Microsatellite diversity and genetic structure of fragmented populations of the rare,fire-dependent shrub Grevillea macleayana

Recent habitat loss and fragmentation superimposed upon ancient patterns of population subdivision are likely to have produced low levels of neutral genetic diversity and marked genetic structure in many plant species. The genetic effects of habitat fragmentation may be most pronounced in species that form small populations, are fully self-compatible and have limited seed dispersal. However, long-lived seed banks, mobile pollinators and long adult lifespans may prevent or delay the accumulation of genetic effects. We studied a rare Australian shrub species, Grevillea macleayana (Proteaceae), that occurs in many small populations, is self-compatible and has restricted seed dispersal. However, it has a relatively long adult lifespan (c. 30 years), a long-lived seed bank that germinates after fire and is pollinated by birds that are numerous and highly mobile. These latter characteristics raise the possibility that populations in the past may have been effectively large and genetically homogeneous. Using six microsatellites, we found that G. macleayana may have relatively low within-population diversity (3.2-4.2 alleles/locus; Hexp = 0.420-0.530), significant population differentiation and moderate genetic structure (FST = 0.218) showing isolation by distance, consistent with historically low gene flow. The frequency distribution of allele sizes suggest that this geographical differentiation is being driven by mutation. We found a lack mutation-drift equilibrium in some populations that is indicative of population bottlenecks. Combined with evidence for large spatiotemporal variation of selfing rates, this suggests that fluctuating population sizes characterize the demography in this species, promoting genetic drift. We argue that natural patterns of pollen and seed dispersal, coupled with the patchy, fire-shaped distribution, may have restricted long-distance gene flow in the past.     

Understanding the population genetic structure of Gleditsia triacanthos L.: seed dispersal and variation in female reproductive success

Fine-scale genetic structuring is influenced by a variety of ecological factors and can directly affect the evolutionary dynamics of plant populations by influencing effective population size and patterns of viability selection. In many plant species, genetic structuring within populations may result from highly localized patterns of seed dispersal around maternal plants or by the correlated dispersal and recruitment of siblings from the same fruit. This fine-scale genetic structuring may be enhanced if female parents vary significantly in their reproductive success. To test these hypotheses, we used genetic data from 17 allozyme loci and a maximum-likelihood, ‘maternity-analysis’ model to estimate individual female fertilities for maternal trees across a large number of naturally established seedlings and saplings in two populations of Gleditsia triacanthos L. (Leguminosae). Maximum-likelihood fertility estimates showed that the three highest fertility females accounted for 58% of the 313 progeny at the first site and 46% of the 651 progeny at the second site, whereas 18 of 35 and 16 of 34 females, respectively, had fertility estimates that did not exceed 1%. Additional analyses of the second site found individual female fertility to vary significantly both within and among juvenile age classes. Female fertility at the first site was weakly correlated with maternal tree size and spatial location relative to the open, old-field portions of the population, where the great majority of seedlings and saplings were growing, but no such correlations were found at the second site. Estimates of realized seed dispersal distances indicated that dispersal was highly localized at the first site, but was nearly random at the second site, possibly reflecting differences between the two sites in the behaviour of animal dispersers. The combined estimates of seed dispersal patterns and fertility variation are sufficient to explain previously described patterns of significant fine-scale spatial genetic structure in these two populations. In general, our results demonstrate that effective seed dispersal distributions may vary significantly from population to population of a species due to the unpredictable behaviour of secondary dispersers. Consequently, the effects of seed dispersal on realized fine-scale genetic structure may also be relatively unpredictable.     

Quantifying gene flow from spatial genetic structure data in a metapopulation of Chamaecrista fasciculata (Leguminosae)

Abstract An extensive allozyme survey was conducted within a natural "meta" population of the native North American annual legume, Chamaecrista fasciculata (Leguminosae) to quantify genetic structure at different spatial scales. Gene flow was then estimated by a recently developed indirect method based on a continuous population model, using pairwise kinship coefficients between individuals. The indirect estimates of gene flow, quantified in terms of neighborhood size, with an average value on the order of 150 individuals, were concordant among different spatial scales (subpopulation, population, metapopulation). This gene-flow value lies within the range of direct estimates previously documented from observations of pollen and seed dispersal for the same metapopulation. Monte Carlo simulations using the direct measures of gene flow as parameters further demonstrated that the observed spatial pattern of allozyme variation was congruent with a model of isolation by distance. Combining previously published estimates of pollen dispersal distances with kinship coefficients from this study, we quantified biparental inbreeding relative to either a single subpopulation or the whole metapopulation. At the level of a neighborhood, little biparental inbreeding was observed and most departure from Hardy-Weinberg genotypic proportions was explained by self-fertilization, whereas both selfing and biparental inbreeding contributed to nonrandom mating at the metapopulation level. Gene flow was also estimated from indirect methods based on a discontinuous population structure model. We discuss these results with respect to the effect of a patchy population structure on estimation of gene flow.     

Contrasting patterns of pollen and seed flow influence the spatial genetic structure of sweet vernal grass (Anthoxanthum odoratum) populations

The spatial genetic structure of plant populations is determined by a combination of gene flow, genetic drift, and natural selection. Gene flow in most plants can result from either seed or pollen dispersal, but detailed investigations of pollen and seed flow among populations that have diverged following local adaptation are lacking. In this study, we compared pollen and seed flow among 10 populations of sweet vernal grass (Anthoxanthum odoratum) on the Park Grass Experiment. Overall, estimates of genetic differentiation that were based on chloroplast DNA (cpDNA) and, which therefore resulted primarily from seed flow, were lower (average F(ST) = 0.058) than previously published estimates that were based on nuclear DNA (average F(ST) = 0.095). Unlike nuclear DNA, cpDNA showed no pattern of isolation by adaptation; cpDNA differentiation was, however, inversely correlated with the number of additions (nutrients and lime) that each plot had received. We suggest that natural selection is restricting pollen flow among plots, whereas nutrient additions are increasing seed flow and genetic diversity by facilitating the successful germination and growth of immigrant seeds. This study highlights the importance of considering all potential gene flow mechanisms when investigating determinants of spatial genetic structure, and cautions against the widespread assumption that pollen flow is more important than seed flow for population connectivity in wind-pollinated species.     

Migration load in plants: role of pollen and seed dispersal in heterogeneous landscapes

Evolution of local adaptation depends critically on the level of gene flow, which, in plants, can be due to either pollen or seed dispersal. Using analytical predictions and individual-centred simulations, we investigate the specific influence of seed and pollen dispersal on local adaptation in plant populations growing in patchy heterogeneous landscapes. We study the evolution of a polygenic trait subject to stabilizing selection within populations, but divergent selection between populations. Deviations from linkage equilibrium and Hardy-Weinberg equilibrium make different contributions to genotypic variance depending on the dispersal mode. Local genotypic variance, differentiation between populations and genetic load vary with the rate of gene flow but are similar for seed and pollen dispersal, unless the landscape is very heterogeneous. In this case, genetic load is higher in the case of pollen dispersal, which appears to be due to differences in the distribution of genotypic values before selection.     

DIRECT AND INDIRECT ESTIMATES OF NEIGHBORHOOD AND EFFECTIVE POPULATION SIZE IN A TROPICAL PALM,ASTROCARYUM MEXICANUM

To estimate the relative importance of genetic drift, the effective population size ???(N_e) can be used. Here we present estimates of the effective population size and related measures in Astrocaryum mexicanum, a tropical palm from Los Tuxtlas rain forest, Veracruz, Mexico. Seed and pollen dispersal were measured. Seeds are primarily dispersed by gravity and secondarily dispersed by small mammals. Mean primary and secondary dispersal distances for seeds were found to be small (0.78 m and 2.35 m, respectively). A. mexicanum is beetle pollinated and pollen movements were measured by different methods: a) using fluorescent dyes, b) as the minimum distance between active female and male inflorescences, and c) using rare allozyme alleles as genetic markers. All three estimates of pollen dispersal were similar, with a mean of approximately 20 m. Using the seed and pollen dispersal data, the genetic neighborhood area (A) was estimated to be 2,551 m². To obtain the effective population size, three different overlapping generation methods were used to estimate an effective density with demographic data from six permanent plots. The effective density ranged from 0.040 to 0.351 individuals per m². The product of effective density and neighborhood area yields a direct estimate of the neighborhood effective population size (N_b). N_b ranged from 102 to 895 individuals. Indirect estimates of population size and migration rate (Nm) were obtained using F_st for five different allozymic loci for both adults and seeds. We obtained a range of Nm from 1.2 to 19.7 in adults and a range of Nm from 4.0 to 82.6 for seeds. We discuss possible causes of the smaller indirect estimates of Nm relative to the direct and compare our estimates with values from other plant populations. Gene dispersal distances, neighborhood size, and effective population size in A. mexicanum are relatively high, suggesting that natural selection, rather than genetic drift, may play a dominant role in patterning the genetic variation in this tropical palm.     

Using genetic markers to directly estimate gene flow and reproductive success parameters in plants on the basis of naturally regenerated seedlings

Estimating seed and pollen gene flow in plants on the basis of samples of naturally regenerated seedlings can provide much needed information about "realized gene flow," but seems to be one of the greatest challenges in plant population biology. Traditional parentage methods, because of their inability to discriminate between male and female parentage of seedlings, unless supported by uniparentally inherited markers, are not capable of precisely describing seed and pollen aspects of gene flow realized in seedlings. Here, we describe a maximum-likelihood method for modeling female and male parentage in a local plant population on the basis of genotypic data from naturally established seedlings and when the location and genotypes of all potential parents within the population are known. The method models female and male reproductive success of individuals as a function of factors likely to influence reproductive success (e.g., distance of seed dispersal, distance between mates, and relative fecundity--i.e., female and male selection gradients). The method is designed to account for levels of seed and pollen gene flow into the local population from unsampled adults; therefore, it is well suited to isolated, but also wide-spread natural populations, where extensive seed and pollen dispersal complicates traditional parentage analyses. Computer simulations were performed to evaluate the utility and robustness of the model and estimation procedure and to assess how the exclusion power of genetic markers (isozymes or microsatellites) affects the accuracy of the parameter estimation. In addition, the method was applied to genotypic data collected in Scots pine (isozymes) and oak (microsatellites) populations to obtain preliminary estimates of long-distance seed and pollen gene flow and the patterns of local seed and pollen dispersal in these species.     

Dispersal, gene flow, and population structure

The accuracy of gene flow estimates is unknown in most natural populations because direct estimates of dispersal are often not possible. These estimates can be highly imprecise or even biased because population genetic structure reflects more than a simple balance between genetic drift and gene flow. Most of the models used to estimate gene flow also assume very simple patterns of movement. As a result, multiple interpretations of population structure involving contemporary gene flow, departures from equilibrium, and other factors are almost always possible. One way to isolate the relative contribution of gene flow to population genetic differentiation is to utilize comparative methods. Population genetic statistics such as FST, heterozygosity and Nei's D can be compared between species with differing dispersal abilities if these species are otherwise phylogenetically, geographically and demographically comparable. Accordingly, the available literature was searched for all groups that meet these criteria to determine whether broad conclusions regarding the relationships between dispersal, population genetic structure, and gene flow estimates are possible. Allozyme and mtDNA data were summarized for 27 animal groups in which dispersal differences can be characterized. In total, genetic data were obtained for 333 species of vertebrates and invertebrates from terrestrial, freshwater and marine habitats. Across these groups, dispersal ability was consistently related to population structure, with a mean rank correlation of -0.72 between ranked dispersal ability and FST. Gene flow estimates derived from private alleles were also correlated with dispersal ability, but were less widely available. Direct-count heterozygosity and average values of Nei's D showed moderate degrees of correlation with dispersal ability. Thus, despite regional, taxonomic and methodological differences among the groups of species surveyed, available data demonstrate that dispersal makes a measurable contribution to population genetic differentiation in the majority of animal species in nature, and that gene flow estimates are rarely so overwhelmed by population history, departures from equilibrium, or other microevolutionary forces as to be uninformative.     

Natural selection on body size is mediated by multiple interacting factors: a comparison of beetle populations varying naturally and experimentally in body size

Body size varies considerably among species and among populations within species, exhibiting many repeatable patterns. However, which sources of selection generate geographic patterns, and which components of fitness mediate evolution of body size, are not well understood. For many animals, resource quality and intraspecific competition may mediate selection on body size producing large-scale geographic patterns. In two sequential experiments, we examine how variation in larval competition and resource quality (seed size) affects the fitness consequences of variation in body size in a scramble-competing seed-feeding beetle, Stator limbatus. Specifically, we compared fitness components among three natural populations of S. limbatus that vary in body size, and then among three lineages of beetles derived from a single base population artificially selected to vary in size, all reared on three sizes of seeds at variable larval density. The effects of larval competition and seed size on larval survival and development time were similar for larger versus smaller beetles. However, larger-bodied beetles suffered a greater reduction in adult body mass with decreasing seed size and increasing larval density; the relative advantage of being large decreased with decreasing seed size and increasing larval density. There were highly significant interactions between the effects of seed size and larval density on body size, and a significant three-way interaction (population-by-density-by-seed size), indicating that environmental effects on the fitness consequences of being large are nonadditive. Our study demonstrates how multiple ecological variables (resource availability and resource competition) interact to affect organismal fitness components, and that such interactions can mediate natural selection on body size. Studying individual factors influencing selection on body size may lead to misleading results given the potential for nonlinear interactions among selective agents.     

Fine-scale genetic structure of the common Primula elatior (Primulaceae) at an early stage of population fragmentation

Many rare species are threatened by habitat fragmentation; however, less is known about effects of fragmentation on common species, despite their potential role in ecosystem productivity and functioning. We identified key factors and processes influencing gene flow in a large population of Primula elatior, a common distylous perennial herb, at an early stage of the fragmentation process, i.e., when fragmentation is taking place. Using 19 allozyme loci, we investigated genetic variation and fine-scale spatial genetic structure (SGS) at seedling and adult life stages in relation to fragmentation history (recent bottlenecks), selection, clonal propagation, sexual reproduction (seed and pollen dispersal, distyly), and patchy structure (patch size, plant density, and morph ratio). The main factors contributing to the strong SGS are seed and (to a lesser extent) pollen dispersal, through a spatial Wahlund effect and biparental inbreeding. Significant differences in allele frequencies between seedlings and adults indicate a temporal Wahlund effect. Patch plant density and biased morph ratio also affect the genetic patterns. Our results show that if P. elatior populations evolve into patchworks of small, isolated remnants, genetic erosion, reduced gene flow, and increased inbreeding can be expected, suggesting that such common plant species might require large population sizes to remain viable.     

POPULATION STRUCTURE AND LEVELS OF GENE FLOW IN THE MEDITERRANEAN LAND SNAIL ALBINARIA CORRUGATA (PULMONATA: CLAUSILIIDAE)

The amount of gene flow among local populations partly determines the relative importance of genetic drift and natural selection in the differentiation of such populations. Land snails, because of their limited powers for dispersal, may be particularly likely to show such differentiation. In this study, we directly estimate gene flow in Albinaria corrugata, a sedentary, rock-dwelling gastropod from Crete, by mark-recapture studies. In the same area, 23 samples were taken and studied electrophoretically for six polymorphic enzyme loci. The field studies indicate that the population structure corresponds closely to the stepping-stone model: demes are present on limestone boulders that are a few meters apart, and dispersal takes place mainly between adjacent demes. Average deme size (N) is estimated at 29 breeding individuals and the proportion of migrants per generation at 0.195 (Nm = 5.7). We find no reason to assume long-distance dispersal, apart from dispersal along occasional stretches of suitable habitat. Genetic subdivision of the population, as derived from F_ST values, corresponds to the direct estimate only at the lowest spatial level (distance between sample sites < 10 m), where values for Nm of 5.4 and 17.6 were obtained. In contrast, at the larger spatial scales, F_ST values give gene-flow estimates that are incompatible with the expected amount of gene flow at these scales. We explain these discrepancies by arguing that gene flow is in fact extremely limited, making correct estimates of Nm from F_ST impossible at the larger spatial scales. In view of these low levels of gene flow, it is concluded that both genetic drift and natural selection may play important roles in the genetic differentiation of this species, even at the lowest spatial scales.     

Gene flow and the measurement of dispersal in plant populations

Despite theoretical assumptions, estimates of dispersal distance in natural plant populations have only recently been undertaken. Since plants are non-motile, gene flow depends on the dispersal of pollen and seed. Methods of estimating pollen and seed dispersal are reviewed and their implications for studies of evolutionary processes discussed. Practical advice is provided for designing exercises suitable for estimating dispersal distance in natural plant populations.     

Gene flow and melanism in garter snakes revisited: a comparison of molecular markers and island vs. coalescent models

Within populations, the stochastic effect of genetic drift and deterministic effect of natural selection are potentially weakened or altered by gene flow among populations. The influence of gene flow on Lake Erie populations of the common garter snake has been of particular interest because of a discontinuous colour pattern polymorphism (striped vs. melanistic) that is a target of natural selection. We reassessed the relative contributions of gene flow and genetic drift using genetic data and population size estimates. We compared all combinations of two marker systems and two analytical approaches to the estimation of gene flow rates: allozymes (data previously published), microsatellite DNA (new data), the island model ( F _ST-based approach), and a coalescence-based approach. For the coalescence approach, mutation rates and sampling effects were also investigated. While the two markers produced similar results, gene flow based on F _ST was considerably higher (Nm > 4) than that from the coalescence-based method (Nm < 1). Estimates of gene flow are likely to be inflated by lack of migration-drift equilibrium and changing population size. Potentially low rates of gene flow (Nm < 1), small population size at some sites, and positive correlations of number of microsatellite DNA alleles and island size and between M , mean ratio of number of alleles to range in allele size, and island size suggest that in addition to selection, random genetic drift may influence colour pattern frequencies. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79, 389–399.     

How species evolve collectively: implications of gene flow and selection for the spread of advantageous alleles

The traditional view that species are held together through gene flow has been challenged by observations that migration is too restricted among populations of many species to prevent local divergence. However, only very low levels of gene flow are necessary to permit the spread of highly advantageous alleles, providing an alternative means by which low-migration species might be held together. We re-evaluate these arguments given the recent and wide availability of indirect estimates of gene flow. Our literature review of F(ST) values for a broad range of taxa suggests that gene flow in many taxa is considerably greater than suspected from earlier studies and often is sufficiently high to homogenize even neutral alleles. However, there are numerous species from essentially all organismal groups that lack sufficient gene flow to prevent divergence. Crude estimates on the strength of selection on phenotypic traits and effect sizes of quantitative trait loci (QTL) suggest that selection coefficients for leading QTL underlying phenotypic traits may be high enough to permit their rapid spread across populations. Thus, species may evolve collectively at major loci through the spread of favourable alleles, while simultaneously differentiating at other loci due to drift and local selection.     

COLOR-PATTERN VARIATION IN LAKE ERIE WATER SNAKES: THE ROLE OF GENE FLOW

In an effort to clarify the evolutionary processes influencing color-pattern variation in Lake Erie island water snake (Nerodia sipedon) populations, rates of gene flow among island and mainland populations were estimated from patterns of allozymic variation detected using electrophoresis. Rates of gene flow were high with Nm, the number of migrants per generation, averaging 25.5 among island sites, 9.2 between the Ontario mainland and the islands, and 3.6 between the Ohio mainland and the islands. Based on estimates of current population size from mark-recapture work and of past population size extrapolated from the extent of shoreline habitat, values of m between island and mainland populations ranged from 0.0008–0.01. Synthesis of estimates of the rate of gene flow with information on inheritance of color pattern, the strength of natural selection, and population history supports the hypothesis that color-pattern variation in island populations results from a balance between gene flow and natural selection. However, depending on the mode of inheritance of color pattern, stochastic processes such as drift may have been important in the initial stages of differentiation between island and mainland populations.     

Flow regulation affects temporal patterns of riverine plant seed dispersal: potential implications for plant recruitment

1. Changes to the natural flow regime of a river caused by flow regulation may affect waterborne seed dispersal (hydrochory), and this may be an important mechanism by which regulation affects riverine plant communities. We assessed the effect of altered timing of seasonal flow peaks on hydrochory and considered the potential implications for plant recruitment. 2. We sampled hydrochory within five lowland rivers of temperate Australia, three of which are regulated by large dams. These dams are operated to store winter and spring rains and release water in summer and autumn for agriculture. At three sites on each river, hydrochory was sampled monthly for 12 months using passive drift nets. The contents of the drift samples were determined using the seedling‐emergence method. 3. More than 33 000 seedlings from 142 taxa germinated from the samples. In general, more seeds and taxa were observed in the drift at higher flows. By altering the period of peak flows from winter–spring to summer–autumn, flow regulation similarly affected the period of peak seed dispersal. The effect of regulation on seed dispersal varied between taxa depending on their timing of seed release and whether or not they maintain a persistent soil seed bank. 4. Hydrochory in rivers is a product of flow regime and the life history of plants. By altering natural flow regimes and thus hydrochorous dispersal patterns, flow regulation is likely to affect adversely the recruitment of native plant species with dispersal phenologies adapted to natural flow regimes (such as many riparian trees and shrubs) and encourage the spread of non‐native (exotic) species. 5. Changes to hydrochorous dispersal patterns are an important mechanism by which altered flow timing affects riverine plant communities. Natural seasonal flow peaks (in this case spring) are likely to be important for the recruitment of many native riparian woody taxa.     

LONG-DISTANCE GENE FLOW IN THE SEDENTARY BUTTERFLY,EUPHILOTES ENOPTES (LEPIDOPTERA: LYCAENIDAE)

The relationship between gene flow and geographic proximity has been assessed for many insect species, but dispersal distances are poorly known for most of these. Thus, we are able to assess the concordance between vagility and gene flow for only a few species. In this study, I documented variation at six allozyme loci among Washington and Oregon populations of the sedentary, patchily distributed, lycaenid butterfly, Euphilotes enoptes (Boisduval) to assess whether the relationship between gene flow and geographic distance is consistent with the dispersal biology of this species. Both a phenogram based on genetic distances between populations and a regression analysis of gene flow estimates on geographic distances showed a pattern consistent with genetic isolation by distance. Many estimates of gene flow among pairs of populations separated by more than 100 km exceeded the equivalent of 10 individuals exchanged per generation, a value much greater than would be predicted from the limited dispersal ability of this species. However, based on the allozyme data, genetic neighborhood size was estimated to be approximately 39 individuals, a value that is consistent with poor vagility. The results of this study speak to the power of stepping-stone gene flow among populations and are compared to the results of other studies that have examined the relationship between dispersal and gene flow in sedentary insects.     

Long‐term ‘islands’ in the landscape: low gene flow,effective population size and genetic divergence in the shrub Hakea oldfieldii (Proteaceae)

The genetic structure of disjunct populations is determined by founding genetic properties, demographic processes, gene flow, drift and local selection. We aim to identify the genetic consequences of natural population disjunction at regional and local scales in Hakea oldfieldii using nuclear and plastid markers to investigate long‐term effective population sizes and gene flow, and patterns of diversity and divergence, among populations. Regional divergence was significant as shown by a consistent pattern in principal coordinates, neighbor‐joining and Bayesian analyses, but divergence at the local level was also significant with localized distribution of plastid haplotypes and populations clustering separately in Bayesian analyses. Historical, recent and first‐generation gene flow was low, suggesting that recent habitat fragmentation has not reduced gene migration significantly. Genetic bottlenecks were detected in three populations. Long‐term effective population size was significantly correlated with the number of alleles/locus and observed heterozygosity, but not with census population size, suggesting that the loss of diversity is associated with long‐term changes rather than recent fragmentation. Inbreeding coefficients were significant in only three populations, suggesting that the loss of diversity is linked to drift and bottlenecks associated with demographic processes (local extinction by fires) rather than inbreeding. Historical disjunction as a result of specific ecological requirements, contraction of habitats following drying during the Pleistocene, low gene flow and changes in population size are likely to have been important forces driving divergence through isolation by distance and drift. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 319–334.     

Gene flow and effective population size in Lepanthes (Orchidaceae): a case for genetic drift

Genetic drift can play an important role in population differentiation, particularly when effective population sizes are small and gene flow is limited. Such conditions are suspected to be common in the species-rich Orchidaceae. We investigated the likelihood of genetic drift in natural populations of three endemic species of Lepanthes (Orchidaceae) from Puerto Rico. We estimated effective population size, Ne, using three ecologically based methods. Two of the three estimates were based on variance in reproductive potential and the third was based on coalescence time. All estimates of Ne were usually <40% of the standing population size, resulting in values of <20 individuals per population. Based on starch gel electrophoresis of isozymes, Nm estimates suggest restricted gene flow among populations in the range of one or less successful migrant per generation. Genetic differentiation among populations is expected under such conditions from random genetic drift. Indeed we observed high genetic differentiation among populations (L. rubripetala, F_ST> G_ST, 6; θ.248, 0.266, 0.293; L. rupestris, 0.148, 0.169, 0.138; L. eltoroensis, 0.251, 0.219, 0.218, respectively). Genetic drift is likely to be important for population differentiation in Lepanthes as a result of small effective population sizes and restricted gene flow.     

Fine-Scale Spatial Genetic Structure in Emmer Wheat and the Role of Population Range Position

The extent of spatial genetic structure (SGS) within plant populations depends on seed and pollen dispersal distance, breeding type, level of self-fertilization and effective plant density. Self-fertilizing species with gravity-dispersed seeds are expected to have both small effective population sizes and low pollen movement leading to high genetic structure. Higher SGS can be expected in more patchy and peripheral populations because of lower plant density and population sizes, and lower intensity of gene flow. We tested these predictions analyzing SGS in two core and two peripheral populations of predominantly self-fertilizing emmer wheat. Analysis of SGS with 11 nuclear microsatellites revealed (1) a negative linear relationship between kinship coefficients, calculated for pairs of individuals, and the logarithm of geographical distance between members of the pairs, in all studied populations; and (2) a significant autocorrelation for a distance up to 5 m (core populations) or 20 m (peripheral populations). Pollen flow, estimated from comparison of nuclear and chloroplast variation, was spatially limited, as was seed dispersal. Our results support a hypothesized relationship between SGS intensity and breeding system, the mode of seed dispersal and the population range position (core vs. periphery).