Low Within-population Genetic Diversity Research Articles

Phylogeographic analysis reveals early Pleistocene divergence and strong genetic structure in Euphorbia phosphorea (Euphorbiaceae), an endemic species to the semi-arid Caatinga Biome

Abstract The largest continuous patch of Seasonally Dry Tropical Forests (SDTF) is the Caatinga Biome in north-eastern Brazil, which has the greatest biodiversity and highest degree of endemism compared to any other SDTF patch of South America. Our aim was to conduct a phylogeographic study with Euphorbia phosphorea to understand the evolutionary history of this species endemic to the Caatinga Biome. We studied genetic diversity and structure, demographic dynamics, as well as intraspecific phylogenetic relationships, based on plastidial and nuclear DNA. In addition, we conducted species distribution modelling (SDM) to infer connectivity since the Last Interglacial. Our results recovered a stronger genetic structure for plastidial than for nuclear genome, which probably result from distinct species capability to seed and pollen dispersal, and low levels of genetic diversity within populations. Although SDM results suggest higher connectivity during Last Maximum Glacial, demographic analyses detected no signs of demographic expansion in this period. In agreement with previous studies with rock specialists, our data reveal remarkable among-population differentiation and low within-population genetic diversity, which make populations more prone to drift effects. Furthermore, our results shed light on the importance of early Pleistocene events in the diversification of lineages within the Caatinga Biome, although further comparative work is required.

Genetic diversity of the threatened <i>Saussurea dorogostaiskii</i> (Asteraceae) in the Khuvsgul region of Mongolia

<i>Saussurea dorogostaiskii</i> Palib. (Asteraceae) is a critically endangered medicinal plant in Mongolia and Russia. We studied the genetic variation of <i>S. dorogostaiskii</i> from three mountains of northern Mongolia. The genetic profile was assessed in 70 individuals from eight populations using five inter-simple sequence repeat markers, producing 53 loci with 96.4% polymorphism across all bands. Shannon’s index (<i>I</i>) and Nei’s gene diversity (<i>H</i>) value at the species level of <i>S. dorogostaiskii</i> are 0.25 and 0.17, respectively. An AMOVA showed high genetic variation among the populations (22% of populations and 32% of mountains), consistent with the high genetic differentiation (<i>G<sub>ST</sub></i> = 0.49) and low gene flow (<i>N<sub>m</sub></i> = 0.51) in <i>S. dorogostaiskii</i> populations. Eight populations were clustered into two groups, corresponding to their geographic locations. The low within-population genetic diversity and high genetic differentiation among <i>S. dorogostaiskii</i> populations factor into their endangered designation. This genetic analysis reveals that all populations are equally threatened, and community-based conservation is appropriate for these species.

Genetic diversity and gene expression diversity shape the adaptive pattern of the aquatic plant Batrachium bungei along an altitudinal gradient on the Qinghai-Tibet plateau.

It is an intriguing issue of evolutionary biology how genetic diversity and gene expression diversity shape the adaptive patterns. Comparative transcriptomic studies of wild populations in extreme environments provide critical insights into the relative contribution of genetic and expressive components. In this study, we analyzed the genetic diversity and gene expression diversity of 20 populations of the aquatic plant Batrachium bungei along elevations ranging from 2690 to 4896m on the Qinghai-Tibet plateau (QTP). Based on single nucleotide polymorphisms (SNPs) and gene expression data from 100 individuals of B. bungei, we found that variation in genetic sequence was more sensitive to detect weak differentiation than gene expression. Using 292,613 high-quality SNPs, we documented a significant phylogeographical structure, a low within-population genetic diversity, and a high inter-population genetic differentiation in B. bungei populations. Analysis of relationship between geographic distance, genetic distance, and gene expression similarity showed that geographic isolation shaped gene flow patterns but not gene expression patterns. We observed a negative relationship between genetic diversity and gene expression diversity within and among B. bungei populations, and we demonstrated that as environmental conditions worsen with increasing altitude, genetic diversity played an increased role in maintaining the stability of populations, while the corresponding role of gene expression diversity decreased. These results suggested that genetic diversity and gene expression diversity might act as a complementary mechanism contributing to the long-term survival of B. bungei in extreme environments.

Recent, Late Pleistocene fragmentation shaped the phylogeographic structure of the European black pine (Pinus nigra Arnold)

Fragmentation acting over geological times confers wide, biogeographical scale and genetic diversity patterns to species, through demographic and natural selection processes. To test the effects of historical fragmentation on the genetic diversity and differentiation of a widespread forest tree, Pinus nigra Arnold, the European black pine, and to resolve its demographic history, we described and modelled its spatial genetic structure and gene genealogy. We then tested which Pleistocene event, whether recent or ancient, could explain its widespread but patchy geographic distribution. We used a set of different genetic markers, both neutral and potentially adaptive, and either bi-parentally or paternally only inherited, and we sampled natural populations across the entire species range. We analysed the data using both frequentist population genetic and Bayesian inference methods to calibrate realistic, demographic timed scenarios. We also considered how habitat suitability might have affected demography by correlating climate variables at different recent Pleistocene ages with genetic diversity estimates. Species with geographically fragmented distribution areas are expected to display significant among-population genetic differentiation and low within-population genetic diversity. Contrary to these expectations, we show that the current diversity of Pinus nigra and its weak genetic spatial structure result from the Late Pleistocene or Early Holocene fragmentation of one ancestral population into six distinct genetic lineages. Gene flow among the different lineages is strong across forests and many current populations are admixed between lineages. We propose to modify the currently accepted international nomenclature made of five sub-species and name these six lineages using regionally accepted sub-species-level names.

Population genomic evidence for plant glacial survival in Scandinavia.

Quaternary glaciations have played a major role in shaping the genetic diversity and distribution of plant species. Strong palaeoecological and genetic evidence supports a postglacial recolonization of most plant species to northern Europe from southern, eastern and even western glacial refugia. Although highly controversial, the existence of small in situ glacial refugia in northern Europe has recently gained molecular support. We used genomic analyses to examine the phylogeography of a species that is critical in this debate. Carex scirpoidea Michx subsp. scirpoidea is a dioecious, amphi-Atlantic arctic-alpine sedge that is widely distributed in North America, but absent from most of Eurasia, apart from three extremely disjunct populations in Norway, all well within the limits of the Weichselian ice sheet. Range-wide population sampling and variation at 5,307 single nucleotide polymorphisms show that the three Norwegian populations comprise unique evolutionary lineages divergent from Greenland with high between-population divergence. The Norwegian populations have low within-population genetic diversity consistent with having experienced genetic bottlenecks in glacial refugia, and host private alleles that probably accumulated in long-term isolated populations. Demographic analyses support a single, pre-Weichselian colonization into Norway from East Greenland, and subsequent divergence of the three populations in separate refugia. Other refugial areas are identified in North-east Greenland, Minnesota/Michigan, Colorado and Alaska. Admixed populations in British Columbia and West Greenland indicate postglacial contact. Taken together, evidence from this study strongly indicates in situ glacial survival in Scandinavia.

Low within-population genetic diversity and high genetic differentiation among populations of the endangered plant Tetracentron sinense Oliver revealed by inter-simple sequence repeat analysis

Tetracentron sinense Oliver, an endangered species from China, displays a low within-population genetic diversity and high genetic differentiation among populations, and the existing populations could be divided into three conservation and management units. The endangered tree Tetracentron sinense Oliver has great value; however, little is known regarding the within-population genetic diversity and differentiation among T. sinense populations. We examined the genetic diversity and differentiation of T. sinense wild populations, and we tested the effect of small-size population on the level of genetic diversity within these populations. Using inter-simple sequence repeat (ISSR), we assessed the genetic variation and structure among 174 individuals from 26 natural populations of T. sinense sampled across its distribution range in China. The ISSR primers yielded 180 amplified loci (123 were polymorphic). At the species level, the percentage of polymorphic loci (PPL), Nei’s gene diversity (H), and Shannon’s information index (I) were 68.3%, 0.196 and 0.300, respectively. The average population level PPL was 20.0%, and the Na, Ne, H, and I were 1.20, 1.13, 0.076, and 0.112, respectively. AMOVA revealed high genetic differentiation among populations (52.0% of total variance, P = 0.001), consistent with the gene differentiation coefficient (Gst = 0.607) and gene flow (Nm = 0.326). The 174 individuals of the 26 T. sinense populations clustered into three groups, and T. sinense geographic and genetic distance were significantly correlated. T. sinense exhibited intermediate within-species genetic diversity, indicating preserved evolutionary potential. The low within-population genetic diversity and high genetic differentiation among T. sinense populations may be one of important factors causing endangerment. Three conservation units were determined based on genetic difference and structure. Inter-population introduction of individuals within units via appropriate propagation and seedling management might be an effective strategy for increasing T. sinense within-population genetic diversity and population size.

Regional population structure of the endangered Bridle Shiner (Notropis bifrenatus)

In the last 100 years, the Bridle Shiner has declined over significant parts of its range. We used mitochondrial and nuclear microsatellite markers to investigate population structure of this species. Tissue samples were collected from populations in six drainages in PA, NJ, NY and CT. One predominant haplotype was observed in the Delaware, Housatonic, Passaic, and Raritan specimens. Specimens from the Hudson and St. Lawrence drainages had a separate unique haplotype. Microsatellite loci revealed low within-population genetic diversity and pairwise population comparisons of allelic divergence showed significant genetic differences among most drainages. Population structure analysis of microsatellite loci recover the same population clusters inferred using mtDNA. Within the Delaware drainage, there was significant differentiation among areas separated by 128 km. Several alternate scenarios of population divergence and population bottlenecks were investigated using approximate Bayesian computing. These supported a scenario with a bottleneck occurring in the ancestral population of Bridle Shiner followed by recent divergence of Northern and Southern drainages.

Genetic diversity and demographic history of the Siberian lime (Tilia sibirica)

Tilia sibirica Bayer (Siberian lime) is endemic to the low mountain systems north of the Altai Mountains of southern Siberia, approximately 1,000 kms to the east of the natural range limit of its closest congeneric, the small-leaved lime (Tilia cordata). Some consider the taxon to be a subspecies of T. cordata. This putative pre-Last Glacial Maximum (LGM) relict may have had a stronghold in a refugium of the Altai Mountains and survived various waves of fluctuating climatic changes that occurred in the region. With continued climatic changes expected, these hardy but isolated populations can be important sources for population expansion. To date, we do not know the genetic status or history of this forest tree.This study uses standard population genetic and Approximate Bayesian Computation (ABC) analyses of microsatellite data to determine the genetic diversity and differentiation, clonal occurrence and date of divergence of the two lime taxa. The results show that T. sibirica and T. cordata are distinct biological units with significant genetic differentiation. The ABC analysis suggests a (Middle) Pleistocene divergence. We have revealed low within-population genetic diversity as well as high levels of clonality in T. sibirica. The focus now should be on restoring and conserving these small and isolated relict populations.

Distribution and Genetic Structure of Fucus distichus Linnaeus 1953 (formerly F. gardneri) within Central San Francisco Bay

Fucus distichus, a rockweed common to the mid-intertidal shoreline within the San Francisco Estuary (previously known as F. gardneri), was injured during the Cosco Busan oil spill in November 2007 and subsequent clean-up actions. Restoration planning activities are underway to help recover F. distichus at sites within central San Francisco Bay where damage occurred. As a first step, we conducted shoreline surveys during the summers of 2012–2013 to map the occurrence of this rockweed. Of the 151.73 km of rocky shoreline within the central bay, F. distichus covered 32.16 km of shoreline. The alga generally occurred in narrow bands but formed expansive beds at locations with natural, flat bedrock benches. We also observed F. distichus on artificial substrata such as seawalls and riprap, but not on pilings. Samples of F. distichus from 11 sites throughout the central / east San Francisco Bay were genetically analyzed (microsatellite genotyping). The populations analyzed (1) had low genetic diversity, (2) the frequency of homozygotes was higher than expected (suggesting high inbreeding), and (3) also displayed geographic population structure, in part driven by very small differences in the midst of extremely low within-population genetic diversity. However, these genetic data do not raise concerns for restoration methods in terms of choosing donor populations and mixing F. distichus from different sites within the central bay. The choice of donor populations should be based on practical criteria for effective restoration; individuals will nonetheless be taken from locations as nearby to donor sites as possible. Various locations throughout the central San Francisco Bay are composed of cobble or small riprap that are populated with F. distichus, which could provide efficient means of translocating rockweed for future restoration activities.

Genetic structure and demographic history of Cycas chenii (Cycadaceae), an endangered species with extremely small populations

Geological activities and climate oscillations during the Quaternary period profoundly impacted the distribution of species in Southwest China. Some plant species may be harbored in refugia, such as the dry-hot valleys of Southwest China. Cycas chenii X. Gong & W. Zhou, a critically endangered cycad species, which grows under the canopy in subtropical evergreen broad-leaved forests along the upstream drainage area of the Red River, is endemic to this refugium. In this study, 60 individuals of C. chenii collected from six populations were analyzed by sequencing two chloroplast intergenic spacers (cpDNA: psbA-trnH and trnL-trnF) and two nuclear genes (PHYP and RBP-1). Results showed high genetic diversity at the species level, but low within-population genetic diversity and high interpopulation genetic differentiation. A Bayesian phylogenetic tree based on cpDNA showed that five chloroplast haplotypes were clustered into two clades, which corresponds to the division of the western and eastern bank of the Red River. These data indicate a possible role for the Red River as a geographic barrier to gene flow in C. chenii. Based on our findings, we propose appropriate in situ and ex situ conservation strategies for C. chenii.

Genetic diversity and structuring across the range of a widely distributed ladybird: focus on rear-edge populations phenotypically divergent.

Population genetics and phenotypic structures are often predicted to vary along the geographic range of a species. This phenomenon would be accentuated for species with large range areas, with discontinuities and marginal populations. We herein compare the genetic patterns of central populations of Coccinella septempunctata L. with those of two phenotypically differentiated populations considered as rear‐edge populations and subspecies based on phenotype (Algeria and Japan). According to the central‐marginal model and expected characteristics of rear‐edge populations, we hypothesize that these rear‐edge populations have (1) a reduced genetic diversity, resulting from their relative isolation over long periods of time, (2) a higher population genetic differentiation, explained by low contemporary gene flow levels, and (3) a relationship between genetic diversity characteristics and phenotypes, due to historical isolation and/or local adaptation. Based on genotyping of 28 populations for 18 microsatellite markers, several levels of regional genetic diversity and differentiation are observed between and within populations, according to their localization: low within‐population genetic diversity and higher genetic differentiation of rear‐edge populations. The genetic structuring clearly dissociates the Algerian and Eastern Asia populations from the others. Geographical patterns of genetic diversity and differentiation support the hypothesis of the central‐marginal model. The pattern observed is in agreement with the phenotypic structure across species range. A clear genetic break between populations of Algeria, the Eastern Asia, and the remaining populations is a dominant feature of the data. Differential local adaptations, absence of gene flow between marginal and central populations, and/or incapacity to mate after colonization, have contributed to their distinct genotypic and phenotypic characteristics.

Genetic diversity and structure of the threatened species Sinopodophyllum hexandrum (Royle) Ying.

Sinopodophyllum hexandrum is an important medicinal plant that has been listed as an endangered species, making the conservation of its genetic diversity a priority. Therefore, the genetic diversity and population structure of S. hexandrum was investigated through inter-simple sequence repeat analysis of eight natural populations. Eleven selected primers generated 141 discernible fragments. The percentage of polymorphic bands was 37.59% at the species level, and 7.66-24.32% at the population level. Genetic diversity of S. hexandrum was low within populations (average HE = 0.0366), but higher at the species level (HE = 0.0963). Clear structure and high genetic differentiation were detected between populations using unweighted pair groups mean arithmetic and principle coordinate analysis. Clustering approaches clustered the eight sampled populations into three major groups, and AMOVA confirmed there to be significant variation between populations (63.27%). Genetic differentiation may have arisen through limited gene flow (Nm = 0.3317) in this species. Isolation by distance among populations was determined by comparing genetic distance versus geographical distance using the Mantel test. The results revealed no correlation between spatial pattern and geographic location. Given the low within-population genetic diversity, high differentiation among populations, and the increasing anthropogenic pressure on this species, in situ conservation measures, in addition to sampling and ex situ preservation, are recommended to preserve S. hexandrum populations and to retain their genetic diversity.

Genetic diversity and structure of Sinopodophyllum hexandrum (Royle) Ying in the Qinling Mountains, China.

Sinopodophyllum hexandrum is an important medicinal plant whose genetic diversity must be conserved because it is endangered. The Qinling Mts. are a S. hexandrum distribution area that has unique environmental features that highly affect the evolution of the species. To provide the reference data for evolutionary and conservation studies, the genetic diversity and population structure of S. hexandrum in its overall natural distribution areas in the Qinling Mts. were investigated through inter-simple sequence repeats analysis of 32 natural populations. The 11 selected primers generated a total of 135 polymorphic bands. S. hexandrum genetic diversity was low within populations (average He = 0.0621), but higher at the species level (He = 0.1434). Clear structure and high genetic differentiation among populations were detected by using the unweighted pair group method for arithmetic averages, principle coordinate analysis and Bayesian clustering. The clustering approaches supported a division of the 32 populations into three major groups, for which analysis of molecular variance confirmed significant variation (63.27%) among populations. The genetic differentiation may have been attributed to the limited gene flow (Nm = 0.3587) in the species. Isolation by distance among populations was determined by comparing genetic distance versus geographic distance by using the Mantel test. Result was insignificant (r = 0.212, P = 0.287) at 0.05, showing that their spatial pattern and geographic locations are not correlated. Given the low within-population genetic diversity, high differentiation among populations and the increasing anthropogenic pressure on the species, in situ conservation measures were recommended to preserve S. hexandrum in Qinling Mts., and other populations must be sampled to retain as much genetic diversity of the species to achieve ex situ preservation as a supplement to in situ conservation.

Assessment of genetic diversity in the endangered populations of Breonadia salicina (Rubiaceae) growing in the kingdom of Saudi Arabia using inter-simple sequence repeat markers.

BackgroundBreonadia salicina (Rubiaceae) is a critically endangered plant at the local scale native to southwestern Saudi Arabia. To understand the levels and partitioning of genetic variation across populations and geographical regions of this species, we assessed its genetic diversity using inter-simple sequence repeat (ISSR) markers.ResultsFourteen ISSR primers selected from 43 primers gave rise to 211 amplified loci, of which 68 were polymorphic. The percentage of polymorphic loci (PPL) at the population level ranged from 17.1 to 23.7%, with an average of 21.3%. Nei’s gene diversity (h) and Shannon’s information index (I) were 0.086 and 0.125, respectively. At the species level, PPL was 32.2%, while h and I were 0.116 and 0.172, respectively. A hierarchical analysis of molecular variance revealed a high level of genetic differentiation among populations (17% of total variance, P = 0.001), consistent with the gene differentiation coefficient (GST = 0.256). Nevertheless, the evaluated genetic diversity was very low within populations; while relatively high among populations, levels were insufficient for long-term survival. Saudi Arabian accessions were also compared to accessions of a population from Yemen, where the species is more widespread. The Yemeni population also showed low genetic diversity but clustered separately.ConclusionsBreonadia salicina in Saudi Arabia is characterized by low within-population genetic diversity and high among-population genetic differentiation. Based on our findings, this locally endangered species is on the verge of local extinction. The species’ survival depends on successful implementation of suggested strategies for its long-term conservation.

Genetic diversity and structure of Lilium pumilum DC. in southeast of Qinghai–Tibet plateau

Lilium pumilum DC. is a valuable species not only for its showy flowers but also for its edible and medicinal values. As one of the distribution areas of L. pumilum, Qinghai–Tibet plateau has unique environmental features which have high impact on the evolution of the species. No population genetic studies have been done for L. pumilum so far. To provide the first reference data for evolutionary study and understanding the influence of eco-geographic factors on the distribution of genetic variation in L. pumilum, interspecific simple sequence repeat markers were used to investigate genetic diversity and population structure of 28 populations sampled from southeast of Qinghai–Tibet plateau. Fifteen selected primers generated a total of 147 polymorphic bands. The genetic diversity was low within populations (average He = 0.173), but higher at the species level (He = 0.392). A clear population structure and high level of genetic differentiation (FST = 0.518) were detected by unweighted pair group method for arithmetic averages, principle coordinate analysis and Bayesian clustering. All clustering approaches supported a division of the 28 populations into 4 major groups for which analysis of molecular variance confirmed a significant variation among groups (34.3 %). These population genetic parameters suggest limited gene flow among populations and evidence for isolation by distance (r = 0.272, P < 0.0001) was found in this study. Altitude, AMT and AMP explained 9.5, 11.5 and 14.0 % of the total variance among populations indicating that eco-geographic factors have a significant effect. Considering the low within-population genetic diversity, high differentiation among populations and the increasing anthropogenic pressure on the species, in situ conservation measures were recommended to preserve L. pumilum in Qinghai–Tibet plateau.

Population Genetic Structure of Laurus nobilis L. Inferred From Transferred Nuclear Microsatellites

Abstract Species with fragmented populations and low population size often display low within-population genetic diversity and strong among-population differentiation. Laurus nobilis L. (Lauraceae), common laurel, has a scattered distribution throughout the Mediterranean, with only few autochthonous populations. Our goal was to elucidate if this species has range-wide genetic structure and if planted material can be traced back to its origin. Genetic diversity was investigated using 4 polymorphic nuclear microsatellites (nSSR) transferred from two species of Lauraceae. Sixty-six laurel trees were selected from 7 widely separated populations within the Mediterranean distribution area of the species. A total of 34 alleles (9 alleles per locus on average) were found. Mean genetic diversity within-population (Hs), was 0.558. Genetic differentiation among populations (GST = 0.243) was high compared to that of other angiosperms. Laurus nobilis can be separated into two main gene pools, one from western (Tunisia, Algeria and France) and the other from eastern Mediterranean (Turkey). The Algerian, Tunisian and French populations presented a strong genetic similarity, compatible with the fact that North African laurel populations could be recently introduced from north-western Mediterranean stock.

Genetic races associated with the genera and sections of host species in the holoparasitic plant Cytinus (Cytinaceae) in the Western Mediterranean basin

* Speciation via race formation is an important evolutionary process in parasites, producing changes that favour their development on particular host species. Here, the holoparasitic plant Cytinus, which has diverse host species in the family Cistaceae, has been used to study the occurrence of such races. * Amplified fragment length polymorphism (AFLP) analyses were performed on 174 individuals of 22 populations parasitizing 10 Cistaceae species in the Western Mediterranean basin. * Neighbour-joining, multivariate ordination analyses, and individual-based Bayesian analyses, clustered Cytinus populations into five well-characterized genetic races that, overall, agreed with the taxonomic sections of their hosts. In the AMOVA, among-races differences accounted for almost 50% of the genetic variation. The isolation-by-distance model was not supported by a Mantel test among Cytinus populations (r = 0.012; P = 0.456). All races showed low within-population genetic diversity, probably as a result of restricted pollen flow aggravated by flowering asynchrony, restricted seed dispersion, or stochastic processes. * The genetic differentiation among the five races of Cytinus is congruent with the view that these races are well-characterized lineages that have evolved independently as a result of selective pressures imposed by their hosts. This pattern, with genetically distinctive groups associated with the infrageneric sections of the host species, has not been reported previously for parasitic angiosperms.

Loss of genetic variation in geographically marginal populations of Atriplex tatarica (Chenopodiaceae).

Genetic variability was estimated for Atriplex tatarica from 25 populations in the Czech Republic. Since its north-western range margin is in central Europe, a relationship between marginality and low within-population genetic diversity was tested in accordance with the Central-Marginal Model. Population genetic diversity was expressed by assessing patterns of variation at 13 putatively neutral allozyme loci (comprising 30 putative alleles) within and between 25 natural populations of A. tatarica along a north-west-south-east transect in the Czech Republic. Atriplex tatarica is a species of human-made habitats with a mixed mating system and wide geographic distribution. Overall, A. tatarica displayed moderate levels of genetic diversity in comparison with other herbaceous plants. The percentage of loci that were polymorphic was 47.1%, with average values of 1.55, 0.151 and 0.155 for the average number of alleles per polymorphic locus (A), observed heterozygosity (Ho) and expected heterozygosity (He), respectively. There was only weak evidence of inbreeding within populations (FIS=0.031) and significant population differentiation (FST=0.214). Analysis of the data provides no evidence for isolation-by-distance for the whole study area. However, Mantel tests were highly significant for the marginal Bohemian region and non-significant for the central Moravian region. While northern populations of A. tatarica showed significantly lower allelic richness (A=1.462) than populations from the southern part of the study area (A=1.615), they did not differ in observed heterozygosity (Ho), gene diversity (HS), inbreeding within populations (FIS) or population differentiation (FST), despite generally lower values of particular genetic measurements in the marginal region. Genetic diversity, with the exception of allelic richness, was not significantly lower at the margins of the species' range. This, therefore, provides only weak support for the predictions of the Central-Marginal Model.

Allozyme diversity and genetic structure of marginal and central populations of Corylus avellana L. (Betulaceae) in Europe

Corylus avellana L. (hazel, Betulaceae) is a long-lived, widespread shrub in Europe, having its northern range margin in Fennoscandia and a postglacial history involving range-expansion from refugial areas in southern Europe. In this study, we tested for a relationship between marginality and low within-population genetic diversity by assessing patterns of variation at 14 putatively neutral allozyme loci (comprising 43 putative alleles) within and between 40 natural populations of C. avellana along a north-south transect in Europe. Geographically marginal populations (central Sweden) showed lower levels of within-population diversity than populations in more central regions, as indicated by significant negative correlations between latitude and the percentage of polymorphic loci (rS=–0.47, P < 0.001), the average number of alleles per locus (rS=–0.65, P < 0.001), the expected heterozygosity (rS=–0.19, P < 0.05), and the proportion of distinguishable genotypes (rS=–0.56, P < 0.001). These patterns, combined with the unusually high between-population component of gene diversity (GST=19.7%) and allelic richness (AST=24%) in the marginal region, can be attributed to historical bottlenecks during the species’ postglacial range-expansion, but may also reflect a history of genetic drift in the small, isolated populations occupying the marginal region. Information on the spatial distribution of genotypes provide further support for a role of vegetative reproduction (layering) in the structuring of genetic variation within populations.