.     

Allozyme variation in the diploid (A genome) populations ofScilla scilloides (Hyacinthaceae)

Allozyme variation at eleven loci encoding seven enzyme systems were examined in 20 populations of diploid (genome AA, 2n = 16)Scilla scilloides in China. In comparison with the average species of seed plants studied, populations of this species display a high amount of genetic variation (A = 2.0, P = 58.6%, H_o = 0.172, and H_e = 0.185). Allozyme variation pattern revealed predominant outcrossing within populations and considerable differentiation (F_ST = 0.314) among populations as well as between the subtropic and temperate regions. The wide distribution, long existence and outcrossing are presumably the main factors responsible for the high genetic diversity within populations. But the gravity dispersal of seeds and pollination by small insects set limits to the increase of genetic variation within populations and promote differentiation between populations and regions. In addition, allozyme variation does not distinguishS. scilloides var.albo-viridis and suggests that subtropic populations may be considered as a genetic entity.     

Evolutionary implications of allozyme and RAPD variation in diploid populations of dioecious buffalograss Buchloë dactyloides

Buffalograss, Buchloë dactyloides, is widely distributed throughout the Great Plains of North America, where it is an important species for rangeland forage and soil conservation. The species consists of two widespread polyploid races, with narrowly endemic diploid populations known from two regions: central Mexico and Gulf Coast Texas. We describe and compare the patterns of allozyme and RAPD variation in the two diploid races, using a set of 48 individuals from Texas and Mexico (four population samples of 12 individuals each). Twelve of 22 allozyme loci were polymorphic, exhibiting 35 alleles, while seven 10-mer RAPD primers revealed 98 polymorphic bands. Strong regional differences were detected in the extent of allozyme polymorphism: Mexican populations exhibited more internal gene diversity (H_e= 0.20, 0.19) than did the Texan populations (H_e= 0.08, 0.06), although the number of RAPD bands in Texas (n= 62) was only marginally smaller than in Mexico (n= 68). F-statistics for the allozyme data, averaged over loci, revealed strong regional differentiation (mean F_RT=+ 0.30), as well as some differentiation among populations within regions (mean F_PR=+ 0.09). In order to describe and compare the partitioning of genetic variation for multiple allozyme and RAPD loci, we performed an Analysis of Molecular Variance (AMOVA). AMOVA for both allozyme and RAPD data revealed similar qualitative patterns: large regional differences and smaller (but significant) population differences within regions. RAPDs revealed greater variation among regions (58.4% of total variance) than allozymes (45.2%), but less variation among individuals within populations (31.9% for RAPDs vs. 45.2% for allozymes); the proportion of genetic variance among populations within regions was similar (9.7% for RAPDs vs. 9.6% for allozymes). Despite this large-scale concordance of allozyme and RAPD variation patterns, multiple correlation Mantel techniques revealed that the correlations were low on an individual by individual basis. Our findings of strong regional differences among the diploid races will facilitate further study of polyploid evolution in buffalograss.     

Genetic diversity and population structure of<Emphasis Type="Italic">Kalopanax pictus</Emphasis> (Araliaceae)

Kalopanax pictus (Araliaceae) is a long-lived woody species primarily distributed throughout East Asia. This species is regarded as medically and ecologically important in Korea. We used starch gel electrophoresis to investigate its allozyme variation and genetic structure in samples from Korean populations (both natural and cultivated). Our analysis of 10 enzymes revealed 18 loci, of which 10 were polymorphic (35.6%). Genetic diversity at the species and population levels were 0.200 and 0.149, respectively, with the mean for cultivated populations (0.124) being lower than for natural (wild) populations (0.181). Asexual and sexual reproduction modes, perennial habitat, and longevity all were possible factors contributing to such high diversity. An indirect estimate of the number of migrants per generation (N _m = 1.08) indicated that gene flow was not extensive among these Korean populations. Therefore, we suggest that geographical distance as well as reproductive isolation between wild and cultivated plants may play roles in shaping the population structure of this species.     

A cline of allozyme variation inAbies mariesii

Genetic variation at 22 allozyme loci was examined for 1,003 trees from 11 isolated natural populations ofAbies mariesii covering all except the southernmost region of its geographic range. Genetic diversity within species (H _es=0.063) was low compared to many other long-lived woody species. Most of the genetic variation is found within populations (G _ST=0.144) despite their isolated distribution. Genetic distance between populations was positively correlated with geographic distance. Genetic diversity within populations was generally low (meanH _ep=0.054), but varied across populations in a clinal fashion such that genetic variation decreased with increasing latitude. These genetic characteristics may reflect the distribution history of this species.     

Genetic Variation in the Endangered Endemic Species Cycas fairylakea (Cycadaceae) in China and Implications for Conservation

Cycas fairylakea is an endangered endemic species in China. Genetic diversity within and among four natural populations of this species in China was investigated using amplified fragments length polymorphism (AFLP). A moderate to low level of intraspecific genetic diversity was detected in this species (at population level: P = 39.57 %, H₀ = 0.244; at species level: P = 60.22%, H₀ = 0.356). The among-population component accounted for, respectively, 25.7 and 31.5% of the genetic variation, according to AMOVA and Shannon’s index, indicating most of the genetic variation was found between individuals within populations. All four populations have opposite pyramid age structure, and few coning individuals, which is still decreasing. Possibly because of habitat degradation and environmental pollution, plant diseases and insect pests in the populations were extremely serious, suggesting that the main factors threatening the survival of C. fairylakea populations were not genetic variation, but human activities and the breeding system of this species.     

Population Genetic structure ofPotentilla fragariodes var.major (Rosaceae) in Korea

The genetic diversity and population structure of eighteenPotentilla fragariodes var.major (Rosaceae) populations in Korea were determined using genetic variations at 22 allozyme loci. The percent of polymorphic loci within the enzymes was 66.7%. Genetic diversity at the species level and at the population level was high (Hes = 0.203; Hep = 0.185, respectively), whereas the extent of the population divergence was relatively low (G_ST = 0.069). F_IS, a measure of the deviation from random mating within the 18 populations, was 0.075. An indirect estimate of the number of migrants per generation (Nm = 3.36) indicated that gene flow was high among Korean populations of the species. In addition, analysis of fixation indices revealed a slight heterozygote deficiency in some populations and at some loci. Wide geographic ranges, perennial herbaceous nature and the persistence of multiple generations are associated with the high level of genetic variation. AlthoughP. fragariodes var.major usually propagated by asexually-produced ramets, we could not rule out the possibility that sexual reproduction occurred at a low rate because each ramet may produce terminal flowers. Mean genetic identity between populations was 0.983. It is highly probable that directional movement toward genetic uniformity in a relatively homogeneous habitat operates among Korean populations ofP. fragariodes var.major.     

Allozyme Diversity and Population Genetic Structure of Pinus densata Master in Northwestern Yunnan, China

We investigated the levels and patterns of genetic diversity of Pinus densata Master in Yunnan. Horizontal starch-gel electrophoresis was performed on macrogametophytes collected from nine populations in northwestern Yunnan, China. Compared with other gymnosperm species, P. densata has higher mean values for all measures of genetic diversity. Allozyme polymorphism (0.99 criterion) was 97.0% and 71.4% at the species and population levels, respectively. The average number of alleles per locus was 3.1 and 2.0 at the species and population levels. Mean expected heterozygosity was substantially higher in P. densata than average values investigated for other gymnosperms both at the population (H _ep = 0.174±0.031) and at the species (H _es = 0.190) levels. Of the total genetic variation, less than 12% was partitioned among populations (G _ST = 0.112). Our allozyme survey supports the suggestion that the observed higher diversity in P. densata may be attributed partly to its hybrid origin between two genetically distinct species, P. yunnanensis and P. tabulaeformis. In addition, we suggest that introgression would give rise to the increase in genetic diversity occurring in P. densata.     

Genetic variation and island biogreography: Microsatellite and mitochondrial DNA variation in island populations of the Australian bush rat, Rattus fuscipes greyii

To understand the impact of various factors on the maintenance of genetic variation in natural populations, we need to focus on situations where at least some of these factors are removed or controlled. In this study, we used highly variable, presumably neutral, microsatellite and mtDNA markers to assess the nature of genetic variation in 14 island and two mainland populations of the Australian bush rat, where there is no migration between islands. Thus we are controlling for selection and gene flow. Both marker sets revealed low levels of diversity within the small island populations and extreme differentiation between populations. For six microsatellite loci, all of the small island populations had less genetic variation than the mainland populations; reduction in allelic diversity was more pronounced than loss of heterozygosity. Kangaroo Island, the large island population, had similar levels of diversity to the mainland populations. A 442 base pair (bp) section of the mtDNA control region was screened for variation by outgroup heteroduplex analysis/temperature gradient gel electrophoresis (OHA/TGGE). Only three of the 13 small island populations showed haplotypic diversity: Gambier (2), Waldegrave (2), and Eyere (3). The level of haplotypic diversity in the small island populations was similar to that on the mainland, most likely reflecting a recent population bottleneck on the mainland. In contrast, Kangaroo Island had 9 mtDNA haplotypes. The dominant factor influencing genetic diversity on the islands was island size. No correlation was detected between genetic diversity and the time since isolation or distance form the mainland. The combination of genetic drift within and complete isolation among the small island populations has resulted in rapid and extreme population divergence. Population pair-wise comparisons of allele frequency distributions showed significant differences for all populations for all loci (F _st = 0.11–0.84, R _st = 0.07–0.99). For the mtDNA control region, 92.6% of variation was apportioned between populations; only the Pearson islands shared a haplotype. Mantel tests of pair-wise genetic distance with pair-wise geographic distance showed no significant geographical clustering of haplotypes. However, population substructuring was detected within populations where sampling was conducted over a broader geographical range, as indicated by departures from Hardy-Weinberg equilibrium. Thus substructuring in the ancestral population cannot be ruled out. The dominant evolutionary forces on the islands, after the initial founder event, are stochastic population processes such as genetic drift and mutation. This revised version was published online in July 2006 with corrections to the Cover Date.     

Population genetic structure of island and mainland populations of the quokka, Setonix brachyurus (Macropodidae): a comparison of AFLP and microsatellite markers

Translocation and reintroduction are important tools for the conservation or recovery of species threatened with extinction in the wild. However, an understanding of the potential genetic consequences of mixing populations requires an understanding of the genetic variation within, and similarities among, donor and recipient populations. Genetic diversity was measured using two independent marker systems (microsatellites and AFLPs) for one island and four small remnant mainland populations of Setonix brachyurus, a threatened medium sized macropod restricted to fragmented habitat remnants and two off-shore islands in southwest Australia. Microsatellite diversity in the island population (R _s = 3.2, H _e = 71%) was similar to, or greater than, all mainland populations (R _s = 2.1–3.9, H _e = 34-71%). In contrast, AFLP diversity was significantly lower in the island population (PPL = 20.5; H _j = 0.118) compared to all mainland populations (mean PPL = 79.5–89.7; mean H _j = 0.23–0.29). Microsatellites differentiated all (mainland and island) populations from each other. However, AFLP only differentiated the island population from the mainland populations—all mainland populations were not significantly differentiated from each other for this marker. Given a known time since isolation of the island population from the mainland (6,000 years ago), and an overall more conservative rate of evolution of AFLP markers, our results are consistent with mainland populations fragmenting thousands of years ago (but <6,000 years), probably as a consequence of reduced rainfall and the constriction of the preferred mesic habitat of quokkas. Our results also support a recent history of severe population bottlenecks in mainland populations, and a long history of bottlenecks of the island population, but reflect a recent explosion in numbers since European occupation of the island. Our results indicate that translocation of island populations to supplement mainland populations would introduce genetically markedly differentiated, and possibly maladapted, individuals.     

Maintenance of Phenotypic and Genotypic Diversity in Managed Populations of Stenocereus Stellatus (Cactaceae) by Indigenous Peoples in Central Mexico

The columnar cactus Stenocereus stellatus is used in Central Mexico for its edible fruits which are harvested in wild, managed in situ and cultivated populations. Management in situ of wild populations is conducted by selectively sparing and enhancing the abundance of plants with desirable phenotypes when fields are cleared for agricultural use. Cultivation of desirable phenotypes is carried out by vegetative propagation in homegardens. Effects of human management on morphological and genetic variation of S. stellatus were analyzed by comparing morphological diversity indices (MD, based on Simpson’s index) and expected (H_e) heterozygosity indices from allozyme analysis, in wild, managed in situ, and cultivated populations from La Mixteca and the Tehuacán Valley regions. Morphological diversity was similar among regions, but populations from the wetter La Mixteca region averaged higher genetic variation (H_e = 0.279) than populations from Tehuacán (H_e = 0.265). On average, populations manipulated by people had higher levels of variation (MD = 0.479 ± 0.012, H_e = 0.289 in cultivated populations; MD = 0.461 ± 0.014, H_e = 0.270 in managed in situ populations) than wild populations (MD = 0.408 ± 0.017, H_e = 0.253), which is apparently due to a continual introduction and replacement of plant materials in the manipulated populations. The results illustrate that human management may not only maintain but also increase both morphological and genetic diversity of manipulated plant populations in relation to that existing in the wild. Managed in situ and cultivated populations of S. stellatus are important reservoirs of variation, and are crucial for the general maintenance of diversity in wild populations. These populations may play a principal role in designing strategies for the conservation of variation of this cactus.     

Low levels of genetic variation inPhenakospermum guyannense (Strelitziaceae), a widespread bat-pollinated Amazonian herb

Phenakospermum guyannense is a monotypic, arborescent, long-lived monocot that is widespread in Amazonian South America. This outcrossing species is pollinated primarily by phyllostomid bats. Given these life-history characteristics,P. guyannense is expected to exhibit high levels of genetic variation and gene flow. We used isozyme electrophoresis and randomly amplified polymorphic DNA (RAPD) to characterize genetic variation in populations ofP. guyannense from French Guiana. Both measures detected a surprisingly low level of genetic variation, with only five out of twenty (25%) allozyme loci polymorphic (P), 1.35 alleles per locus (A), and an expected heterozygosity (H_e) of 0.090 at the species level. Isozymic genetic variation was even lower within populations (P = 17.5, A = 1.24, H_e = 0.074), and was corroborated by a RAPD assay that used 26 arbitrary primers (P = 3.61, A = 1.04, H_e = 0.014). Although overall levels of variation were low, the detectable variation was distributed as would be expected for an outcrossing species with extensive gene flow (mean G_ST = 0.230). We suspect thatP. guyannense is depauperate in genetic variation because of a series of bottlenecks that affected the species over this portion of its range.     

TEMPORAL ALLOZYME FREQUENCY CHANGES IN DENSITY FLUCTUATING POPULATIONS OF WILLOW GROUSE (LAGOPUS LAGOPUS L.)

The population density of willow grouse (Lagopus lagopus L.) in northern Scandinavia changes in synchrony with the cyclic density variations in populations of microtine rodents. To assess the genetic changes accompanying the variations in population number, allozyme variation was studied at 23 loci in 640 willow grouse, representing four mainland and one island locality sampled during high and low population density. The average heterozygosity (H = 8.3%) and proportion of polymorphic loci (P = 26%) is not lower in willow grouse than in avian species with a more stable demography; the recurrent population density changes do not appear to affect drastically the long term effective population size, presumably because of extensive migration. Significant allele frequency differences were found both between populations and between different density phases. The genetic distance (D; Nei, 1972) was, in about 50% of the cases, larger between two consecutive time periods than between two localities in a certain year. Spatial and temporal allele frequency variation each represented around 3% of the gene diversity. The temporal heterogeneity may be caused by nonrandom sampling of family groups, rather than drift of allele frequencies between generations due to small effective population size, as has been suggested for microtine species.     

Genetic diversity and morphological differentiation in Liatris helleri (Asteraceae), a threatened plant species

Liatris helleri (Asteraceae) is an insect-pollinated herbaceous perennial endemic to several high-elevation sites in the Blue Ridge Mountains of North Carolina. Allozymes were used to describe the genetic diversity and population structure in nine populations of this rare, federally listed species. Differences in leaf morphology were also examined for greenhouse-grown plants representing several populations. The proportion of the total genetic diversity found among populations, as indicated by the allozyme data, was 16%. Higher levels of population differentiation were found for differences in leaf shape; population of origin accounted for 37% of the variation in maximum leaf width, while families within populations accounted for 7%. In contrast to many endemic species, L. helleri maintains fairly high levels of genetic diversity. For the species, the percent polymorphic loci was 87.5, the average number of alleles at variable loci was 3.00 and the gene diversity was 0.276. Mean population values were percent polymorphic loci =58.4, mean number of alleles per polymorphic locus =2.59 and gene diversity =0.219. The estimated gene flow was low (Nm=0.85–1.32) and a relatively high correlation (r=0.55; p<0.005) was found between linear geographic and genetic distance. This suggests that the populations are partially isolated by distance, despite the limited range (<60 km) of the species. We recommend that population distinetiveness be maintained in restoration efforts.     

Genetic Variation in Remnant Populations of the Woolly Spider Monkey (Brachyteles arachnoides)

The muriqui or woolly spider monkey (Brachyteles arachnoids) is an endangered primate endemic to the Atlantic Forest of Brazil, <5% of which remains. The known muriqui population consists of <700 individuals separated into approximately 15 geographically isolated forest fragments. I present data on the distribution of genetic variation within and between two such remnant populations (FE and FBR) and summarize the implications of these results for long-range management of species genetic diversity. Eleven of 32 allozyme loci were polymorphic, representing an overall level of polymorphism of 34.4% and a mean heterozygosity per locus of 11%. Both values are among the highest reported for New World monkeys. Genetic differentiation between the two localities is highly significant (F_ST = 0.413, p < 0.001). Genetic distance between them is an order of magnitude greater than that between other populations of platyrrhine subspecies, but this could be an artifact of the small sample size from FBR. High levels of genetic diversity apparently characteristic of this species persist because (1) fragmentation and size reduction of muriqui populations has occurred very rapidly relative to the muriqui life span—although both polymorphism and heterozygosity were lost between generations in the largest population, the high genetic diversity present in the parent population was still in evidence; and (2) genetic diversity before population fragmentation by human activity was not distributed uniformly throughout the species' historic distribution. Thus, remnant muriqui populations are important genetic reservoirs of alleles that are unique or rare in the species gene pool as a whole. These results emphasize the need for the integration of conservation management efforts throughout the species range.     

Large mainland populations of South Island robins retain greater genetic diversity than offshore island refuges

For conservation purposes islands are considered safe refuges for many species, particularly in regions where introduced predators form a major threat to the native fauna, but island populations are also known to possess low levels of genetic diversity. The New Zealand archipelago provides an ideal system to compare genetic diversity of large mainland populations where introduced predators are common, to that of smaller offshore islands, which serve as predator-free refuges. We assessed microsatellite variation in South Island robins (Petroica australis australis), and compared large mainland, small mainland, natural island and translocated island populations. Large mainland populations exhibited more polymorphic loci and higher number of alleles than small mainland and natural island populations. Genetic variation did not differ between natural and translocated island populations, even though one of the translocated populations was established with five individuals. Hatching failure was recorded in a subset of the populations and found to be significantly higher in translocated populations than in a large mainland population. Significant population differentiation was largely based on heterogeneity in allele frequencies (including fixation of alleles), as few unique alleles were observed. This study shows that large mainland populations retain higher levels of genetic diversity than natural and translocated island populations. It highlights the importance of protecting these mainland populations and using them as a source for new translocations. In the future, these populations may become extremely valuable for species conservation if existing island populations become adversely affected by low levels of genetic variation and do not persist.     

Genetic variation and differentiation in Nordic populations of Elymus alaskanus (Scrib. ex Merr.) Löve (Poaceae)

 To gain information on the extent and nature of genetic variation in Elymus alaskanus, levels and distribution of genetic variation were assessed within and among 13 populations originating from Iceland, Norway, Sweden and Russia using allozymes. The results showed that four (30.7%) of the 13 loci were polymorphic within the species, while the mean percentage of polymorphic loci within the populations was 1.9%. The mean number of alleles per locus for the species was 1.8 and 1.02 across the populations. Genetic diversity at the species level was low (H _es=0.135), and mean population diversity was notably lower (H _ep=0.005). A high degree of genetic differentiation was observed among populations. The salient points emerging from this study are: (1) statistically significant differences were found in allele frequencies among populations for every polymorphic locus (P<0.001), (2) the high mean coefficient of gene differentiation (G _ST) showed that 95% of the total allozyme variation was attributable to differences among populations, and (3) relatively high genetic distances between the populations were obtained (mean D=0.16). The Norwegian populations had the highest genetic diversity as compared with the other populations. Geographical comparisons revealed three different groups of populations clearly differentiated, i.e. Scandinavia (Norway and Sweden), Iceland and Russia. Cluster and principal coordinates analyses revealed the same genetic patterns of relationships among populations. Generally, this study indicates that E. alaskanus contains low allozymic variation in its populations. The implications of these results for the conservation of the species are discussed. Received: 23 October 1998 / Accepted: 19 December 1998     

Concordance between life history traits, invasion history, and allozyme diversity of the Everglades invader Melaleuca quinquenervia

During the century following its initial introduction in 1886, the Australian tree Melaleuca quinquenervia (Myrtaceae) dispersed from a few introduction points to occupy over 200,000 ha, primarily in historic Everglades wetlands of southern Florida. Cellulose acetate gel electrophoresis (CAGE) was used to investigate the allozyme diversity and population genetic structure of 208 individuals in a dozen populations resulting from this invasion. The analyses showed that these populations have a high (82%) rate of polymorphic loci and an average of 2 alleles/locus. There was substantial heterozygosity (mean H_e = 0.356), which concords well with recent studies reporting a greater number of introduction events and sources than generally recognized. The introduction history and distributional patterns within Florida have led to geographic structuring (G_ST = 0.419) in which the Gulf Coast metapopulation has a greater effective number of alleles and greater heterozygosity than the Atlantic Coast metapopulation. The gene diversity in M. quinquenervia was comparable to other tropical woody species. Its strong population divergence was reminiscent of pioneer species and consistent with its status as a plant invader in Florida.     

Genetic diversity in the rare orchid Gymnadenia odoratissima and a comparison with the more common congener, G. conopsea

Patterns and levels of genetic diversity mayhave significant influence on the long termpersistence of local populations and revealingsuch information is important in protectingrare species. In this study we investigated thegenetic pattern in five microsatellite lociwithin five Swedish populations of the rareorchid species Gymnadenia odoratissima. Thegeographic distribution of G. odoratissima isrestricted to Europe and in Scandinavia it isonly found in three provinces in southernSweden; Östergötland,Västergötland and on the island ofGotland.Compared with the more widespread congener G.conopsea our results indicate lower levels ofgenetic variation within and higher degrees ofgenetic differentiation among populations ofG. odoratissima (H_EL = 0.6–0.8 in G. conopseaand 0.3–0.7 in G. odoratissima; F_ST over allpopulations = 0.06 in G. conopsea and 0.19 inG. odoratissima). Also, we found a cleardistinction among mainland and islandpopulations of G. odoratissima wherepopulations on the island of Gotland seem toexhibit higher levels of gene flow andintragenetic variation, probably as a result ofa larger number of existing populations.Future conservation of this species shouldfocus on facilitation on colonisation events,especially on the mainland, and preservation ofthe genetically more variable Gotlandpopulations.