Pollination ecology in sympatric winter flowering Helleborus (Ranunculaceae)

We studied biotic and abiotic factors that influence pollination in two sympatric winter flowering species. Helleborus foetidus and Helleborus bocconei flower simultaneously in winter. Although climatic conditions are not favorable for biotic pollination both species rely mainly on large bees of the genus Bombus. At the beginning of flowering, harsh climatic conditions are restrictive for insect visits. As flowering continues and temperatures rise, pollinator activity increases. The two plant species share pollinators that visit them indiscriminately. The flowers of the two species differ in form and insects visit H. foetidus by inserting their heads and H. bocconei ventrally: pollen load on insects is highly specialized. With the arrival of spring, many other species start to bloom and in spite of the large number of flowers still on the plants insects abandon Helleborus species. At the end of spring increasing biotic interactions take away pollinators from the Helleborus species.     

Die Tricks der Stinkenden Nieswurz

The tricks of stinking hellebore The stinking hellebore (Helleborus foetidus) uses ingenious tricks in order to attract pollinators and to achieve the dispersal of its seeds. Yeasts metabolise sugar available in nectar und thereby warm it up to attract bees and bumble bees in cold, early spring. This remarkable heating system allows the flowers to already attract pollinators at a time, when competition by other flower species is still low and visits by pollinators are often limited by low temperatures. Early flowering in turn allows seed set in late spring and early summer. The seeds have a fleshy structure attached, an elaiosome that contains lipids, sugars and vitamins. This makes them highly attractive to ants during a season when high-quality foods are still in short supply and leads to dispersal of the seeds (myrmecochory). The elaiosome also contains a fatty acid that acts as an olfactory attractant. But what attracts ants, is also attractive for other seed predators such as mice. Long-term survival of seeds (and thus germination) therefore greatly depends on the detachment of the elaiosome from the seed by the ants. There is much more than meets the eye with respect of stinking hellebore pollination and seed dispersal. It is an example of the highly complex interdependencies between plants, their blossoms, different animal species as well as yeasts and bacteria.     

Floral integration,phenotypic covariance structure and pollinator variation in bumblebee‐pollinated Helleborus foetidus

By analysing patterns of phenotypic integration and multivariate covariance structure of five metric floral traits in nine Iberian populations of bumblebee‐pollinated Helleborus foetidus (Ranunculaceae), this paper attempts to test the general hypothesis that pollinators enhance floral integration and selectively modify phenotypic correlations between functionally linked floral traits. The five floral traits examined exhibited significant phenotypic integration at all populations, and both the magnitude and the pattern of integration differed widely among populations. Variation in extent and pattern of integration was neither distance‐dependent nor significantly related to between‐population variation in taxonomical composition and morphological diversity of the pollinator assemblage. Patterns of floral integration were closer to expectations derived from consideration of developmental affinities between floral whorls than to expectations based on a pollinator‐mediated adaptive hypothesis. Taken together, results of this study suggest that between‐population differences in magnitude and pattern of floral integration in H. foetidus are probably best explained as a consequence of random genetic sampling in the characteristically small and ephemeral populations of this species, rather than reflecting the selective action of current pollinators.     

Indirect effects of alternative food resources in an ant–plant interaction

The seeds of many plant species present a food body that is consumed by animal dispersers. In theory, if the animals are polyphagous, the availability of alternative food resource other than the diaspore itself may influence its dispersal and survival. We used the myrmecochore Helleborus foetidus L. (Ranunculaceae), the seeds of which are attached to a lipid-rich elaiosome that is attractive to ants, as a model system to investigate (1) whether alternative foods that are present along with the plant affect ant foraging behavior and diaspore removal and (2) whether food availability in an ant nest affects seed predation and germination. In a field experiment, artificial diaspore depots were offered together with either sugar, insect corpses, seed, or no food (control). Contrary to the prediction that ants would rather concentrate their foraging effort on the highly rewarding alternative foods only, many workers, attracted by the sugar, switched to the hellebore diaspores, which significantly enhanced removal rate. Results obtained in the laboratory further indicated that the larvae of Aphaenogaster iberica (a major seed disperser) predated more on the H. foetidus embryos when no alternative food was available. This, in turn, slightly reduced seed germination. Overall, these results shed light, for the first time, on the potential indirect effects of alternative resources on the fate of diaspores adapted for ant dispersal.     

Variation in structural gender in the hermaphrodite Helleborus foetidus (Ranunculaceae): within- and among-population patterns

In hermaphrodite plants, variations in structural gender (defined as the ratio between male and female gametes) may occur at different levels (among flowers, plants, and populations). In this study, we investigated variation in four traits influencing structural gender (number of carpels, ovules per carpel, stamens, and pollen grains per stamen) within and among six distant populations of the hermaphrodite perennial herb Helleborus foetidus (Ranunculaceae) in the Iberian Peninsula. Our results show that the four traits investigated varied significantly at all levels considered. Traits influencing the female sexual component (number of carpels and ovules per carpel) showed greater variation at the lowermost levels (within flower and plant) than traits influencing the male component, which in turn varied more markedly among populations. Number of carpels per flower and number of pollen grains per anther were the most important traits affecting between-plant variation in structural gender. There was no evidence of significant plant-level trade-offs or correlations between the various male and female traits, which covaried differently across populations. The observed between-population variation in structural gender of Helleborus foetidus can be explained as a consequence of differences in self-pollination levels related to a flowers mating environment.     

Seed- vs. microsite-limited recruitment in a myrmecochorous herb

Dispersal in myrmecochorous herbs is frequently short-distance and this might limit patch occupancy. Here we ask whether recruitment and patch occupancy in the myrmecochere Helleborus foetidus are seed- or microsite-limited and what role dispersal plays in recruitment limitation. In pine forests from two widely separated mountain systems in the Iberian Peninsula, we performed two seed addition experiments to explore the nature of recruitment limitation in this perennial herb. In an introduction experiment, we added seeds in patches far from reproductive individuals, which rarely receive any seed. In an augmentation experiment, we added seeds in patches beneath reproductive plants. Each experiment considered three main factors: seed addition (added or not added), rodent exclusion (excluded or not excluded), and microhabitat (scrub-covered or not scrub-covered). The augmentation experiment tests whether recruitment is limited by seed availability (exploring also the possible role of rodent granivory in such limitation) while the introduction experiment tests whether the empty patches occur either because they are unsuitable for the species (microsite limitation) or due to a low probability of seed arrival (dispersal limitation). Adding seeds almost always increased seedling emergence and recruitment, demonstrating that recruitment in this species is seed-limited and refuting the possibility of microsite limitation. The comparison of the introduction and augmentation experiments indicated that Helleborus recruitment dynamics and patch occupancy are constrained by additive effects of fecundity and dispersal limitation.     

Epigenetic variation predicts regional and local intraspecific functional diversity in a perennial herb

The ecological significance of epigenetic variation has been generally inferred from studies on model plants under artificial conditions, but the importance of epigenetic differences between individuals as a source of intraspecific diversity in natural plant populations remains essentially unknown. This study investigates the relationship between epigenetic variation and functional plant diversity by conducting epigenetic (methylation‐sensitive amplified fragment length polymorphisms, MSAP) and genetic (amplified fragment length polymorphisms, AFLP) marker–trait association analyses for 20 whole‐plant, leaf and regenerative functional traits in a large sample of wild‐growing plants of the perennial herb Helleborus foetidus from ten sampling sites in south‐eastern Spain. Plants differed widely in functional characteristics, and exhibited greater epigenetic than genetic diversity, as shown by per cent polymorphism of MSAP fragments (92%) or markers (69%) greatly exceeding that for AFLP ones (41%). After controlling for genetic structuring and possible cryptic relatedness, every functional trait considered exhibited a significant association with at least one AFLP or MSAP marker. A total of 27 MSAP (13.0% of total) and 12 AFLP (4.4%) markers were involved in significant associations, which explained on average 8.2% and 8.0% of trait variance, respectively. Individual MSAP markers were more likely to be associated with functional traits than AFLP markers. Between‐site differences in multivariate functional diversity were directly related to variation in multilocus epigenetic diversity after multilocus genetic diversity was statistically accounted for. Results suggest that epigenetic variation can be an important source of intraspecific functional diversity in H. foetidus, possibly endowing this species with the capacity to exploit a broad range of ecological conditions despite its modest genetic diversity.     

Pre- and postdispersal seed predation by rodents: balance of food and safety

Seed presentation and availability for seed predators changeduring every plant reproductive cycle. We know very little abouthow those changes impinge on both the ability of seed predatorsto impact plant populations and the foraging costs associatedwith seed consumption. Therefore, we conducted several fieldexperiments to evaluate whether wood mice Apodemus sylvaticusbalance food and safety while foraging on Helleborus foetidusseeds during both the pre- and early postdispersal phases ofthe plant reproductive cycle. Both food and safety were keydeterminants of mouse foraging on H. foetidus seeds, thoughtheir roles were not consistent along the plant reproductivecycle. Thus, augmenting ambient food reduced fruit removal bymice during the predispersal phase. During the postdispersalphase, seeds in sheltered microsites experienced higher removalrates than those located in nonsheltered microsites; however,no effect of food augmentation was detected. This apparent reversedrole of food and safety on decision making by mice seemed closelylinked to both the dramatic changes in accessibility and presentationof H. foetidus seeds and the coupled changing foraging costsfaced by mice at different phases of the plant reproductivecycle. For instance, because the cost of foraging for predispersalseeds was higher than for postdispersal seeds, the effect offood augmentation on foraging by wood mice was greater duringthe predispersal phase. Thus, our study illustrates the needof considering differences between pre- and postdispersal seedpredation in the study of granivore rodents and their impacton plant populations.     

Geographic variation in the adaptive landscape for seed size at dispersal in the myrmecochorous <Emphasis Type="Italic">Helleborus foetidus</Emphasis>

Literature on seed dispersal mutualisms suggests that plant populations should hardly adapt to their current dispersers. We address the predictions that selection pressures exerted by ants on dispersal-related diaspore traits of the ant-dispersed Helleborus foetidus are highly variable in space, and that geographic (inter-population) variation in these traits is unrelated to selection by current dispersers. To test these predictions we use the concept of the quantitative adaptive landscape for seed size at dispersal. Such landscape depicts the relationship between the population’s mean trait value (mean seed size in the present study) and the population’s mean fitness (mean dispersal probability in the present study). Adaptive landscapes make it possible to assess whether the mean population’s phenotype agrees with one favored by selection. We first analyse, in 12 populations of H. foetidus from southern Spain, the extent of divergence among populations in seed and elaiosome size, and the abundance, composition, and behavior of the ant communities. Seeds from a fixed set of five of these populations were offered to ants in all the study sites to fit the adaptive landscape for seed size. In addition, seeds from the local population were also offered in each site. Our results show that seed size has undergone a larger divergence among populations than elaiosome size. Despite geographic variation in ant assemblages, the adaptive landscapes for seed size at dispersal were remarkably similar among sites: ants create disruptive selection on seed size in 10 out of 12 study sites. As predicted, the basic features of these adaptive landscapes (curvature and location of the minimum) varied geographically in accordance with variation in the size of seed dispersers. Also as predicted, in most populations, the observed mean seed size does not agree with that expected from the adaptive landscapes at dispersal. However, the relevance of dispersers for seed size evolution should not be neglected since the agreement between observed and optimum seed size was stronger where dispersers were more abundant. Thus, against the general view, our results evidence that, in H. foetidus, the observed geographic variation in dispersal-related plant traits is partly linked to selection exerted by current dispersers. Geographic variation in ant assemblages determines both the existence of a selection mosaic and the degree of adjustment of populations to the patterns of selection in the mosaic.     

铁筷子提取物对肿瘤细胞增殖及COX-2 mRNA表达的抑制作用

研究不同铁筷子提取物对肿瘤细胞增殖及 COX-2 mRNA 表达的抑制作用。以铁筷子醇总提取物(TKZ1)、正丁醇萃取部位(TKZ2)、乙酸乙酯萃取部位(TKZ3)分别作用于 DU145、PC3、HeLa、HT-29、HepG2等肿瘤细胞,应用噻唑蓝实验(MTT 法)计算其对细胞增殖的抑制作用,应用荧光定量 PCR 技术检测TKZ1、TKZ2、TKZ3处理后的各肿瘤细胞中 COX-2 mRNA 的表达情况。结果表明：TKZ1、TKZ2、TKZ3均能显著抑制多种肿瘤细胞的增殖,与阴性对照组比较,其可以在 mRNA 水平上抑制 COX-2的表达,且呈明显的量效关系。说明铁筷子提取物对体外肿瘤细胞的增殖具有显著的抑制作用,其抗瘤机制可能与抑制肿瘤细胞中 COX-2 mRNA 的表达有关。     

Frugivorous birds mediate sex‐biased facilitation in a dioecious nurse plant

Abstract. Facilitation by dispersal occurs if the nurse plant acts as a focus which is actively selected by seed dispersers and enhances the fitness of the facilitated plant. Sex‐biased facilitation may be produced if seed dispersers tend to concentrate the seeds under female, fruit‐bearing plants of dioecious species more often than under conspecific males. Juniperus sabina is a dioecious shrub with a prostrate growth form from Mediterranean high mountains that modifies many microhabitat characteristics related to seedling establishment and survival. Soil water availability, maximum soil temperature in summer, organic matter and total nitrogen content, were different on open ground as compared with beneath J. sabina shrubs, irrespective of its sex. Other studied characteristics such as soil bulk density and soil compaction after rain did not differ between the microhabitats considered. Some species, such as Juniperus communis, Pinus nigra, Helleborus foetidus and Euphorbia nicaeensis, are spatially associated to J. sabina shrubs, strongly suggesting a facilitative role. The anemochorous P. nigra and myrme‐chorous H. foetidus and E. nicaeensis did not associate preferentially to any sex of J. sabina. Only J. communis, an endozoochorous species sharing the same bird dispersers as J. sabina, presented a female‐biased spatial association with the nurse plant. Seed dispersal mediated by birds attracted by the fruit‐rewarding females of J. sabina explains the sex‐biased spatial pattern of Juniperus communis.     

MSAP markers and global cytosine methylation in plants: a literature survey and comparative analysis for a wild‐growing species

Methylation of DNA cytosines affects whether transposons are silenced and genes are expressed, and is a major epigenetic mechanism whereby plants respond to environmental change. Analyses of methylation‐sensitive amplification polymorphism (MS‐AFLP or MSAP) have been often used to assess methyl‐cytosine changes in response to stress treatments and, more recently, in ecological studies of wild plant populations. MSAP technique does not require a sequenced reference genome and provides many anonymous loci randomly distributed over the genome for which the methylation status can be ascertained. Scoring of MSAP data, however, is not straightforward, and efforts are still required to standardize this step to make use of the potential to distinguish between methylation at different nucleotide contexts. Furthermore, it is not known how accurately MSAP infers genome‐wide cytosine methylation levels in plants. Here, we analyse the relationship between MSAP results and the percentage of global cytosine methylation in genomic DNA obtained by HPLC analysis. A screening of literature revealed that methylation of cytosines at cleavage sites assayed by MSAP was greater than genome‐wide estimates obtained by HPLC, and percentages of methylation at different nucleotide contexts varied within and across species. Concurrent HPLC and MSAP analyses of DNA from 200 individuals of the perennial herb Helleborus foetidus confirmed that methyl‐cytosine was more frequent in CCGG contexts than in the genome as a whole. In this species, global methylation was unrelated to methylation at the inner CG site. We suggest that global HPLC and context‐specific MSAP methylation estimates provide complementary information whose combination can improve our current understanding of methylation‐based epigenetic processes in nonmodel plants.     

Phenotypic gender variation within inflorescences of the protogynous species Helleborus foetidus L. (Ranunculaceae)

In the protogynous species H. foetidus, I investigated if investment in sexual structures and seed set shows any trend with position in the flower-opening sequence. In four-flower inflorescences, stamen number decreased with flower position i.e. was highest for the earliest flower and lowest for the latest flower. Ovule number was significantly higher in the latest flower. Stamen and ovule number did not covary, indicating that there is no structural gender trade-off in this species. Seed set (i.e. percentage of ovules producing seed) did not differ between control plants and pollen-supplemented plants and the effect of pollen supplementation did not vary among positions. Seed set showed marginally significant differences between control plants and simulated-herbivory plants, but the effects of this treatment varied significantly among positions. Significant among-position variation in seed set was observed in the control plants but not the simulated-herbivory plants. Seed set of latest-opening flowers of simulated-herbivory plants was significantly higher than that of latest-opening flowers of control plants. H. foetidus support Brunet and Charlesworth's (1995) prediction that in plants with protogynous flowers, later-opening flowers should specialize as female flowers, at least under conditions of high resource availability.     

Some permeability properties of angiosperm pollen grains,pollen tubes and generative cells

Summary The permeability of pollen grains, pollen tubes and generative cells of Helleborus foetidus and Galanthus nivalis has been investigated using four probes spanning a wide range of molecular weights: 4,6-diamidino-2-phenyl indole (DAPI; mol.wt. 350). Evans blue (mol.wt. 960), FITC-dextran (average mol.wt. 19400) and FITC-albumin (average mol.wt. 67000). DAPI penetrated into the vegetative cells of desiccated and hydrated pollen, and also entered growing pollen tubes. In contrast, the generative cells of hydrated pollen and of pollen tubes were highly resistant to penetration, as they were when isolated in osmotically balancing medium. Evans blue failed to enter intact generative cells under any of the conditions tested. The dye ultimately entered the vegetative cells of some pollen grains, but these were non-germinable. Growing pollen tubes invariably resisted penetration. Neither of the high molecular weight conjugates entered germinable pollen grains or intact pollen tubes. The results suggest that it is highly unlikely that DNA fragments of high molecular weight can enter viable pollen, pollen tubes or generative cells under any normal conditions.     

The Comportment of the Vegetative Nucleus and Generative Cell in the Pollen and Pollen Tubes of Helleborus foetidus L

It has been reported that in species of Plumbaginaceae, Chenopodiaceae,Cruciferae and Amaryllidaceae a ‘male germ unit’is formed in which the two male gametes remain inter-connected,with one of the pair linked intimately to the vegetative nucleus.In two species the unit has been shown to remain intact in thepollen tube, and some accounts imply that it is polarized inits movement, the vegetative nucleus leading in the tube. Evidence given in this paper indicates that such a unit is unlikelyto be present in Helleborus foetidus L. (Ranunculaceae). Applicationof an optical sectioning technique has shown that at no timeis there a persistent linkage between the generative cell andthe vegetative nucleus in unhydrated, hydrated and germinatingpollen, nor is one present in the early pollen tube. Furthermore,no inter-connections between the two entities were seen in protoplastsfrom living, hydrated and incipiently germinating grains isolatedmechanically in an osmotically balancing medium. Following germination,the vegetative nucleus leaves the grain in advance of the generativecell in most instances, but in the samples examined the generativecell led in about 30 per cent of the tubes. Assembling a polarisedmale germ unit in these circumstances would require (a) theformation of an inter-connection between the vegetative nucleusand the generative cell or one of the gametes derived from itduring passage through the tube, and (b) where the generativecell initially leads in the tube, an exchange in relative positions.It is considered improbable that these conditions could consistentlybe met. Mature, incipiently germinating pollen of H. foetidus releasesa fibrillar component when extruded into suitable media. Websor clusters of fibrils are commonly seen to be associated withboth the vegetative nucleus and the generative cell. The possibilitythat the fibrils are composed of aggregates of microfilamentsis considered. Helleborus foetidus L., pollen germination, generative cell, vegetative nucleus, male germ unit     

Lipid content and fatty acid composition in hypogeous organs ofHelleborus species (Ranunculaceae)

Root and rhizome of fourHelleborus species:H. viridis, H. odorus, H. niger, H. foetidus were examined throughout a two-year period in order to determine the variation of lipid content and fatty acid composition during the ontogenetic cycle. In the deciduous geophytesH. viridis andH. odorus subsp.laxus the lipid content reaches its maximum during the quiescent phase, whereas in the evergreen geophyteH. niger the lipid content is highest during the main growth period. In the chamaephyteH. foetidus lipid content is always very low and it does not show variation throughout the year. In all species the fatty acids detected in the neutral fraction are myristic, palmitic, stearic, oleic and linoleic acid. Palmitic and linoleic acid are always present in larger quantities. The pattern of the relative content of saturated/unsaturated fatty acids seems to be linked to the life form of these species.     

Land-use and edge effects unbalance seed dispersal and predation interactions under habitat fragmentation

The process of fragmentation can greatly influence plant–animal interactions. To assess the degree to which it affects the balance between two interactions of opposite sign, namely seed dispersal and post-dispersal seed predation, we selected 16 patches of chestnut forest in O Courel and El Bierzo, northwestern Spain. We assessed the effect of fragmentation over two different seed dispersal–predation systems using Helleborus foetidus and Ilex aquifolium as model species. In the first case, field experiments consisted of seed-offering trays with selective exclusion of rodents and ants in a two-way orthogonal design. In the second experiment, we placed experimental branches and trays on the floor to assess seed dispersal and predation. The interactions between several fragment traits and the relative contribution of rodents, ants and birds to seed removal were analyzed by means of generalized linear mixed models. Results show that for H. foetidus, differences in seed dispersal–predation were accounted for by patch shape, which affected mainly the dispersal phase. Major seed dispersal took place in patches with a smaller edge to core ratio and high plant cover (abandoned patches), whilst the latter also showed maximum seed predation. For I. aquifolium, fragmentation effects were significant only for seed predation, which was increased in abandoned patches. This shows that the effects of habitat fragmentation can emerge at different phases depending on specific traits of the interacting animals. It also highlights the importance of traditional land-use practices in species interactions.     

The role of post‐floral persistent perianth in Helleborus viridis subsp. occidentalis (Ranunculaceae)

Species within the genus Helleborus differ in the relative location of their sepals. Previous studies have proved the existence of post‐floral functionality of sepals in H. foetidus, which leads us to consider the possibility of differences in the functionality of the sepals in H. viridis subsp. occidentalis. In this study, we analyzed their influence on the number and weight of seeds through experimental manipulations, which involve a progressive reduction of sepals following fertilization. Our results show that different levels of perianth reduction have no effect on the number and weight of seeds in Helleborus viridis subsp. occidentalis. We propose that differences in the timing of leaf development and changes in the position of floral organs among the different Helleborus species, underlie a distinctive response to the ever‐changing weather conditions of the European winters and springs.     

Within-carpel and among-carpel competition during seed development, and selection on carpel number, in the apocarpous perennial herb Helleborus foetidus L. (Ranunculaceae)

Apocarpous flowers share opportunities for post-fertilization ovule selection among more functional levels than syncarpous flowers, because the occurrence of a variable number of unfused carpels adds a new source of variation to the likelihood of successful female reproduction. The extent to which post-fertilization events might differ among these unfused carpels may promote variations in the reproductive strategies of plants. We report a study of the variation, within and among carpels and flowers, in seed production and mass in the apocarpous Helleborus foetidus (Ranunculaceae), in relation to the number of carpels per flower. Differences within and among carpels in female reproductive success were affected by carpel number and pollination environment. When analysing whole flowers as functional units we also found that the magnitude of the differences related to carpel number and pollination treatment actually depended on the “distribution” of pollen types within flowers. Thus, variable within-flower pollination environments, more likely to occur in apocarpous than in syncarpous flowers, may affect the strategies of resource allocation for fruit development at different stages of the reproductive process. Regarding seed production, we found that producing more flowers with four carpels was under directional; however, when mean diaspore mass was considered as a measure of fitness, directional selection was found on producing flowers with two and three carpels (the modal carpel number found in wild populations). We discuss ecological and developmental reasons which could explain the observed pattern, and conclude that selection on an optimum carpel number may be very variable across the species range, as the discussed reasons may impose constraints on eventual evolutionary response, thus contributing to the maintenance of the intra-individual variability in carpel number.     

Variation in DNA methylation transmissibility,genetic heterogeneity and fecundity‐related traits in natural populations of the perennial herb Helleborus foetidus

Inferences about the role of epigenetics in plant ecology and evolution are mostly based on studies of cultivated or model plants conducted in artificial environments. Insights from natural populations, however, are essential to evaluate the possible consequences of epigenetic processes in biologically realistic scenarios with genetically and phenotypically heterogeneous populations. Here, we explore associations across individuals between DNA methylation transmissibility (proportion of methylation‐sensitive loci whose methylation status persists unchanged after male gametogenesis), genetic characteristics (assessed with AFLP markers), seed size variability (within‐plant seed mass variance), and realized maternal fecundity (number of recently recruited seedlings), in three populations of the perennial herb Helleborus foetidus along a natural ecological gradient in southeastern Spain. Plants (sporophytes) differed in the fidelity with which DNA methylation was transmitted to descendant pollen (gametophytes). This variation in methylation transmissibility was associated with genetic differences. Four AFLP loci were significantly associated with transmissibility and accounted collectively for ~40% of its sample‐wide variance. Within‐plant variance in seed mass was inversely related to individual transmissibility. The number of seedlings recruited by individual plants was significantly associated with transmissibility. The sign of the relationship varied between populations, which points to environment‐specific, divergent phenotypic selection on epigenetic transmissibility. Results support the view that epigenetic transmissibility is itself a phenotypic trait whose evolution may be driven by natural selection, and suggest that in natural populations epigenetic and genetic variation are two intertwined, rather than independent, evolutionary factors.