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.     

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.     

Population history of the terrestrial orchid Cremastra appendiculata var. variabilis from Korea,inferred from levels and distribution of genetic diversity

Cremastra appendiculata var. variabilis is a self‐compatible, insect‐pollinated, terrestrial orchid that is a typical member of the warm‐temperate vegetation in the Korean Peninsula. Here we examine levels and partitioning of allozyme diversity (22 loci) in 12 populations of this orchid to gain insight into its genetic structure and post‐glacial colonization history in Korea. It harboured considerably higher levels of genetic variation within populations (%P = 48.1, A = 1.70 and H_e = 0.217) and lower degree of differentiation among populations (F_ST = 0.068) than those typical of allozyme‐based studies in other terrestrial orchid species. These patterns suggest that extant populations were derived from multiple source populations (i.e. from multiple glacial refugia), although further studies are needed to confirm this scenario. In addition to population history, traits such as high potential of seed dispersal, a mixed mating system and its occurrence in large and continuous populations would have contributed to the current levels and distribution of genetic diversity in Korean populations of C. appendiculata var. variabilis. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 721–732.     

Contemporary habitat loss reduces genetic diversity in an ecologically specialized butterfly

Aim This study investigated the influence of contemporary habitat loss on the genetic diversity and structure of animal species using a common, but ecologically specialized, butterfly, Theclinesthes albocincta (Lepidoptera: Lycaenidae), as a model. Location South Australia. Methods We used amplified fragment length polymorphism (AFLP) and allozyme datasets to investigate the genetic structure and genetic diversity among populations of T. albocincta in a fragmented landscape and compared this diversity and structure with that of populations in two nearby landscapes that have more continuous distributions of butterflies and their habitat. Butterflies were sampled from 15 sites and genotyped, first using 363 informative AFLP bands and then using 17 polymorphic allozyme loci (n = 248 and 254, respectively). We complemented these analyses with phylogeographic information based on mitochondrial DNA (mtDNA) haplotype information derived from a previous study in the same landscapes. Results Both datasets indicated a relatively high level of genetic structuring across the sampling range (AFLP, F_ST = 0.34; allozyme, F_ST = 0.13): structure was greatest among populations in the fragmented landscape (AFLP, F_ST = 0.15; allozyme, F_ST = 0.13). Populations in the fragmented landscape also had significantly lower genetic diversity than populations in the other two landscapes: there were no detectable differences in genetic diversity between the two continuous landscapes. There was also evidence (r² = 0.33) of an isolation by distance effect across the sampled range of the species. Main conclusions The multiple lines of evidence, presented within a phylogeographic context, support the hypothesis that contemporary habitat fragmentation has been a major driver of genetic erosion and differentiation in this species. Theclinesthes albocincta populations in the fragmented landscape are thus likely to be at greater risk of extinction because of reduced genetic diversity, their isolation from conspecific subpopulations in other landscapes, and other extrinsic forces acting on their small population sizes. Our study provides compelling evidence that habitat loss and fragmentation have significant rapid impacts on the genetic diversity and structure of butterfly populations, especially specialist species with particular habitat preferences and poor dispersal abilities.     

Mating system and population genetic structure of Bruguiera gymnorrhiza (Rhizophoraceae), a viviparous mangrove species in China

In order to explain the diversity patterns and develop the conservation strategies, the population genetic structures and the mating systems of Bruguiera gymnorrhiza from the coastlines of south China were investigated in this study. The mating system parameters were analyzed using progeny arrays for allozyme markers. The multilocus outcrossing rates (tm) ranged from 0.845 (Fugong) to 0.267 (Dongzhai harbor). High allozyme variations within the five collected populations were determined and compared with the published data of other plant species with the mixed mating systems. At species level, the percentage of polymorphic loci (P) was 80%, the average number of alleles per locus (A) was 2.440, and the heterozygosity (He) was 0.293. The total gene diversity within each population (H_S = 0.2782) and the coefficient of genetic differentiation (G_ST = 0.0579) among the populations were estimated. On the basis of this population genetic structure, it is suggested that the gene flow (Nm = 3.85) is quite high, which is possibly related to its water-dispersed hypocotyls. It is also suggested that the mating system of this species is of mixed mating.     

Genetics of Brazil nut (Bertholletia excelsa Humb. & Bonpl.: Lecythidaceae)

Summary We provide an estimate of genetic variation within and between two populations of Bertholletia excelsa (Brazil nut), a large canopy tree found in the rain forests of South America. Average heterozygosity is 0.190, and 54.3% of the sampled loci are polymorphic. The population structure deviates significantly from Hardy-Weinberg expectations for Fest2 and Pgm2 (F =0.405 and 0.443, respectively) in one population, and highly significantly (F=-0.341) for Gdh in the other population. Although allele frequencies of the two populations differ significantly for Aat2, Est5, Mdh1, and Mdh2B, Nei's coefficient of gene differentiation (G_st) indicates that the between-population component (D_st) of genic diversity represents only 3.75% of the size of the within-population component (H_s). The implications of these findings in terms of conservation genetics are that much of the genetic diversity of this species may be preserved within one or a few populations. However, such populations must be very large because it appears that the large amount of genetic variation in Brazil nut populations is maintained by extensive gene flow and bonds of mating over a large area. The genetic architecture of Bertholletia excelsa is similar to that expected for an extensively diploidized paleopolyploid species.     

Allozyme variation and population genetic structure of common wild rice Oryza rufipogon Griff. in China

In order to determine the genetic diversity and genetic structure of populations in common wild rice Oryza rufipogon, an endangered species, allozyme diversity was analyzed using 22 loci in 607 individuals of 21 natural populations from the Guangxi, Guangdong, Hainan, Yunnan, Hunan, Jiangxi and Fujian provinces in China. The populations studied showed a moderate allozyme variability (A=1.33, P=22.7%, Ho=0.033 and He=0.068), which was relatively high for the genus Oryza. The levels of genetic diversity for Guangxi and Guangdong were significantly higher than those for the other regions, and thus South China appeared to be the center of genetic diversity of O. rufipogon in China. A moderate genetic differentiation (F_ST=0.310, I=0.964) was found among the populations studied. Interestingly, the pattern of population differentiation does not correspond to geographic distance. An estimate of the outcrossing rate (t=0.324) suggests that the species has a typical mixed-mating system. The deficit of heterozygotes (F=0.511) indicates that some inbreeding may have taken place in outcrossing asexual populations because of intra-clone outcrossing events and ”isolation by distance” as a result of human disturbance. In order to predict the long-term genetic survival of fragmented populations, further studies on gene flow among the remaining populations and the genetic effects of fragmentation are proposed. Finally, some implications for the conservation of endangered species are suggested. Received: 22 June 1999 / Accepted: 20 December 1999     

Genetic structure of Polytrichum formosum in relation to the breeding system as revealed by microsatellites

Microsatellite variation was determined for three Danish and three Dutch populations of the haploid moss species Polytrichum formosum to gain insight into the relative importance of sexual vs. asexual reproduction for the amount and structure of genetic variation. In general, low levels of microsatellite variation were observed within this species. Even when estimated for polymorphic loci only, the levels of microsatellite variability (P=90.6, A=4.3 and H_S=0.468) within populations were on average lower than those reported for most other plant species. In contrast, genotypic diversity was high within each of the examined populations, indicating that sexual reproduction is a very important determinant of the genetic structure of P. formosum within populations. In agreement with previous findings for allozyme data, no significant genetic differentiation (F_ST=0.028, R_ST=0.015) was observed neither between populations nor between regions approximately 450 km apart (Denmark vs. the Netherlands). These low levels of population differentiation observed for both types of genetic markers are probably best explained by a high level of effective spore dispersal (gene flow) between populations. Therefore, also on a large geographical scale sexual reproduction is the most important determinant of the genetic structure of P. formosum, despite the high potential to reproduce clonally.     

Mating system and genetic diversity of a rare desert legume Ammopiptanthus nanus (Leguminosae)

Ammopiptanthus nanus is an endangered evergreen shrub endemic to the deserts of central Asia and plays an important role in delaying further desertification. We examined allozyme variation and AFLP diversity in A. Nanus populations and investigated the mating system of this species using progeny arrays assayed for poly-morphic allozyme loci. Mating system analysis in the Keyi'eryongke'er population showed low levels of out-crossing, and strong inbreeding depression. Low levels of genetic variation were detected at both population (allozyme, Pp=14.0%,A=1.14, He=0.031; AFLP, Pp=14.5%, Shannon's information index I=0.063) and species (allozyme, Pp=21.1%,A=1.21, He=0.040; AFLP, Pp=20.9%, I=0.083) levels; while moderate genetic differentia-tion existed among populations, as indicated by allozymes (GST=0.081) and AFLP (GST=0.151-0.193). Founder effect, bottlenecks in evolutionary history, the mixed mating system and co-ancestry may have influenced the level of genetic diversity in A. Nanus. Markers of both types provide new insights for conservation management, indicating that the Biao'ertuokuoyi and Keyi'eryongke'er populations should be given priority for in situ conser-vation and regarded as seed sources for ex situ conservation.     

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 Structure and Gene Flow in Elymus glaucus (blue wildrye): Implications for Native Grassland Restoration

Interest in using native grass species for restoration is increasing, yet little is known about the ecology and genetics of native grass populations or the spatial scales over which seed can be transferred and successfully grown. The purpose of this study was to investigate the genetic structure within and among populations of Elymus glaucus in order to make some preliminary recommendations for the transfer and use of this species in revegetation and restoration projects. Twenty populations from California, Oregon, and Washington were analyzed for allozyme genotype at 20 loci, and patterns of variation within and among populations were determined. Allozyme variation at the species level was high, with 80% of the loci polymorphic and an average expected heterozygosity (an index of genetic diversity) of 0.194. All but two of the populations showed some level of polymorphism. A high degree of population differentiation was found, with 54.9% of the variation at allozyme loci partitioned among populations (F_st= 0.549). A lesser degree of genetic differentiation among closely spaced subpopulations within one of the populations was also demonstrated (F_st= 0.124). Self-pollination and the patchy natural distribution of the species both likely contribute to the low level of gene flow (N_m= 0.205) that was estimated. Zones developed for the transfer of seed of commercial conifer species may be inappropriate for transfer of E. glaucus germplasm because conifer species are characterized by high levels of gene flow. Limited gene flow in E. glaucus can facilitate the divergence of populations over relatively small spatial scales. This genetic differentiation can be due to random genetic drift, localized selective pressures, or both. In order to minimize the chances of planting poorly adapted germplasm, seed of E. glaucus may need to be collected in close proximity to the proposed restoration site.     

Patterns of Genetic Diversity and Population Structure of the Clonal Herb,Potentilla fragarioides var. sprengeliana (Rosaceae) in Korea

The genetic diversity and population structure of Potentilla fragarioides var. sprengeliana(Rosaceae) in Korea were investigated using genetic variation at 22 allozyme loci. The percent of polymorphic loci within the enzymes was 59.1%. The genetic diversity at the species level and at the population level was high (Hes=0.210; Hep=0.199, respectively), whereas the extent of the population divergence was relatively low (GST=0.074). FIS, a measure of the deviation from random mating within the 19 populations, was 0.331. An indirect estimate of the number of migrants per generation (Nm=3.15) indicates that gene flow is 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. The mean genetic identity between populations was 0.985. It is highly probable that the trend of genetic uniformity in a relatively homogeneous habitat is thought to be operated among Korean populations of P. fragarioides var.sprengeliana.     

在韩国境内Potentilla fragarioides var .sprengeliana的遗传多样性和种群结构(英文)

根据 2 2个等位酶位点遗传变异 ,探讨了韩国境内委陵菜 (PotentillafragarioidesL .var.sprengeliana)的遗传多样性和种群结构。酶位点的多态位点百分比为 5 9 1%。种和种群水平上的遗传多样性比较高 ,分别为Hes=0 .2 10 ,Hep=0 .199;而种群的分化水平则相对较低 (GST=0 .0 74)。 19个种群中随机交配的偏差为FIS=0 .331。每代迁移数的间接估计 (Nm=3.15 )表明该种在韩国的种群间基因流较高。另外 ,固定指数分析显示在一些种群和位点有轻微的杂合子缺乏。种群间平均遗传一致度为 0 985。这些韩国委陵菜种群存在于较为均一的生境 ,这很有可能是造成其种群遗传一致性较高的原因。     

Genetic variation and the mating system in the rare Acacia sciophanes compared with its common sister species Acacia anfractuosa (Mimosaceae)

Acacia sciophanes is an extremely rare and Critically Endangered species known from two small populations separated by less than 7 km. Specifically we aimed to investigate whether rarity in A. sciophanes is associated with decreased levels of genetic variation and increased levels of selfing by comparing patterns of genetic variation and mating system parameters with its widespread and common sister species A. anfractuosa. Fourteen polymorphic allozyme loci were used to assess genetic diversity with four of these used in the estimation of mating system parameters. At the species level A. sciophanes has lower allelic richness, polymorphism, observed heterozygosity and gene diversity than A. anfractuosa and significantly lower levels of gene diversity at the population level. Both species have a mixed mating system but the largest population of A. sciophanes has lower levels of outcrossing, higher correlated paternity and increased bi-parental inbreeding compared with two A. anfractuosa populations. However, both correlated paternity and bi-parental inbreeding appear to be at least partly influenced by population size regardless of the species. We suggest that A. sciophanes is likely to be an intrinsically rare species and that in particular the lower levels of genetic diversity and increased selfing are a feature of a species that has the ability to persist in a few localised small populations. Despite recent extensive habitat destruction our comparative study provided no clear evidence that such events have contributed to the lower genetic diversity and increased selfing in A. sciophanes and we believe its ability to exist in small populations may not only be an important factor in its survival as a rare species but also indicates that it may be less susceptible to the impacts of habitat loss and fragmentation. The key to this species conservation will be the maintenance of suitable habitat, particularly through improved fire regimes and control of invasive weeds, that will allow the two small populations to continue to persist in extremely restricted areas of remnant vegetation.     

Genetic analysis of 50-year old Brazilian pine (Araucaria angustifolia) plantations: implications for conservation planning

In order to determine the implications of including planted populations in conservation planning, we investigate the structure and genetic diversity, mating system, and effective population size within families in three planted and one natural Araucaria angustifolia populations. The study area is a Brazilian National Forest in southern Brazil, established in the 1950s. Genetic analysis was investigated using nine allozyme loci. Significant and positive fixation index was detected for the adults of all studied populations. Although we observed lower levels of genetic diversity and biparental inbreeding in some populations, their pattern of mating, similar to that expected in panmixia, indicates that all populations can be used in conservation planning. Furthermore, the presence of unique alleles and the significant genetic differentiation among all adult populations (F _ST = 0.404) reinforces that all populations are important conservation resources. The average paternity correlation was high in almost all populations, showing that open-pollinated families are composed of mixtures of half- and full-sibs. With regards to conservation strategies, our results suggest that seed collection from at least 133 seed trees is necessary to retain a reference effective population size of 500. Our results confirm that the planted Araucaria forests in southern Brazil can be useful as a genetic resource for conservation, breeding and reforestation strategies for this threatened species.     

Reproductive biology and genetic diversity of a cryptoviviparous mangrove aegiceras corniculatum (Myrsinaceae) using allozyme and intersimple sequence repeat (ISSR) analysis

Mangroves consist of a group of taxonomically diverse species representing about 20 families of angiosperms. However, little is known about their reproductive biology, genetic structure, and the ecological and genetic factors affecting this structure. Comparative studies of various mangrove species are needed to fill such gaps in our knowledge. The pollination biology, outcrossing rate, and genetic diversity of Aegiceras corniculatum were investigated in this study. Pollination experiments suggested that the species is predominantly pollinator-dependent in fruit setting. A quantitative analysis of the mating system was performed using progeny arrays assayed for intersimple sequence repeat (ISSR) markers. The multilocus outcrossing rate (tm) was estimated to be 0.653 in a wild population. Both allozyme and ISSR were used to investigate genetic variation within and among populations. The combined effects of founder events and enhanced local gene flow through seedling dispersal by ocean currents apparently played an important role in shaping the population genetic structure in this mangrove species. Both allozyme variation (P = 4.76%, A = 1.05, HE = 0.024) and ISSR diversity (P = 16.18%, A = 1.061, HE = 0.039) were very low at the species level, in comparison with other woody plants with mixed-mating or outcrossing systems. Gene differentiation among populations was also low: allozyme GST = 0.106 and ISSR GST = 0.178. The unusually high genetic identities (0.997 for allozyme and 0.992 for ISSR loci), however, suggest that these populations are probably all descended from a common ancestral population with low polymorphism.     

Population-Genetic Structure of Beaver (Castor fiber L., 1758) Communities and Estimation of Effective Reproductive Size N e of an Elementary Population

The absence of panmixia at all hierarchical levels of the European beaver communities down to individual families implies a complex organization of the population-genetic structures of the species, in particular, a large intergroup component of gene diversity in the populations. Testing this assumption by analysis of 39 allozyme loci in the communities of reintroduced beaver from the Vyatka river basin (Kirov oblast) has shown that only the beaver colonies exhibit high intergroup gene diversity (G _st = 0.32) whereas this parameter is much lower when estimated among beaver groups from individual Vyatka River tributaries and among localities of one of the tributaries (0.07 and 0.11, respectively). The data suggesting genetic heterogeneity among individual settles within colonies have been obtained. The factors affecting the structure of the beaver communities of the lower hierarchical ranks are considered: the common origin, founder effect, selection, gene drift, assortative mating, and social and behavior features of the species. The conclusion is drawn that the founder effect could be the primary factor of population differentiation only at the time of their formation. The heterogeneity among colonies and among settles is maintained largely by isolation of colonies from one another. The strong interspecific competition for food resources, which is behaviorally implemented in the species at the level of minimal structural units (individual settles) creates a profound and unique population-genetic subdivision of the species. These results substantiate the suggestion that an elementary population (micropopulation) of European beaver is a colony, i.e., a set of related settles of different types. Based on ecological and genetic parameters, the effective reproductive size N _e of the minimum beaver population was estimated to be equal to three animals. This extremely low value of effective reproductive population size largely explains the high tolerance of European beaver to inbreeding and striking viability of the species, which from the early 19th century has been for more than hundred years on the brink of survival in the condition which would made any other mammalian species vanish from the Earth.     

Mating system and pollen flow between remnant populations of the endangered tropical tree, <Emphasis Type="Italic">Guaiacum sanctum</Emphasis> (Zygophyllaceae)

Tropical trees are generally long-lived making it difficult to assess the long-term effects of habitat fragmentation on genetic diversity. Maintenance of genetic diversity in fragmented landscapes is largely dependent on the species’ mating system and the degree of genetic connectivity (seed and pollen flow) among fragments. Currently, these parameters are largely unknown for many endangered tropical tree species. Additionally, landscape fragmentation may isolate tropical tree populations from larger, more continuous populations. The role of isolated individuals in pollen transfer within and between remnant populations is not clear. In this study, we estimate the mating system and pollen flow patterns in continuous and remnant populations of the endangered tropical tree Guaiacum sanctum (Zygophyllaceae). Fractional paternity analyses were used to estimate average gene flow distances between fragmented remnant populations and the siring success of an intermediately located, but isolated individual. In these populations, G. sanctum is a mixed-mating species (t _m = 0.72 − 0.95) whose pollen is transported over large distances (>4 km). An isolated tree may have functioned as a stepping-stone between two clusters of individuals, assisting long-distance pollen movement. This individual also sired a disproportionately high number of seeds (13.9%), and is thus an important component of the reproductive success of these populations, thus rejecting Janzen’s “living-dead” hypothesis. The high levels of genetic diversity maintained as a consequence of long-distance pollen-flow suggest that this endangered species may have the potential for future adaptation and population expansion if suitable habitats become available.     

Contrasting levels of genetic diversity between the historically rare orchid Cypripedium japonicum and the historically common orchid Cypripedium macranthos in South Korea

Many terrestrial orchids are historically rare and occur in small, spatially isolated populations. Theory predicts that such species will harbour low levels of genetic variation within populations and will exhibit a high degree of population genetic divergence, primarily as a result of genetic drift. If the origin of the present‐day populations is relatively recent from the same genetically depauperate source population, a complete lack of genetic differentiation between conspecific populations is expected. If a terrestrial orchid was historically common with moderate or high levels of genetic diversity, but has experienced more recent anthropogenic disturbance as a result of over‐collection, it would still exhibit initial levels of genetic variation within populations and a low degree of genetic divergence between populations. To test these predictions, we examined the genetic diversity in six populations (N = 131) of the historically and currently rare Cypripedium japonicum and in four populations (N = 94) of the historically common but now rare C. macranthos from South Korea. Fourteen putative allozyme loci resolved from eight enzyme systems revealed no variation either within or among populations of C. japonicum, which supports the first prediction. In contrast, populations of C. macranthos harboured high levels of genetic variation (mean percentage of polymorphic loci %P = 46.7; mean expected heterozygosity H_e = 0.185) and exhibited a low degree of population genetic divergence (G_ST = 0.059), supporting the second prediction. The lack of genetic variation both within and among conspecific populations of C. japonicum may suggest that populations originated from the same genetically depauperate ancestral population. The high levels of genetic diversity maintained in populations of C. macranthos suggest that the collection‐mediated decrease in the number of individuals is still too recent for long‐term effects on genetic variation. Based on current demographic and genetic data, in situ and ex situ conservation strategies should be provided to preserve genetic variation and to ensure the long‐term survival of the two species in the Korean Peninsula. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160 , 119–129.