Fine-scale clonal structure and diversity within patches of a clone-forming dioecious shrub, Ilex leucoclada (Aquifoliaceae)

BACKGROUND AND AIMS: The mode of reproduction (sexual vs. asexual) is likely to have important effects on genetic variation and its spatial distribution within plant populations. An investigation was undertaken of fine-scale clonal structure and diversity within patches of Ilex leucoclada (a clone-forming dioecious shrub). METHODS: Six patches were selected in a 1-ha plot previously established in an old-growth beech forest. Two of the selected patches were composed predominantly of stems with male flowers (male patch), and two contained stems with predominantly female flowers (female patch). The remaining two patches contained stems with male flowers and stems with female flowers in more or less equal proportions (mixed patch). Different genets were distinguished using random amplified polymorphic DNA (RAPD) markers. KEY RESULTS: One hundred and fifty-six genets with different RAPD phenotypes were identified among 1928 stems from the six patches. Among the six patches, the male patches had the lowest clonal diversity, and the mixed patches had the highest. Distribution maps of the genets showed that they extended downhill, reflecting natural layering that occurred when stems were pressed to the ground by heavy snow. In every patch, there were a few large genets with many stems and many small genets with a few stems. CONCLUSION: The differences in clonal diversity among patches may be due to differences in seedling recruitment frequencies. The skewed distribution of genet size (defined as the number of stems per genet) within patches may be due to differences in the timing of germination, or age (with early-establishing genets having clear advantages for acquiring resources) and/or intraspecific competition.     

Sex-different response in growth traits to resource heterogeneity explains male-biased sex ratio

In dioecious plants, differences in growth traits between sexes in a response to micro-environmental heterogeneity may affect sex ratio bias and spatial distributions. Here, we examined sex ratios, stem growth traits and spatial distribution patterns in the dioecious clonal shrub Aucuba japonica var. borealis, in stands with varying light intensities. We found that male stems were significantly more decumbent (lower height/length ratio) but female stems were upright (higher height/length ratio). Moreover, we found sex-different response in stem density (no. of stems per unit area) along a light intensity gradient; in males the stem density increased with increases in canopy openness, but not in females. The higher sensitivity of males in increasing stem density to light intensity correlated with male-biased sex ratio; fine-scale sex ratio was strongly male-biased as canopy openness increased. There were also differences between sexes in spatial distributions of stems. Spatial segregation of sexes and male patches occupying larger areas than female patches might result from vigorous growth of males under well-lit environments. In summary, females and males showed different growth responses to environmental variation, and this seemed to be one of possible causes for the sex-differential spatial distributions and locally biased sex ratios.     

Intrapopulation sex ratio variation in the salt grass Distichlis spicata

ABSTRACT In many dioecious plant populations, males and females appear to be spatially segregated, a pattern that is difficult to explain given its potentially high costs. However, in asexually propagating species, spatial segregation of the sexes may be indistinguishable from superficially similar patterns generated by random establishment of a few genets followed by extensive clonal spread and by gender-specific differences in rates of clonal spread. In populations where a significant fraction of individuals are not flowering and gender cannot be assigned to this fraction, apparent spatial segregation of the sexes may be due to differential flowering between the sexes. We confirm reports that flowering ramets of the clonal, perennial grass Distichlis spicata are spatially segregated by sex. We extend these studies in two fundamental ways and demonstrate that this species exhibits true spatial segregation of the sexes. First, using RAPD markers, we estimated that at least 50% of ramets in patches with biased sex ratios represent distinct genotypes. Second, we identified a RAPD marker linked to female phenotype (eliminating the possibility that gender is environmentally determined) and used it to show that the majority of patches exhibit significantly biased sex ratios for both ramets and genets, regardless of flowering status.     

Flower bud abortion influences clonal growth and sexual dimorphism in the understorey dioecious shrub Aucuba japonica (Cornaceae)

Sexual differences were investigated to determine the significance of flower bud abortion in the dioecious shrub Aucuba japonica Thunb. The mean number of flowers per inflorescence and the mean number of flowering inflorescences (as opposed to aborted inflorescences) per individual were greater in males than in females in 1997 and 1998. Reproductive investment by males was 0.4-times (1997) and 1.4-times (1998) that by females. In addition, females aborted 30.9% (1997) and 42.7% (1998) of their total flower buds without blooming, whereas no male flower buds aborted. One of the architectural traits of this shrub is that in the year that a flower bud is produced at the shoot apex, the shoot will branch into two or more shoots. Thus, there was less sexual difference in the number of current shoots per individual than there was in the number of flowering inflorescences. The relationship between annual growth and reproduction, and the probability of reproduction in the following year, suggested that the higher investment in female reproduction was manifested as a cost for reproductive frequency rather than as a cost for annual growth. The spatial distribution of both males and females was clumped, which may be the result of clonal growth. In addition, overall sex ratios were not skewed and the number of sprouts did not differ significantly between sexes. These results suggested that flower bud abortion by females might reduce sexual dimorphism in terms of clonal growth.     

Sex ratio variation in dioecious plant species: A comparative ecological study of six species of Lindera (Lauraceae)

Sex ratio variation was investigated in natural populations of six dioecious shrub species of Lindera in Japan. Interspecific differences in sex ratio were examined in relation to patterns of population structure, floral dimorphism, fruit production and intersexual differences in herbivory. Sex ratios tended towards equality or bias in favour of males, except for populations of L. glauca , in which no male plants were found. Sex switching in individual plants was not observed. Although male flowers were generally larger in size than flowers of conspecific females, costs of flowering did not appear to influence sex ratio. However, a relation was found between high allocation to fruit production and increasing male fraction of populations. In addition, girths of flowering plants tended to be larger in females, which may indicate delay in age of the first reproduction compared to males. No intersexual difference was found in degree of leaf damage due to herbivory. Results of this study are discussed in the light of other studies on sex ratio variation in dioecious plant species.     

Nutrient deficiency promotes male-biased apparent sex ratios at the ramet level in the dioecious plant <Emphasis Type="Italic">Myrica gale</Emphasis> var. <Emphasis Type="Italic">tomentosa</Emphasis> in oligotrophic environments in bogs

In populations of dioecious plants, the differences in the cost of reproduction between male and female plants can promote a male-biased sex ratio. In this study, we examine the macronutrient levels in tissues of the dioecious wetland shrub Myrica gale to identify the cost of reproduction for male and female plants and to examine the effect of nutrients on the apparent sex ratio at the ramet level. We examined plants across 12 populations of M. gale inhabiting bogs and fens in Japan. For each population, we used line transects to estimate the apparent sex ratio and measured the concentrations of nitrogen (N), phosphorus (P), and potassium (K) in the leaves sampled from male and female plants and in the fruits from female plants. For five of the populations, we calculated the flowering frequency, mortality, and the recruitment rate (as the rate of clonal propagation). We found that the proportion of females was positively affected, and the male bias of sex ratios reduced, by increases in P concentration in leaves sampled from female plants. Neither mortality nor recruitment was affected by sex or by the nutrient concentration (P, K). The flowering frequency was not affected by sex or by K concentration, but decreased with decreases in the P concentration measured in leaves. This study confirmed that reproduction in M. gale is P-limited. We found no distinct differences in the flowering frequency, mortality, or recruitment rate between the male and female plants.     

Sexual dimorphism and sex ratios of two Australian dioecious species of alpine pineapple grass, Astelia alpina var. novaehollandiae and Astelia psychrocharis (Asteliaceae)

Sexual dimorphism and male biased sex ratios have been predicted for dioecious plants experiencing the limited conditions for growth and reproduction found in many alpine environments. To test these predictions, the reproductive ecology of two congeneric, co-occurring, dioecious, clonal, species was examined in the subalpine and alpine zones of Kosciuszko National Park, southeastern Australia. Specifically, plant size (vegetative cover of plants in quadrats), floral display (number of flowers per inflorescence, number of inflorescences per quadrat) and sex ratios (proportion of females in quadrats with flowers) were examined in ten populations of Astelia psychrocharis (Asteliaceae) and nine populations of Astelia alpina var. novae-hollandiae (Asteliaceae). Sexual dimorphism did occur, with males having more flowers per inflorescence (106% more flowers for A.alpina males and 12% more for A.psychrocharis males compared to females) and more inflorescences per quadrat than females (78% more inflorescences for A.alpina males and 46% more inflorescences for A.psychrocharis males compared to females). Plant size did not differ between male and female quadrats of either species, nor were there male biased sex ratios. However, plant size was related to flowering status in A.psychrocharis with the 65 quadrats that did not flower having lower vegetative cover than the 175 flowering quadrats indicating that there may be a minimum size/ cover required prior to flowering in this species. For A.alpina, all but two of the 185 quadrats randomly sampled flowered. There was no effect of altitude on plant size and very little effect of altitude on floral display for either species, apart from a slight increase in the number of inflorescences per quadrat with increasing altitude for A.psychrocharis, and slight decrease in number of flowers per inflorescence with increasing altitude for A.alpina females.     

Clonal genetic structure and diversity in populations of an aquatic plant with combined vs. separate sexes

Clonality is often implicated in models of the evolution of dioecy, but few studies have explicitly compared clonal structure between plant sexual systems, or between the sexes in dioecious populations. Here, we exploit the occurrence of monoecy and dioecy in clonal Sagittaria latifola (Alismataceae) to evaluate two main hypotheses: (i) clone sizes are smaller in monoecious than dioecious populations, because of constraints imposed on clone size by costs associated with geitonogamy; (ii) in dioecious populations, male clones are larger and flower more often than female clones because of sex‐differential reproductive costs. Differences in clone size and flowering could result in discordance between ramet‐ and genet‐based sex ratios. We used spatially explicit sampling to address these hypotheses in 10 monoecious and 11 dioecious populations of S. latifolia at the northern range limit in Eastern North America. In contrast to our predictions, monoecious clones were significantly larger than dioecious clones, probably due to their higher rates of vegetative growth and corm production, and in dioecious populations, there was no difference in clone size between females and males; ramet‐ and genet‐based sex ratios were therefore highly correlated. Genotypic diversity declined with latitude for both sexual systems, but monoecious populations exhibited lower genotypic richness. Differences in life history between the sexual systems of S. latifolia appear to be the most important determinants of clonal structure and diversity.     

Patterns of sex ratio variation and genetic diversity in the dioecious forest perennial <Emphasis Type="Italic">Mercurialis perennis</Emphasis>

In small populations of plant species with separate sexes, it can be expected that besides the local environment also stochastic events influence population sex ratios. Biased sex ratios may in turn negatively affect genetic diversity due to increased genetic drift and, in clonal plants, due to reduced sexual reproductive output. Empirical evidence for these processes is scarce, however. We investigated the pattern of sex ratio variation and the distribution of genetic variation of the dioecious clonal forest herb Mercurialis perennis using AFLP markers. Analysis of molecular variance indicated a pronounced genetic structure. Overall within-population genetic diversity was moderate and local sex ratios were slightly male biased. The proportion of male to female plants in large populations slightly increased with increasing light penetration to the herb layer. Small populations, on the contrary, displayed high variability in sex ratios, unrelated to the local light environment. Genotypic diversity decreased with more male-biased sex ratios. We conclude that stochastic events related to small population size and the local forest environment, related to canopy closure, affect the proportion of female plants and indirectly influence local genotypic diversity, likely through the degree of sexual reproduction. This is one of the first studies to report a clear association between gender proportions and genetic diversity of a dioecious plant species in a fairly large survey.     

Population sex ratios, sex-specific clonal traits and tradeoffs among these traits in the liverwort Marchantia inflexa

Unisexual, female- or male-biased populations are common among some clonal plants. Within and among populations, the relative frequencies of males and females can be influenced by sex-specific demographic patterns that, in turn, can be a consequence of life-history characteristics. The objectives of this study were to describe (1)  population sex-ratio patterns among habitat patches along a river, and (2)  sex-specific patterns of and correlations among life-history clonal traits including: growth rates, number of meristematic tips and asexual reproduction in the dioecious liverwort, Marchantia inflexa . In Trinidad, a section of a stream was surveyed for the occurrence of female and male sex expressing thalli among habitat patches, and habitat characteristics (canopy openness and patch size) were recorded. The numbers of female and male inflorescences were obtained also. Of the 209 patches observed in the field, 83% did not contain sex expressing thalli, 9% contained thalli of both sexes, and the rest contained only female- (4%) or male-expressing (4%) thalli. Sex expression was less common among small patches, and there was a tendency for sex expression to be less likely among patches with the lowest canopy openness. The proportion of male inflorescences among the bisexual patches ranged from 0.22 to 0.80. In a greenhouse, we planted gemmae (asexual propagules) from field-collected isolates: ten female and eight male. On six occasions, we harvested replicates of each isolate to estimate isolate trait means. Females grew faster, produced more meristematic tips and had lower levels of asexual reproduction than males. Number of meristematic tips was negatively correlated with asexual reproduction and positively correlated with growth rate. These sex-specific patterns may lead to the high frequency of single sex patches as well as biased adult population sex ratios that are observed in the field.     

Sex ratios and population persistence in the rare shrub Lindera subcoriacea Wofford

Biased sex ratios can have conservation consequences for dioecious plant species with small population sizes because of an increased risk of single sex populations. Biased sex ratios have been observed in two of the three species of Lindera (Lauraceae) in the eastern United States, but have not been documented for Lindera subcoriacea, a rare shrub of the southeastern USA. We inventoried 78 of 118 populations in North Carolina over a 3 year period, documenting the location, community type, and sex, of 299 individuals. In addition, we measured the stem height and diameter for 245 individuals. We examined population persistence relative to historical population size estimates. Average population size was 7.9 individuals and 72 % of visited populations were extant. There was a significant positive correlation between historical estimates of population size and persistence. Lindera subcoriacea consistently had male-biased (58 %) sex ratios across all population sizes and vegetation communities. Males and females had similar stem heights (mean 200.4 vs. 187.8 cm, respectively) and diameters (1.3 vs. 1.2 cm, respectively) across years and were not spatially segregated within populations. It is unclear at what stage biased sex ratios arise in L. subcoriacea, but results suggest that the causes operate across vegetation communities and population sizes. The weak bias exhibited in L. subcoriacea sex ratios has limited implications for the species’ conservation except where spatially isolated populations are unisexual. Given the vulnerability of small L. subcoriacea populations to extirpation, they should be high priority targets for management.     

Maintenance of the sexes and persistence of a clonal organism in spatially complex metapopulations

Clonal organisms persist at a range of population sex ratios, from equal numbers of males and females to single-sex systems. When intersexual competition is strong enough to drive one sex locally extinct, the maintenance of the sexes is facilitated by the semi-independent dynamics of populations within a metapopulation. These semi-independent dynamics are influenced by dispersal and recolonization rates, which are affected by the spatial arrangement of populations. To establish the quantitative relationship between spatially complex metapopulations and the maintenance of the sexes, we used a mathematical model of the liverwort Marchantia inflexa. This clonal organism is found in discrete patches on rocks and along the banks of streams, which form single-sex and two-sex metapopulations. In this system, asexual propagules mainly disperse short distances. Long-distance between-patch dispersal and recolonization mainly occurs via sexual propagules, which require both sexes to be present. Dispersal of these two types of propagules could interact with the spatial arrangement of populations to affect the maintenance of the sexes. With our mathematical model, we found that at intermediate distances between populations, metapopulations maintained both sexes, and the spatial arrangement of populations changed the threshold at which one sex was lost. On the other hand, when populations were close to one another, one sex was lost and the single-sex metapopulation persisted through dispersal of asexual propagules. When populations were far apart, one sex was lost, and the metapopulation either went extinct due to lack of recolonization by asexual propagules or persisted because clumped populations facilitated recolonization. These idealized spatial arrangements help clarify the effects of the spatial arrangement on the maintenance of the sexes and the persistence of metapopulations of clonal organisms, which can help explain geographic parthenogenesis and the distribution of asexual populations, the persistence of asexual species, and inform the conservation of clonal organisms.     

Does Landscape Fragmentation Influence Sex Ratio of Dioecious Plants? A Case Study of Pistacia chinensis in the Thousand-Island Lake Region of China

The Thousand-Island Lake region in Zhejiang Province, China is a highly fragmented landscape with a clear point-in-time of fragmentation as a result of flooding to form the reservoir. Islands in the artificial lake were surveyed to examine how population sex ratio of a dioecious plant specie Pistacia chinensis B. was affected by landscape fragmentation. A natural population on the mainland near the lake was also surveyed for comparison. Population size, sex ratio and diameter at breast height (DBH) of individuals were measured over 2 years. More than 1,500 individuals, distributed in 31 populations, were studied. Soil nitrogen in the different populations was measured to identify the relationship between sex ratio and micro-environmental conditions. In accordance with the results of many other reports on biased sex ratio in relation to environmental gradient, we found that poor soil nitrogen areas fostered male-biased populations. In addition, the degree of sex ratio bias increased with decreasing population size and population connectivity. The biased sex ratios were only found in younger individuals (less than 50 years old) in small populations, while a stable 1∶1 sex ratio was found in the large population on the mainland. We concluded that the effects of landscape fragmentation on the dioecious population sex ratio were mainly achieved in relation to changing soil nitrogen conditions in patches and pollen limitation within and among populations. Large populations could maintain a more suitable environment in terms of nutrient conditions and pollen flow, subsequently maintaining a stable sex ratio in dioecious plant populations. Both micro-environmental factors and spatial structure should be considered in fragmented landscape for the conservation of dioecious plant species.     

Mechanisms governing sex-ratio variation in dioecious Rumex nivalis

Sex ratios of flowering individuals in dioecious plant populations are often close to unity, or are male biased owing to gender-specific differences in flowering or mortality. Female-biased sex ratios, although infrequent, are often reported in species with heteromorphic sex chromosomes. Two main hypotheses have been proposed to account for female bias: (1) selective fertilization resulting from differential pollen-tube growth of female- versus male-determining microgametophytes (certation); (2) differences in the performance and viability of the sexes after parental investment. Here we investigate these hypotheses in Rumex nivalis (Polygonaceae), a European alpine herb with female-biased sex ratios in which females possess XX, and males XY1Y2, sex chromosomes. Using field surveys and a glasshouse experiment we investigated the relation between sex ratios and life-history stage in 18 populations from contrasting elevations and snowbed microsites and used a male-specific SCAR-marker to determine the sex of nonflowering individuals. Female bias among flowering individuals was one of the highest reported for populations of a dioecious species (mean female frequency = 0.87), but males increased in frequency at higher elevations and in the center of snowbeds. Female bias was also evident in nonflowering individuals (mean 0.78) and in seeds from open-pollinated flowers (mean 0.59). The female bias in seeds was weakly associated with the frequency of male flowering individuals in populations in the direction predicted when certation occurs. Under glasshouse conditions, females outperformed males at several life-history stages, although male seeds were heavier than female seeds. Poor performance of Y1Y2 gametophytes and male sporophytes in R. nivalis may be a consequence of the accumulation of deleterious mutations on Y-sex chromosomes.     

Genetic variation within and among patches of the clonal species, Vaccinium stamineum L

Once thought to be dominated by a few genets, clonal plant populations can contain high levels of genetic diversity. Sexual reproduction and vegetative growth strategy affect the amount and distribution of genetic diversity within clonal plant populations. We determined the scale of genetic diversity in a population of Vaccinium stamineum, a clonal shrub that forms discrete patches. Using the random amplified polymorphic DNA (RAPD) technique, we surveyed the genetic diversity of V. stamineum within and among patches from a 1-ha site. We found 67 unique RAPD profiles among the 99 sampled individuals from 22 patches. In two patches, all the sampled individuals had the same RAPD profile. In seven patches, every individual sampled had a different RAPD profile. The remaining patches showed mixed RAPD profiles which suggested both clonal and sexual reproduction. Each unique RAPD profile was restricted to one patch (with one exception), which suggests that clonal growth occurs at the patch scale. High levels of genetic variation within some patches may be explained by somatic mutation; however, seedling recruitment is a more likely explanation.     

Sex ratio, reproductive investment and flowering phenology in dioecious Rhamnus alaternus (Rhamnaceae)

Guitián, J. 1995. Sex ratio, reproductive investment and flowering phenology in dioecious Rhamnus alaternus (Rhamnaceae). - Nord. J. Bot. 15: 139–143. Copenhagen. ISSN 0107–055X.
During 1992 and 1993 I investigated the reproductive biology of the dioecious Mediterranean shrub Rhamnus alaternus in a population in the northwest Iberian Peninsula. Reproductive investment was estimated as the mean total dry weight of reproductive organs per branch tip. I also estimated number of flowers produced per plant and population sex ratio, and investigated the spatial distribution of the sexes and flowering phenology. The sex ratio was 1:1, and the spatial distribution of the sexes was random. Male plants produced 2.6 times more flowers than female plants, but the overall reproductive investment by females was 5- to 9-fold higher. In both years of study, male plants commenced flowering first. The male and female flowering peaks coincided closely (in late March) in both years. The results of this study suggest that male and female R. alaternus differ most notably in the amount of resources allocated to reproduction.     

Dioecious plants are more precocious than cosexual plants: A comparative study of relative sizes at the onset of sexual reproduction in woody species

The reproductive capacities of dioecious plant species may be limited by severe pollen limitation and narrow seed shadows for the two reasons. First, they are unable to self‐pollinate, and seed production occurs only with pollinator movement from males to females. Second, only 50% of the individuals in populations contribute to seed production. Despite these handicaps, dioecious plants maintain their populations in plant communities with cooccurring cosexual plants, and no substantial difference in population growth rates has been found between dioecious and cosexual plants. Hence, dioecious plants are thought to mitigate these disadvantages by adopting ecological traits, such as insect pollination, animal‐dispersed fleshy fruits, and precocious flowering. We studied the relationship between flowering and plant size in 30 woody species with different sex expressions, leaf habits, fruit types, and maximum plant sizes. The study site was located in an evergreen broad‐leaved forest on the island of Honshu, Japan. A phylogenetic linear regression model showed that dioecious species tended to mature at smaller sizes than did cosexual taxa. At the population level, given equal plant densities and reproductive efforts, the precocity of dioecious plants could serve as one of the factors that mitigate the limitations of pollen and seed‐shadow handicaps by increasing the density of reproductive individuals in the population. At the individual level, smaller size of onset of flowering may play a role in enhancing reproductive success over a lifetime by increasing reproductive opportunities. We discussed the possible effect of the relationship between precocity and some ecological traits of dioecious plants, such as small flowers pollinated by unspecialized insects, fleshy fruit dispersed by animals, and their preferential occurrence in the tropics and in island habitats. The universality of precocity among dioecious plants should be investigated in diverse plant communities. Such studies will increase our understanding of the evolution of plant breeding systems.     

Effect of sex ratio,habitat factors and neighborhood competition on stem growth in the dioecious tree Fraxinus mandshurica

Growth rates of male and female trees are often different in a dioecious species. In this study, we analyzed sex ratios and the effect of gender, neighborhood competition and habitat factors on the stem growth of dioecious Fraxinus mandshurica trees in a secondary conifer and broad-leaved mixed forest in the Changbai Mountains of northeastern China. The sex ratio in the 5.2-ha study area does not deviate significantly from the expected 1:1 ratio, except for trees in the large diameter classes. For dbh >40 cm, the sex ratio is male-biased. This result suggests that males have a faster rate of stem growth than females, which is usually explained by the higher cost of reproduction in the fruit-bearing females. An analysis of the dbh distributions of two successive measurements showed that the rate of stem growth of the (27) females drops off with increasing dbh and remains below that of the (35) males. A causal model was used to analyze factors affecting the rate of stem growth, showing that these rates are affected significantly and positively by soil moisture and tree size in both genders and that within-gender competition is mainly for nitrogen. Our study suggests that neighborhood competition does not affect stem growth significantly, which is a rather surprising result.     

The influence of demography and local mating environment on sex ratios in a wind‐pollinated dioecious plant

Negative frequency‐dependent selection should result in equal sex ratios in large populations of dioecious flowering plants, but deviations from equality are commonly reported. A variety of ecological and genetic factors can explain biased sex ratios, although the mechanisms involved are not well understood. Most dioecious species are long‐lived and/or clonal complicating efforts to identify stages during the life cycle when biases develop. We investigated the demographic correlates of sex‐ratio variation in two chromosome races of Rumex hastatulus, an annual, wind‐pollinated colonizer of open habitats from the southern USA. We examined sex ratios in 46 populations and evaluated the hypothesis that the proximity of males in the local mating environment, through its influence on gametophytic selection, is the primary cause of female‐biased sex ratios. Female‐biased sex ratios characterized most populations of R.  hastatulus (mean sex ratio = 0.62), with significant female bias in 89% of populations. Large, high‐density populations had the highest proportion of females, whereas smaller, low‐density populations had sex ratios closer to equality. Progeny sex ratios were more female biased when males were in closer proximity to females, a result consistent with the gametophytic selection hypothesis. Our results suggest that interactions between demographic and genetic factors are probably the main cause of female‐biased sex ratios in R. hastatulus. The annual life cycle of this species may limit the scope for selection against males and may account for the weaker degree of bias in comparison with perennial Rumex species.     

Effects of kin-structured seed dispersal on the genetic structure of the clonal dioecious shrub ilex leucoclada

Nonrandom patterns of gene dispersal have been identified as possible causes of genetic structuring within populations. Attempts to model these patterns have generally focused solely on the effects of isolation by distance, but the processes involved are more complex than such modeling suggests. Here, we extend considerations of gene dispersal processes beyond simple isolation by distance effects by directly evaluating the effects of kin-structured gene dispersal mediated by the group dispersal of related seeds within fruits (i.e., kin-structured seed dispersal) by birds on genetic structure in Ilex leucoclada, a clonal dioecious shrub. To examine the genetic structure patterns, we established two 30x30 m plots (one with immature soils in old-growth forest and one in secondary forest, designated IM and SC, respectively) with different I. leucoclada stem densities. In these two plots 145 and 510 stems were found, representing 78 and 85 genets, respectively, identified by analyzing their genotypes at eight microsatellite loci. The clonal structure was stronger in the SC plot than in the IM plot. Correlograms of coancestry for genets in both plots exhibited significant, positive, high values in the shortest distance class, indicating the presence of strong genetic structure. However, Sp statistics revealed that the pattern of the genetic structure differed between the plots. In addition, to estimate the family structure within fruits, we sampled forty fruits, in total, from 15 randomly selected plants in the area around the IM and SC plots, and found that 80% of the fruits were multiseeded and 42-100% of the multiseeded fruits contained at least one pair of full sibs. Simulations based on these estimates demonstrated that the group dispersal of related seeds produced through correlated mating both within and across fruits, but not unstructured half-sib dispersal, could generate the observed magnitude and trends of genetic structure found in the IM plot. Furthermore, in addition to kin-structured seed dispersal, isolation by distance processes is also likely to promote genetic substructuring in the SC plot. After discussing possible ecological factors that may have contributed to the observed genetic structure, we contrast our results with those predicted by general isolation by distance models, and propose that kin-structured seed dispersal should promote some evolutionary phenomena, and thus should be incorporated, where appropriate, in models of gene dispersal in natural plant populations.