Reproductive assurance varies with flower size in Collinsia parviflora (Scrophulariaceae)

A central question in plant evolutionary ecology is how mixed mating systems are maintained in the face of selection against self-pollination. Recently, attention has focused on the potential reproductive assurance (RA) benefit of selfing: the ability to produce seeds via autonomous selfing when the potential for outcrossing is reduced or absent. To date, there is little experimental support for this benefit under natural pollination conditions. In addition, the RA hypothesis has not been tested experimentally in a species displaying morphological variation for traits expected to influence the mating system, such as flower size, which affects both attractiveness to pollinators and ability to self autonomously. Here, we document significant among-population variation in flower size in Collinsia parviflora and show that pollinators preferred large flowers over small flowers in experimental arrays. The pollinator community varied among three study sites, and two small-flowered populations had lower pollinator visitation rates than one large-flowered population. We compared seed production between intact flowers (can self) and experimentally emasculated flowers (require a pollinator) on large- and small-flowered plants. As predicted by the RA hypothesis, small-flowered plants show a greater RA benefit of selfing than large-flowered plants; emasculated, small flowers produced very few seeds, relative to intact, small flowers or either emasculated or intact, large flowers. We also show that the RA benefit is pollination-context dependent, differing between small- and large-flowered test sites, likely due to a combination of pollinator discrimination against small flowers and differences between test sites in the pollinator community. This paper is the first experimental evidence showing a trait-dependent RA benefit of selfing under natural pollination conditions.     

Variation in pollen limitation and floral parasitism across a mating system transition in a Pacific coastal dune plant: evolutionary causes or ecological consequences?

Background and Aims Evolutionary transitions from outcrossing to self-fertilization are thought to occur because selfing provides reproductive assurance when pollinators or mates are scarce, but they could also occur via selection to reduce floral vulnerability to herbivores. This study investigated geographic covariation between floral morphology, fruit set, pollen limitation and florivory across the geographic range of Camissoniopsis cheiranthifolia, a Pacific coastal dune endemic that varies strikingly in flower size and mating system.Methods Fruit set was quantified in 75 populations, and in 41 of these floral herbivory by larvae of a specialized moth (Mompha sp.) that consumes anthers in developing buds was also quantified. Experimental pollen supplementation was performed to quantify pollen limitation in three large-flowered, outcrossing and two small-flowered, selfing populations. These parameters were also compared between large- and small-flowered phenotypes within three mixed populations.Key Results Fruit set was much lower in large-flowered populations, and also much lower among large- than small-flowered plants within populations. Pollen supplementation increased per flower seed production in large-flowered but not small-flowered populations, but fruit set was not pollen limited. Hence inadequate pollination cannot account for the low fruit set of large-flowered plants. Floral herbivory was much more frequent in large-flowered populations and correlated negatively with fruit set. However, florivores did not preferentially attack large-flowered plants in three large-flowered populations or in two of three mixed populations.Conclusions Selfing alleviated pollen limitation of seeds per fruit, but florivory better explains the marked variation in fruit set. Although florivory was more frequent in large-flowered populations, large-flowered individuals were not generally more vulnerable within populations. Rather than a causative selective factor, reduced florivory in small-flowered, selfing populations is probably an ecological consequence of mating system differentiation, with potentially significant effects on population demography and biotic interactions.     

No evidence for a role of pollinator discrimination in causing selection on flower size through female reproduction

The preference for certain floral phenotypes by flower visiting animals may fuel the evolution of floral traits because variation in flower visitation rates may lead to fitness variation within a population. Here, I examine the importance of flower size for pollinator visitation rate, seed set, and seed mass in two alpine populations of the insect-pollinated herb Ranunculus acris L. during two seasons. There was no pollen limitation of seed set or mass. Pollinators discriminated strongly against flowers experimentally reduced in size. Despite this, there were no signs of any significant impact of flower size on female reproductive success. The results show that although pollinators discriminate strongly among floral phenotypes, this may not always result in female fitness differences within a population because seed set or mass is not limited by pollen availability alone. Probably abiotic environmental constraints prevent plants with high pollinator visitation from capitalizing on the high pollen deposition.     

Variation in sex ratio, morph-specific reproductive ecology and an experimental test of frequency-dependence in the gynodioecious Kallstroemia grandiflora (Zygophyllaceae)

An enduring puzzle in gynodioecious species is the great variation in female frequency seen among populations. We quantified sex ratio in 44 populations of gynodioecious Kallstroemia grandiflora. Then, we measured pollinator visitation, pollen deposition, autonomous selfing rate and pollen limitation of females. Finally, using experimental populations, we tested whether female fitness responds to the frequency of female plants. We found broad variability in sex ratio among populations (0-44% female). Hermaphrodite flowers received more pollinator visits and pollen grains than females, and bagged hermaphrodite flowers produced fruits. However, we found no evidence of pollen limitation in females. In experimental populations, female plants showed no evidence of frequency-dependent pollinator visitation, fruit set, seed set or total seed mass. These results do not support frequency-dependent variation in fitness as a major mechanism affecting female frequencies in K. grandiflora. Within the context of this study, pollinators are abundant and pollinator movement appears to operate at a large enough scale to overcome the potential reproductive disadvantages of producing solely female flowers.     

Relative impact of mate versus pollinator availability on pollen limitation and outcrossing rates in a mass‐flowering species

Plant mating systems are driven by several pre‐pollination factors, including pollinator availability, mate availability and reproductive traits. We investigated the relative contributions of these factors to pollination and to realized outcrossing rates in the patchily distributed mass‐flowering shrub Rhododendron ferrugineum. We jointly monitored pollen limitation (comparing seed set from intact and pollen‐supplemented flowers), reproductive traits (herkogamy, flower size and autofertility) and mating patterns (progeny array analysis) in 28 natural patches varying in the level of pollinator availability (flower visitation rates) and of mate availability (patch floral display estimated as the total number of inflorescences per patch). Our results showed that patch floral display was the strongest determinant of pollination and of the realized outcrossing rates in this mass‐flowering species. We found an increase in pollen limitation and in outcrossing rates with increasing patch floral display. Reproductive traits were not significantly related to patch floral display, while autofertility was negatively correlated to outcrossing rates. These findings suggest that mate limitation, arising from high flower visitation rates in small plant patches, resulted in low pollen limitation and high selfing rates, while pollinator limitation, arising from low flower visitation rates in large plant patches, resulted in higher pollen limitation and outcrossing rates. Pollinator‐mediated selfing and geitonogamy likely alleviates pollen limitation in the case of reduced mate availability, while reduced pollinator availability (intraspecific competition for pollinator services) may result in the maintenance of high outcrossing rates despite reduced seed production.     

Pollen limitation,species’ floral traits and pollinator visitation: different relationships in contrasting communities

Failures in the process of pollen transfer among conspecific plants can severely impact female reproductive success. Thus, pollen limitation can cause selection on plant mating systems and floral traits. The relationships between pollen limitation and floral traits might be partly mediated by the quantity and identity of pollinator visits. However, very little is known about the relationship between pollinator visits and pollen limitation. We examined the relationships between pollen limitation and floral traits at the community level to connect them to community ecology processes. We used 48 plant species from two contrasting communities: one species‐rich lowland community and one species‐poor alpine community. In addition, we calculated visitation rates and ecological pollination generalization for 38 of the species to examine the relationship between pollinator visitation and pollen limitation at the community level. We found low overall levels of pollen limitation that did not differ significantly between the alpine and the lowland community. In both communities, species with evolutionary specialized flowers were more pollen limited than species with unspecialized flowers. Species’ visitation rates and selfing capability were negatively related to pollen limitation in the alpine community, where pollinators are scarcer. However, flower size/number, ecological generalization of plants and flowering onset had greater effects on pollen limitation levels at the lowland community, indicating that the identity of the visitors and plant‐plant competitive interactions are more decisive for plant reproduction in this species‐rich community. There, pollen limitation increased with flower size and flowering onset, and decreased with ecological generalization, but only in species with evolutionary specialized flowers. Our study suggests that selection on plant mating system and floral traits may be idiosyncratic to each particular community and highlights the benefits of conducting community‐level studies for a better understanding of the processes underlying evolutionary responses to pollen limitation.     

Floral size variation in Campanula rotundifolia (Campanulaceae) along altitudinal gradients: patterns and possible selective mechanisms

We investigated patterns of flower‐size variation along altitudinal gradients in the bee‐pollinated perennial Campanula rotundifolia (Campanulaceae) by examining 22 Norwegian populations at altitudes between 240 and 1100 m a.s.l. We explored potential mechanisms for the underlying pattern by quantifying pollinator–faunal composition, pollinator‐visitation rates and pollen limitation of seed set in subsets of the study populations. Despite a decrease in plant size, several measures of flower size increased with elevation. Bumble bees were the main pollinators at both alpine and lowland sites in the study area. However, species composition of the pollinator fauna differed, and pollinators were larger in higher‐elevation than in lower‐elevation sites. Pollinator visitation rates were lower at higher‐elevations than at lower elevations. Pollen limitation of seed set did not vary significantly with altitude. Our results are consistent with differences in bumble‐bee size and visitation rates as causal mechanisms for the relatively larger flowers at higher elevations, in three non‐mutually exclusive ways: 1) Larger flowers reflect selection for increased attractiveness where pollinators are rare. 2) Larger and fewer flowers represent a risk avoidance strategy where the probability of pollination is low on any given day. 3) Flower size variation reflects selection to improve the fit of pollinators with fertile structures by matching flower size to pollinator size across sites.     

The inbreeding depression cost of selfing: Importance of flower size and population size in Collinsia parviflora (Veronicaceae)

Inbreeding depression should evolve with selfing rate when frequent inbreeding results in exposure of and selection against deleterious alleles. The selfing rate may be modified by plant traits such as flower size, or by population characteristics such as census size that can affect the probability of biparental inbreeding. Here we quantify inbreeding depression (δ) among different population sizes of Collinsia parviflora, a wildflower with interpopulation variation in flower size, by comparing fitness components and multiplicative fitness of experimentally produced selfed and outcrossed offspring. Selfed offspring had reduced multiplicative fitness compared to outcrossed offspring, but inbreeding depression was low in all combinations of population size and flower size (δ ≤ 0.05) except in large populations of large-flowered plants (δ = 0.45). The decrement to multiplicative fitness with inbreeding was not affected by population size nested within flower size, but differed between small- and large-flowered plants: small-flowered populations had lower overall inbreeding depression (δ = 0.04) compared to large-flowered populations (δ = 0.25). The difference in load with flower size suggests that either selection has removed deleterious recessive alleles or these alleles have become fixed in small-flowered, potentially more selfing populations, but that purging has not occurred to the same extent in presumably outcrossing large-flowered populations.     

Timing of self-compatibility, flower longevity, and potential for male outcross success in Leptosiphon jepsonii (Polemoniaceae)

When fertilization triggers flower senescence, early autonomous selfing may cause flowers to senesce before pollen has dispersed, discounting unused pollen. Selfing-induced flower senescence was examined in Leptosiphon jepsonii, a species that varies in the timing of self-compatibility. In field and greenhouse experiments, fertilization had a large effect on flower senescence; most outcrossed flowers senesced after 1 d whereas emasculated flowers lasted 2-5 d. In a comparison of inbred lines from three populations, longevity of autonomously selfed flowers of early self-compatible individuals was significantly less than that of late self-compatible individuals. In field experiments, autonomously selfed flowers were shorter-lived in a predominantly early-selfing population than in a predominantly late-selfing population. Pollen was available and viable beyond the first day of anthesis, suggesting that reductions in flower longevity caused by autonomous selfing could incur a cost to male outcross fitness. We argue that this effect is likely to be most pronounced under intermediate rates of pollinator visitation. Observed pollinator visitation rates ranged from 0.035-0.775 visits per flower per day, indicating a potential for selfing-induced flower senescence to incur pollen discounting in Leptosiphon jepsonii.     

Ecological implications of a flower size/number trade-off in tropical forest trees

BACKGROUND: In angiosperms, flower size commonly scales negatively with number. The ecological consequences of this trade-off for tropical trees remain poorly resolved, despite their potential importance for tropical forest conservation. We investigated the flower size number trade-off and its implications for fecundity in a sample of tree species from the Dipterocarpaceae on Borneo. METHODOLOGY/PRINCIPAL FINDINGS: We combined experimental exclusion of pollinators in 11 species, with direct and indirect estimates of contemporary pollen dispersal in two study species and published estimates of pollen dispersal in a further three species to explore the relationship between flower size, pollinator size and mean pollen dispersal distance. Maximum flower production was two orders of magnitude greater in small-flowered than large-flowered species of Dipterocarpaceae. In contrast, fruit production was unrelated to flower size and did not differ significantly among species. Small-flowered species had both smaller-sized pollinators and lower mean pollination success than large-flowered species. Average pollen dispersal distances were lower and frequency of mating between related individuals was higher in a smaller-flowered species than a larger-flowered confamilial. Our synthesis of pollen dispersal estimates across five species of dipterocarp suggests that pollen dispersal scales positively with flower size. CONCLUSIONS AND THEIR SIGNIFICANCE: Trade-offs embedded in the relationship between flower size and pollination success contribute to a reduction in the variance of fecundity among species. It is therefore plausible that these processes could delay competitive exclusion and contribute to maintenance of species coexistence in this ecologically and economically important family of tropical trees. These results have practical implications for tree species conservation and restoration. Seed collection from small-flowered species may be especially vulnerable to cryptic genetic erosion. Our findings also highlight the potential for differential vulnerability of tropical tree species to the deleterious consequences of forest fragmentation.     

Differences in floral traits and flower visitation rates in mating systems in Prunella vulgaris (Lamiaceae)

夏枯草交配系统对花特征和访花频率差异的影响 植物花特征和传粉者的访问次数与交配系统类型密切相关。唇形科植物夏枯草(Prunella vulgaris)存 在两种植株类型,分别为柱头伸出花冠和柱头在花冠内部的植株,而且两种植株的比例在不同种群中存在差异。本研究选择柱头伸出花冠外植株占绝大多数、柱头伸出花冠外植株占多数和柱头在花冠内部植株占多数的3个种群,通过比较每个种群中两种植株类型的开花物候、花形态特征、昆虫访问频率、自交能力、传粉者对结实的贡献以及近交衰退的水平,以检验花特征和传粉者访问次数与交配系统类型的关系。研究结果表明,与柱头在花冠内部的植株相比,柱头伸出花冠外的植株具有更大和更多的花,产生更多的花粉和花蜜,具有更高的访花频率,并主要通过异交产生种子。在种群水平,柱头伸出花冠外的植株占多数种群的访花频率显著高于柱头在花冠内部植株占多数的访花频率。柱头在花冠内部的植株比柱头伸出花冠外的植株具有更强的自动自交能力,在传粉者缺乏时为其提供了繁殖保障,但繁殖保障和异交率在不同种群中差异不显著,表明较低的昆虫访问能够满足夏枯草的授粉需求以产生种子,这可能与夏枯草较少的胚珠数量(每朵花仅有4个胚珠)有关。柱头在花冠内部植株的近交衰退水平低于柱头伸出花冠外植株的近交衰退水平,但两种植株类型的近交衰退水平均低于0.5,说明近交衰退不足以阻止该物种中自交的进化。综上所述,柱头在花冠内部的植株能够通过自交为夏枯草提供繁殖保障,而柱头伸出花冠外的植株能够利用昆虫传粉确保异交,表明混合交配系统在该物种中是一个稳定的状态。     

Effects of patch size and density on flower visitation and seed set of wild plants: a pan-European approach

1.  Habitat fragmentation can affect pollinator and plant population structure in terms of species composition, abundance, area covered and density of flowering plants. This, in turn, may affect pollinator visitation frequency, pollen deposition, seed set and plant fitness.
2.  A reduction in the quantity of flower visits can be coupled with a reduction in the quality of pollination service and hence the plants' overall reproductive success and long-term survival. Understanding the relationship between plant population size and/or isolation and pollination limitation is of fundamental importance for plant conservation.
3.  We examined flower visitation and seed set of 10 different plant species from five European countries to investigate the general effects of plant populations size and density, both within (patch level) and between populations (population level), on seed set and pollination limitation.
4.  We found evidence that the effects of area and density of flowering plant assemblages were generally more pronounced at the patch level than at the population level. We also found that patch and population level together influenced flower visitation and seed set, and the latter increased with increasing patch area and density, but this effect was only apparent in small populations.
5.   Synthesis. By using an extensive pan-European data set on flower visitation and seed set we have identified a general pattern in the interplay between the attractiveness of flowering plant patches for pollinators and density dependence of flower visitation, and also a strong plant species-specific response to habitat fragmentation effects. This can guide efforts to conserve plant–pollinator interactions, ecosystem functioning and plant fitness in fragmented habitats.     

Pollination in Verbascum thapsus (Scrophulariaceae): the advantage of being tall

According to the "effective pollination" hypothesis, tall stature resulting from strong apical dominance attracts greater pollinator visitation, thus allowing larger pollen loads and/or greater outcrossing rates, which in turn produces more vigorous offspring with greater genotypic variability and/or less inbreeding depression. Components of this hypothesis were tested in Verbascum thapsus, which commonly grows unbranched to over 2 m tall with strong apical dominance suppressing all axillary meristems. A natural population survey indicated that plants with visiting pollinators were significantly taller than their nearest neighboring individuals not possessing a visiting pollinator. Plants in natural populations with excluded pollinators produced seeds via a delayed selfing mechanism. However, delayed selfing under pollinator exclusion resulted in only 75% of the seed set obtained with natural pollinators. Under natural pollination, emasculated flowers experienced a 50% reduction in pollen deposition by the time of flower closure but only a 5% reduction in seed set relative to intact flowers. Hence, taller plants attracted more pollinators and maximum seed set could not be achieved without pollinators. Comparison of seed set and seed mass in plants that were artificially selfed and artificially crossed (in both the greenhouse and in natural populations) indicated that plants were fully self-compatible with no evidence of early-acting inbreeding depression. However, this does not exclude the possibility that inbreeding depression is manifested in later life stages. The results suggest that V. thapsus has a mixed mating system with potential for reproductive assurance and various levels of outcrossing depending on variables affecting pollinator availability (e.g., population size).     

Within-population spatial variation in pollinator visitation rates, pollen limitation on seed set, and flower longevity in an alpine species

Pollen limitation through insufficient pollen deposition on stigmas caused by too infrequent pollinator visitation may influence the reproductive outcome of plants. In this study we investigated how pollinator visitation rate, the degree of pollen limitation, and flower longevity varied spatially among three sites at different altitudes within a population of the dwarf shrub Dryas octopetala L. in alpine southern Norway. Significant pollen limitation on seed set only occurred at the mid-elevation site, while seed set at the other sites appeared to be mainly resource limited, thus indicating a spatial variation in pollen limitation. There was no association between the spatial variation in the extent of pollen limitation and pollinator visitation rate to flowers. However, pollinator visitation rates were related to flower longevity of Dryas; sites with low visitation rates had long-lived flowers and vice versa. Thus, our results suggest within-population spatial co-variation between pollinator visitation rates, pollen limitation, and a developmental response to these factors, flower longevity.     

Evaluating pollen flow indicators for an insect-pollinated plant species

For insect-pollinated plant species, reproductive success and genetic exchange via the transfer of pollen between flowers depends (i.a.) on the efficiency, abundance and behaviour of floral visitors. These in turn are expected to respond to plant population size and flower density. High floral densities for example usually attract large numbers of pollinators that visit more flowers per plant or patch, which increases pollen deposition at short distances. Thus, population characteristics might serve as indicators for pollen dispersal patterns and help to identify suitable habitat size and quality for conservation measures. To test this hypothesis, we observed floral visitors of a generalist, entomophilous species, Comarum palustre, and compared their abundance and visitation rates in populations of different sizes and flower densities. At the same time, we mimicked pollen flow using fluorescent dye. In the large and dense populations, pollinator abundance and visitation rates were high and dye was dispersed to the edges of the populations (up to 200 m). In the medium-sized population with high flower density, insect abundance and visitation rates were unexpectedly low and dye dispersal declined very quickly. On the contrary, in the smallest population with scattered flowers, especially bumble bee abundance was similar to the large populations and dye dispersal mirrored this high bumble bee activity. Thus, our results indicate that in smaller habitat fragments, the mere size of a population might be insufficient to suggest pollen flow for a plant species. Instead, the abundance of its major pollinators should be considered.     

The effects of floral display on pollinator visitation vary among populations ofPhacelia linearis (Hydrophyllaceae)

Summary Individual plants in gynodioecious populations ofPhacelia linearis (Hydrophyllaceae) vary in flower gender, flower size, and flower number. This paper reports the effects of variation in floral display on the visitation behaviour of this species' pollinators (mainly pollen-collecting solitary bees) in several natural and three experimental plant populations, and discusses the results in terms of the consequences for plant fitness. The working hypotheses were: (1) that because female plants do not produce pollen, pollen-collecting insects would visit hermaphrodite plants at a higher rate than female plants and would visit more flowers per hermaphrodite than per female; and (2) that pollinator arrival rate would increase with flower size and flower number, the two main components of visual display. These hypotheses were generally supported, but the effects of floral display on pollinator visitation varied substantially among plant populations. Hermaphrodites received significantly higher rates of pollinator arrivals and significantly higher rates of visits to flowers than did females in all experimental populations. Flower size affected arrival rate and flower visit rate positively in natural populations and in two of the three experimental populations. The flower size effect was significant only among female plants in one experimental population, and only among hermaphrodites in another. The effect of flower number on arrival rate was positive and highly significant in natural populations and in all experimental populations. In two out of three experimental populations, insects visited significantly more flowers per hermaphrodite than per female and visited more flowers on many-flowered plants than on few-flowered plants, but neither effect was detected in the third experimental population. Because seed production is not pollen-limited in this species, variation in pollinator visitation behaviour should mainly affect the male reproductive success of hermaphrodite plants. These findings suggest that pollinator-mediated natural selection for floral display inP. linearis varies in space and time.     

Pollinator population size and pollination ecosystem service responses to enhancing floral and nesting resources

Modeling pollination ecosystem services requires a spatially explicit, process‐based approach because they depend on both the behavioral responses of pollinators to the amount and spatial arrangement of habitat and on the within‐ and between‐season dynamics of pollinator populations in response to land use. We describe a novel pollinator model predicting flower visitation rates by wild central‐place foragers (e.g., nesting bees) in spatially explicit landscapes. The model goes beyond existing approaches by: (1) integrating preferential use of more rewarding floral and nesting resources; (2) considering population growth over time; (3) allowing different dispersal distances for workers and reproductives; (4) providing visitation rates for use in crop pollination models. We use the model to estimate the effect of establishing grassy field margins offering nesting resources and a low quantity of flower resources, and/or late‐flowering flower strips offering no nesting resources but abundant flowers, on bumble bee populations and visitation rates to flowers in landscapes that differ in amounts of linear seminatural habitats and early mass‐flowering crops. Flower strips were three times more effective in increasing pollinator populations and visitation rates than field margins, and this effect increased over time. Late‐blooming flower strips increased early‐season visitation rates, but decreased visitation rates in other late‐season flowers. Increases in population size over time in response to flower strips and amounts of linear seminatural habitats reduced this apparent competition for pollinators. Our spatially explicit, process‐based model generates emergent patterns reflecting empirical observations, such that adding flower resources may have contrasting short‐ and long‐term effects due to apparent competition for pollinators and pollinator population size increase. It allows exploring these effects and comparing effect sizes in ways not possible with other existing models. Future applications include species comparisons, analysis of the sensitivity of predictions to life‐history traits, as well as large‐scale management intervention and policy assessment.     

Pollination ecology of Silene acutifolia (Caryophyllaceae): floral traits variation and pollinator attraction

BACKGROUND AND AIMS: The floral display influences the composition of pollinators interacting with a plant species. Geographic and temporal variation in pollinator composition complicates the understanding of the evolutionary consequences of floral display variation. This paper analyses the relationships between Silene acutifolia, a hermaphroditic perennial herb, and its pollinators, based on field studies in the north-west of Spain. METHODS: Studies were conducted over three years (1997-1999). Firstly, the main pollinators of this species were determined for two years in one population. Secondly, pollen limitation in fruit and seed production was analysed by supplementary hand pollinations, and counting the pollen grains and tubes growing in styles for two different-sized populations. Finally, the effect of flower size and number on the rate of visitation and total seed number was examined for 15 marked plants. RESULTS AND CONCLUSIONS: The primary pollinators were long-tongued insects, including Hymenoptera, Lepidoptera and Diptera, but the composition and visitation frequencies differed between years. Pollen limitation occurred in one of the years of study. There was between-population variation in the number of pollen grains and pollen tubes found in styles, suggesting pollen limitation in one population. Overall, pollinators visited plants with more open flowers more frequently, and pollinated more flowers within these plants. Conversely, petal and calyx sizes had no effect on insect visitation. Plants with higher rates of visits produced higher number of seeds, suggesting that pollinator-mediated limitation of seed and fruit production may be important in some years.     

Pollinator response to female and male floral display in a monoecious species and its implications for the evolution of floral dimorphism

Pollinator-mediated selection has been hypothesized as one cause of size dimorphism between female and male flowers. Flower number, ignored in studies of floral dimorphism, may interact with flower size to affect pollinator selectivity. In the present study, we explored pollinator response, and estimated pollen receipt and removal, in experimental populations of monoecious Sagittaria trifolia, in which plants were manipulated to display three, six, nine or 12 female or male flowers per plant. In this species, female flowers are smaller but have a more compressed flowering period than males, creating larger female floral displays. Overall, pollinators preferred to visit male rather than female displays of the same size. Both first visit per foraging bout and visitation rates to female displays increased with display size. However, large male displays did not show increased attractiveness to pollinators. A predicted relationship that pollen removal, rather than pollen receipt, is limited by pollinator visitation was confirmed in the experimental populations. The results suggest that the lack of selection on large male displays may affect the evolution of floral dimorphism in this species.     

Floral dimorphism, pollination, and self-fertilization in gynodioecious GERANIUM RICHARDSONII (Geraniaceae)

The selective maintenance of gynodioecy depends on the relative fitness of the male-sterile (female) and hermaphroditic morphs. Females may compensate for their loss of male fitness by reallocating resources from male function (pollen production and pollinator attraction) to female function (seeds and fruits), thus increasing seed production. Females may also benefit from their inability to self-fertilize if selfing and inbreeding depression reduce seed quality in hermaphrodites. We investigated how differences in floral resource allocation (flower size) between female and hermaphroditic plants affect two measures of female reproductive success, pollinator visitation and pollen receipt, in gynodioecious populations of Geranium richardsonii in Colorado. Using emasculation treatments in natural populations, we further examined whether selfing by autogamy and geitonogamy comprises a significant proportion of pollen receipt by hermaphrodites. Flowers of female plants are significantly smaller than those of hermaphrodites. The reduction in allocation to pollinator-attracting structures (petals) is correlated with a significant reduction in pollinator visitation to female flowers in artificial arrays. The reduction in attractiveness is further manifested in significantly less pollen being deposited on the stigmas of female flowers in natural populations. Autogamy is rare in these protandrous flowers, and geitonogamy accounts for most of the difference in pollen receipt between hermaphrodites and females. Female success at receiving pollen was negatively frequency dependent on the relative frequency of females in populations. Thus, two of the prerequisites for the maintenance of females in gynodioecious populations, differences in resource allocation between floral morphs and high selfing rates in hermaphrodites, occur in G. richardsonii.