The timing of visits by large and small bees differentially affects pollination success in Mimulus ringens.

PREMISE: Cross-fertilization in most flowering plants is facilitated by mobile animals that transport pollen while foraging for floral rewards. The contributions of different visitors can vary widely, depending on the amount of pollen transferred during a single visit and on the frequency and timing of the visits of each pollinator taxon. METHODS: We used three approaches to measure the pollination value of bees that visit Mimulus ringens: pollinator interviews, field population observations, and caging studies. RESULTS: The single-visit effectiveness of small bees (primarily Halictidae) was only half that of larger bees (primarily Bombus) for pollen delivery and removal. In five field populations, we found substantial temporal and spatial variation in visitation and pollination. In most sites big bees were active before 08:00 hours, and by 10:00-11:00 hours, stigmas were usually fully pollinated and closed, and little pollen remained in anthers. Small bees seldom visited before 10:00 hours. Excluding big bees from plants confirmed that pollination is reduced and delayed in this ecological context. CONCLUSIONS: Big bees are the primary pollinators of M. ringens, accounting for at least 75% of seed production. Not only are they more effective per visit, in most situations they also visit before small bees become active. Although small bees are not usually important pollinators of M. ringens, they have the potential to partially replace them as a "fail-safe" pollinator in contexts where big bees are not abundant. In a world where pollinator abundance is declining, such backup pollinators may be important for maintaining plant reproduction.

Variation in flower size and shape of Impatiens capensis is correlated with urbanization in Montreal, Canada.

Urbanization is changing the conditions in which many species live, forcing them to adjust to these novel environments. Floral size and shape are critical traits for the reproduction of plants pollinated by animals as they are involved in the attraction of pollinators and in efficient pollination. Variation in size and shape could be affected by urbanization via its modification of the abiotic environment (habitat fragmentation, water availability, temperature, soil properties), or via its impact on the biotic environment of plants (pollination, herbivory). Although numerous studies have assessed the impact of urbanization on pollinator communities and many plant traits, few have investigated its impact on floral size and shape while quantifying the proportion of the total urbanization effect that is due to biotic interactions. In this study, we tested if urbanization and pollinator visitation rates affect the flower shape of the spotted jewelweed, Impatiens capensis. We quantified the size and shape of flowers in frontal and profile views using geometric morphometrics for 228 individuals from six populations from the region of Montreal, Canada. Pollinator visitation rates were estimated at each site and the main pollinators were found to be bumblebees, honeybees and hummingbirds. We found that floral size and shape are significantly correlated with urbanization as measured by the amount of vegetation in the surrounding environment of the plants (mean normalized vegetation index, NDVI) and by the visitation rates of bumblebees and honey bees. Partitioning of the total flower shape variation suggests that urbanization affects flower shape through abiotic factors and via its impact on pollinator visitation rates. While further studies from other cities are necessary to confirm the role of urbanization in shaping the floral shape of I. capensis, these results support the idea that urbanization could affect flower shapes.

The rapid appearance of homostyly in a cultivated distylous population of Primula forbesii.

Evolutionary breakdown from rigorous outbreeding to self-fertilization frequently occurs in angiosperms. Since the pollinators are not necessary, self-compatible populations often reduce investment in floral display characteristics and pollination reward. Primula forbesii is a biennial herb with distribution restricted to southwest China; it was initially a self-incompatible distylous species, but after 20 years of artificial domestication, homostyly appeared. This change in style provides an ideal material to explore the time required for plant mating systems to adapt to new environmental changes and test whether flower attraction has reduced following transitions to selfing. We did a survey in wild populations of P. forbesii where its seeds were originally collected 20 years ago and recorded the floral morph frequencies and morphologies. The floral morphologies, self-incompatibility, floral scent, and pollinator visitation between distyly and homostyly were compared in greenhouse. Floral morph frequencies of wild populations did not change, while the cultivated population was inclined to L-morph and produced homostyly. Evidence from stigma papillae and pollen size supports the hypothesis that the homostyly possibly originated from mutations of large effect genes in distylous linkage region. Transitions to self-compatible homostyly are accompanied by smaller corolla size, lower amounts of terpenoids, especially linalool and higher amounts of fatty acid derivatives. The main pollinators in the greenhouse were short-tongued Apis cerana. However, homostyly had reduced visiting frequency. The mating system of P. forbesii changed rapidly in just about 20 years of domestication, and our findings confirm the hypothesis that the transition to selfing is accompanied by decreased flower attraction.

Continuous inbreeding affects genetic variation, phenology, and reproductive strategy in ex situ cultivated Digitalis lutea.

PREMISE: Ex situ cultivation is important for plant conservation, but cultivation in small populations may result in genetic changes by drift, inbreeding, or unconscious selection. Repeated inbreeding potentially influences not only plant fitness, but also floral traits and interactions with pollinators, which has not yet been studied in an ex situ context. METHODS: We studied the molecular genetic variation of Digitalis lutea from a botanic garden population cultivated for 30 years, a frozen seed bank conserving the original genetic structure, and two current wild populations including the source population. In a common garden, we studied the effects of experimental inbreeding and between-population crosses on performance, reproductive traits, and flower visitation of plants from the garden and a wild population. RESULTS: Significant genetic differentiation was found between the garden population and the wild population from which the seeds had originally been gathered. After experimental selfing, inbreeding depression was only found for germination and leaf size of plants from the wild population, indicating a history of inbreeding in the smaller garden population. Moreover, garden plants flowered earlier and had floral traits related to selfing, whereas wild plants had traits related to attracting pollinators. Bumblebees visited more flowers of outbred than inbred plants and of wild than garden plants. CONCLUSIONS: Our case study suggests that high levels of inbreeding during ex situ cultivation can influence reproductive traits and thus interactions with pollinators. Together with the effects of genetic erosion and unconscious selection, these changes may affect the success of reintroductions into natural habitats.

Pollinators mediate floral microbial diversity and microbial network under agrochemical disturbance.

How pollinators mediate microbiome assembly in the anthosphere is a major unresolved question of theoretical and applied importance in the face of anthropogenic disturbance. We addressed this question by linking visitation of diverse pollinator functional groups (bees, wasps, flies, butterflies, beetles, true bugs and other taxa) to the key properties of the floral microbiome (microbial α- and ß-diversity and microbial network) under agrochemical disturbance, using a field experiment of bactericide and fungicide treatments on cultivated strawberries that differ in flower abundance. Structural equation modelling was used to link agrochemical disturbance and flower abundance to pollinator visitation to floral microbiome properties. Our results revealed that (i) pollinator visitation influenced the α- and ß-diversity and network centrality of the floral microbiome, with different pollinator functional groups affecting different microbiome properties; (ii) flower abundance influenced the floral microbiome both directly by governing the source pool of microbes and indirectly by enhancing pollinator visitation; and (iii) agrochemical disturbance affected the floral microbiome primarily directly by fungicide, and less so indirectly via pollinator visitation. These findings improve the mechanistic understanding of floral microbiome assembly, and may be generalizable to many other plants that are visited by diverse insect pollinators in natural and managed ecosystems.

Selfing rates vary with floral display, pollinator visitation and plant density in natural populations of Mimulus ringens.

Variation in selfing rates within and among populations of hermaphroditic flowering plants can strongly influence the evolution of reproductive strategies and the genetic structure of populations. This intraspecific variation in mating patterns may reflect both genetic and ecological factors, but the relative importance of these factors remains poorly understood. Here, we explore how selfing in 13 natural populations of the perennial wildflower Mimulus ringens is influenced by (a) pollinator visitation, an ecological factor, and (b) floral display, a trait with a genetic component that also responds to environmental variation. We also explore whether genetically based floral traits, including herkogamy, affect selfing. We found substantial variation among populations in selfing rate (0.13-0.55). Selfing increased strongly and significantly with floral display, among as well as within populations. Selfing also increased at sites with lower pollinator visitation and low plant density. However, selfing was not correlated with floral morphology. Overall, these results suggest that pollinator visitation and floral display, two factors that interact to affect geitonogamous pollinator movements, can influence the selfing rate. This study identifies mechanisms that may play a role in maintaining selfing rate variation among populations.

Floral and pollination characteristics of Eriocaulon heleocharioides, an extinct species in the wild, for evidence-based conservation management.

Generally, floral characteristics and pollination are important factors enhancing the quality and quantity of reproductive output for regeneration in plant conservation. However, lack of evidence-based management could decrease fitness under ex-situ conservation. We investigated the capitulum and pollination characteristics of Eriocaulon heleocharioides Satake (Eriocaulaceae), which is extinct in the wild, to develop an evidence-based conservation management plan incorporating previously ignored reproductive characteristics. To evaluate the functional characteristics of capitula, pollen-ovule ratio, and reproductive status (maximum pollination success/florivory damage) were investigated along six flowering sequences of capitulum. To evaluate the effect of plant density on pollen transfer, high- and low-density plots were established. Total deposited pollen on stigma, insect visitation, and visit duration per capitulum were observed. A significantly lower pollen-ovule ratio was observed in the first of six capitula, reflecting higher female functionality. The highest pollination success was found in the second-fourth capitula, whereas florivory increased along the terminal capitula position. High plant density affected the pollen deposited on stigmas via insect visitation and low pollinator visit duration. Different capitula in E. heleocharioides could have different effects: different sexual functionality, enhancement of reproductive output both in quality and quantity through active pollen transfer, and escaping from florivores. High plant density could facilitate outcross-pollen transfer in E. heleocharioides. Multiple perspectives are important for determining potential reproductive success in ex-situ conservation. Thus, density management reflecting capitulum characteristics could improve the efficiency of conservation efforts for E. heleocharioides.

Limited phenological and pollinator-mediated isolation among selfing and outcrossing Arabidopsis lyrata populations.

Transitions from outcrossing to selfing have been a frequent evolutionary shift in plants and clearly play a role in species divergence. However, many questions remain about the initial mechanistic basis of reproductive isolation during the evolution of selfing. For instance, how important are pre-zygotic pre-pollination mechanisms (e.g. changes in phenology and pollinator visitation) in maintaining reproductive isolation between newly arisen selfing populations and their outcrossing ancestors? To test whether changes in phenology and pollinator visitation isolate selfing populations of Arabidopsis lyrata from outcrossing populations, we conducted a common garden experiment with plants from selfing and outcrossing populations as well as their between-population hybrids. Specifically, we asked whether there was isolation between outcrossing and selfing plants and their between-population hybrids through differences in (1) the timing or intensity of flowering; and/or (2) pollinator visitation. We found that phenology largely overlapped between plants from outcrossing and selfing populations. There were also no differences in pollinator preference related to mating system. Additionally, pollinators preferred to visit flowers on the same plant rather than exploring nearby plants, creating a large opportunity for self-fertilization. Overall, this suggests that pre-zygotic pre-pollination mechanisms do not strongly reproductively isolate plants from selfing and outcrossing populations of Arabidopsis lyrata.

Pollinator visitation rate and effectiveness vary with flowering phenology.

PREMISE: Flowering time may influence pollination success and seed set through a variety of mechanisms, including seasonal changes in total pollinator visitation or the composition and effectiveness of pollinator visitors. METHODS: We investigated mechanisms by which changes in flowering phenology influence pollination and reproductive success of Mertensia ciliata (Boraginaceae). We manipulated flowering onset of potted plants and assessed the frequency and composition of pollinator visitors, as well as seed set. We tested whether floral visitors differed in their effectiveness as pollinators by measuring pollen receipt and seed set resulting from single visits to virgin flowers. RESULTS: Despite a five-fold decrease in pollinator visitation over four weeks, we detected no significant difference in seed set among plants blooming at different times. On a per-visit basis, each bumblebee transferred more conspecific pollen than did a solitary bee or a fly. The proportion of visits by bumblebees increased over the season, countering the decrease in visitation rate so that flowering time had little net effect on seed set. CONCLUSIONS: This work illustrates the need to consider pollinator effectiveness, along with changes in pollinator visitation and species composition to understand the mechanisms by which phenology affects levels of pollination.

Continuous variation in herkogamy enhances the reproductive response of Lonicera implexa to spatial variation in pollinator assemblages.

Herkogamy, the spatial separation of sex organs in hermaphroditic plants, has been proposed as a mechanism to reduce self-pollination and the associated processes of inbreeding and gamete wastage. Longitudinal herkogamy is the most frequent type, with two subtypes: approach herkogamy (anthers below the stigma), which is associated with diverse pollinator arrays, and reverse herkogamy (anthers above the stigma), associated with specialized, long-tongued pollinators. By using a herkogamy index that varied continuously from negative (reverse herkogamy) to positive (approach herkogamy) values, we studied the effect of continuous variation in herkogamy on pollinator attraction, selfing capability and plant fitness across three populations of Lonicera implexa differing in the relative abundance of long-tongued vs. short-tongued pollinators. Reverse herkogamy was significantly more frequent in the population where long-tongued pollinators were dominant than in the other two populations. Agreeing with this, the main floral visitors of L. implexa individuals with small and large herkogamy index were, respectively, long-tongued and short-tongued pollinators. Spontaneous selfing was low and increased with increasing herkogamy index (i.e. with approach herkogamy), although most of it occurred when there was close distance between anthers and stigma. Fruit production was unrelated to the herkogamy index in the population with long-tongued pollinators, but it increased with approach herkogamy (higher herkogamy index) in the other two populations. In contrast, seeds of individuals with reverse herkogamy (smaller herkogamy indices) germinated better. In this species, continuous variation in herkogamy might function as a reproductive strategy, as different morphotypes might be favoured by different pollinator assemblages.

Effect of Pollen Limitation and Pollinator Visitation on Pollination Success of Haloxylon ammodendron (C. A. Mey.) Bunge in Fragmented Habitats.

Haloxylon ammodendron (C. A. Mey.) Bunge is an ecologically important species in arid regions. Pollen limitation may decrease plant reproduction due to low levels of pollen transfer and inadequate pollen receipt. In arid regions, pollen limitations of many plant species may be influenced by habitat fragmentation. However, whether pollen limitation and pollinator visitation affect the pollination success of H. ammodendron (Amaranthaceae) in fragmented habitats still needs further study. In this study, we calculated the pollen limitation in natural and fragmented habitats to estimate the effect of habitat fragmentation on pollen limitation. In different habitats, we investigated the relationship between the number of open flowers and pollinator visiting frequency. In addition, we examined how habitat fragmentation affects pollination success through the influence of pollinator visitation rate on seed set. Our results indicated that pollen limitation was the important limiting factor for seed set in fragmented and natural habitats. The results showed higher pollinator visitation rates resulted in a higher percentage of seeds in both habitats. In H. ammodendron, Apis mellifera was found to be the dominant pollinator. These results may support the assertion that plants evolve traits to attract pollinators and pollinators increase their visiting frequency to better exploit the floral resources. We also determined that outcrossing was dominant in the breeding system and that wind pollination played an important role in pollination success. This study aims to contribute to a better understanding of how environmental heterogeneity affects pollen limitation, pollinator visitation, and pollination success in arid regions.

Non-native honey bees disproportionately dominate the most abundant floral resources in a biodiversity hotspot.

Most plant-pollinator mutualisms are generalized. As such, they are susceptible to perturbation by abundant, generalist, non-native pollinators such as the western honey bee ( Apis mellifera), which can reach high abundances and visit flowers of many plant species in their expansive introduced range. Despite the prevalence of non-native honey bees, their effects on pollination mutualisms in natural ecosystems remain incompletely understood. Here, we contrast community-level patterns of floral visitation by honey bees with that of the diverse native pollinator fauna of southern California, USA. We show that the number of honey bees visiting plant species increases much more rapidly with flower abundance than does that of non-honey bee insects, such that the percentage of all visitors represented by honey bees increases with flower abundance. Thus, honey bees could disproportionately impact the most abundantly blooming plant species and the large numbers of both specialized and generalized pollinator species that they sustain. Honey bees may preferentially exploit high-abundance floral resources because of their ability to recruit nest-mates; these foraging patterns may cause native insect species to forage on lower-abundance resources to avoid competition. Our results illustrate the importance of understanding foraging patterns of introduced pollinators in order to reveal their ecological impacts.

Reproductive assurance weakens pollinator-mediated selection on flower size in an annual mixed-mating species.

BACKGROUND AND AIMS: In animal-pollinated plants, direct and indirect selection for large and small flowers in predominantly outcrossing and selfing species, respectively, is a common consequence of pollen limitation (PL). However, many hermaphroditic species show a mixed-mating system known as delayed selfing, which provides reproductive assurance (RA) only when outcrossing is not realized. Although RA is expected to reduce pollinator-mediated selection towards larger flowers, the consequences of delayed selfing for selection on flower size in mixed-mating species remain overlooked. We investigated whether RA weakens selection on flower size in Tuberaria guttata, a mixed-mating annual herb. METHODS: We related pollinator visitation rates to flower size and measured seed production in emasculated, hand cross-pollinated and intact (control) flowers in three natural populations. For each population, we estimated variation in PL and RA across individuals differing in flower size and phenotypic selection on this trait. KEY RESULTS: Pollinator visitation increased and RA decreased with flower size in all populations. Increasing RA diminished but did not fully alleviate PL, because of early-acting inbreeding depression. In the least-visited and most pollen-limited population, RA increased seed production by >200 %, intensely counteracting the strong pollinator-mediated selection for larger corollas. In the most-visited population, however, RA increased seed production by an average of only 9 %. This population exhibited the largest fraction of individuals that showed a decrease in seed production due to selfing and the weakest pollinator-mediated selection on flower size. CONCLUSIONS: The results suggest that the balance between the extent of RA and outcrossing contributes to determine flower size in mixed-mating systems. Pollinator-mediated selection favours larger flowers by increasing outcrossed seeds, but the benefits of RA greatly lessen this effect, especially under severe conditions of pollen limitation. Our findings also indicate that a mixed-mating system can represent an 'evolutionary trap' under an adequate pollinator supply.

Pollination efficiency of cockroaches and other floral visitors of Clusia blattophila.

Cockroaches have rarely been documented as pollinators. In this paper we examine whether this is because they might be inefficient at pollination compared to other pollinators. Clusia blattophila, a dioecious shrub growing on isolated rocky outcrops in French Guiana, is pollinated by Amazonina platystylata cockroaches and provides a valuable system for the study of cockroach pollination efficiency. We examined the species composition of the visitor guild and visitation rates by means of camcorder recordings and visitor sampling. Then, we investigated the capacity for pollen transfer of principal visitors and found correlations between visitation rates and pollen loads on stigmas. In an exclusion experiment we determined the contributions of individual species to pollination success. Amazonina platystylata, crickets and two species of Diptera transferred pollen, but the number of transferred pollen grains was only related to visitation rates in the case of cockroaches. Crickets visited and rarely carried pollen. Dipterans were as frequent as cockroaches, carried similar pollen loads, but transferred much less pollen. An estimated 41% and 17% of ovules were pollinated by cockroaches and dipterans, respectively. The remaining ovules were not pollinated. There was no spatial variation in pollinator guild composition, but cockroaches visited flowers less frequently at the smaller study site. We demonstrate that cockroaches pollinate a large proportion of ovules. Their pollination service is not confined to one study site and, unlike that provided by dipterans, is not limited to certain years. We suggest that cockroach pollination has been overlooked and that cockroach-pollinated plants, which share certain floral features, possess adaptations to pollination by cockroaches.

One for all and all for one: retention of colour-unchanged old flowers increases pollinator attraction in a hermaphroditic plant.

Long-lived flowers increase pollen transfer rates, but these entail high water and carbon maintenance costs. The retention of pollinated and reward-free old flowers enhances pollinator visitation to young receptive flowers by increasing floral display size. This mechanism is associated with acropetal inflorescences or changes in flower colour and openness, but the retention of unchanging solitary flowers remains overlooked. We examined pollination-dependent variation in floral longevity and determined stigmatic receptivity, pollen viability and pollen removal rates among flower ages in Kielmeyera regalis, a Neotropical savanna shrub. We also evaluated the effects of floral display size on pollinator visitation rates. Lastly, we determined whether old flowers are unvisited and exclusively increase pollinator attraction to young flowers through flower removal experiments. Regardless of pollination treatment, flowers lasted fully open with no detectable physical changes for 3 days. Over time, stigmas remained receptive but >95% of pollen was removed. Pollinator visitation significantly increased with floral display size and intermediate percentages (15-30%) of newly opened flowers. Accordingly, the retention of reward-free and unvisited old flowers increased young flower-pollinator interaction. Our results reveal the importance of a prolonged floral longevity in increasing pollinator attraction toward newly opened receptive flowers without changes in flower colour and form. We conclude that the retention of pollinated, reward-free and unvisited colour-unchanged old flowers in K. regalis is a strategy that counteracts the water use costs associated with the maintenance of large flowers with increased mate opportunities in a pollen-limited scenario.

Pollinator activity and pollination success of Medicago sativa L. in a natural and a managed population.

Medicago sativa L. is an important cash crop in the arid region of northwest China. Pollinator activity is an essential aspect of pollination success, but the relationships between pollinator visitation rate and seed set still need further study of M. sativa. We investigated the following characteristics of M. sativa in natural and managed populations: floral traits, pollinator activity, and breeding system. Our results indicated the management could affect the number of flowers produced; however, there was no detectable effect on the seed set per flower. We found the percentage of seeds among pollinated flowers in the managed population was significantly higher than that in the natural population. Moreover, the increase in the proportion of pollinated flowers could significantly increase seed set per flower, and pollinator visitation rate was the important limiting factor for seed set in both populations. Andrena lebedevi Popov was found to be the most frequent pollinator in both populations. Outcrossing was dominant in the breeding system and insect pollination played an important role in outcrossing. Our study suggested that proper management (artificial selection) could promote pollination success of M. sativa.

The effect of removing numerically dominant, non-native honey bees on seed set of a native plant.

Pollination services are compromised by habitat destruction, land-use intensification, pesticides, and introduced species. How pollination services respond to such stressors depends on the capacity of pollinator assemblages to function in the face of environmental disruption. Here, we quantify how pollination services provided to a native plant change upon removal of the non-native, western honey bee (Apis mellifera)-a numerically dominant floral visitor in the native bee-rich ecosystems of southern California. We focus on services provided to clustered tarweed (Deinandra fasciculata), a native, annual forb that benefits from outcross pollination. Across five different study sites in coastal San Diego County, tarweed flowers attracted 70 insect taxa, approximately half of which were native bees, but non-native honey bees were always the most abundant floral visitor at each site. To test the ability of the native insect fauna to provide pollination services, we performed Apis removals within experimental 0.25 m2 plots containing approximately 20 tarweed plants and compared visitation and seed set between plants in removal and paired control plots (n = 16 pairs). Even though 92% of observed floral visits to control plots were from honey bees, Apis removal reduced seed production by only 14% relative to plants in control plots. These results indicate that native insect assemblages can contribute important pollination services even in ecosystems numerically dominated by introduced pollinators.

Pollination success following loss of a frequent pollinator: the role of compensatory visitation by other effective pollinators.

Pollinator abundance is declining worldwide and may lower the quantity and quality of pollination services to flowering plant populations. Loss of an important pollinator is often assumed to reduce the amount of pollen received by stigmas of a focal species (pollination success), yet this assumption has rarely been tested experimentally. The magnitude of the effect, if any, may depend on the relative efficiency of the remaining pollinators, and on whether the loss of one pollinator leads to changes in visitation patterns by other pollinators. To explore how a change in pollinator composition influences pollination of Asclepias verticillata, we excluded bumble bees from plots in large and small populations of this milkweed species. We then quantified pollinator visitation rates, pollen export and pollen receipt for control plots and for plots where bumble bees were experimentally excluded. We found that exclusion of bumble bees did not reduce pollen receipt by A. verticillata flowers. Visitation by Polistes wasps increased markedly following bumble bee exclusion, both in small populations (186 % increase) and in large populations (400 % increase). Because Polistes wasps were as efficient as bumble bees at pollen transfer, increased wasp visitation offset lost bumble bee pollination services. Thus, loss of a frequent pollinator will not necessarily lead to a decline in pollination success. When pollinator loss is followed by a shift in the composition and abundance of remaining pollinators, pollination success will depend on the net change in the quantity and quality of pollination services.

The effects of local variation in light availability on pollinator visitation, pollen and resource limitation of female reproduction in Hosta ventricosa.

BACKGROUND: Light availability may have direct effects on reproduction through resource availability, and indirect effects on female reproduction by influencing plant-pollinator interactions. Floral display size, pollinator visitation per flower, resource and pollen limitation of fruit and seed production were quantified in a forested patch and an adjacent open patch of two populations of the perennial herb Hosta ventricosa. RESULTS: Plants in the open patch produced significantly larger floral displays than those in the forested patch in both populations. Floral display size had a positive effect on pollinator visitation rate per flower in one population, but no effect in the other. Plants in the open patch received approximately 8-11 times more visitation rates and produced significantly more fruit and seeds per flower than those in the forested patch. However, supplemental pollination resulted in significantly more fruit and seed production per flower compared to natural pollination in the forested patch but not in the open patch in one population, and did not enhance fruit and seed production in either the forested or the open patch in the other. In both populations, supplementally pollinated plants in the open patch produced significantly more fruit and seeds per flower than supplementally pollinated plants in the forested patch. CONCLUSIONS: In H. ventricosa, local variation in light conditions could affect pollinator activity and influence female reproduction through resource availability; however differences in the degree of pollen limitation between local habitats were found to be population-specific.

Invasion of a dominant floral resource: effects on the floral community and pollination of native plants.

Through competition for pollinators, invasive plants may suppress native flora. Community-level studies provide an integrative assessment of invasion impacts and insights into factors that influence the vulnerability of different native species. We investigated effects of the nonnative herb Lythrum salicaria on pollination of native species in 14 fens of the eastern United States. We compared visitors per flower for 122 native plant species in invaded and uninvaded fens and incorporated a landscape-scale experiment, removing L. salicaria flowers from three of the invaded fens. Total flower densities were more than three times higher in invaded than uninvaded or removal sites when L. salicaria was blooming. Despite an increase in number of visitors with number of flowers per area, visitors per native flower declined with increasing numbers of flowers. Therefore, L. salicaria invasion depressed visitation to native flowers. In removal sites, visitation to native flowers was similar to uninvaded sites, confirming the observational results and also suggesting that invasion had not generated a persistent build-up of visitor populations. To study species-level impacts, we examined effects of invasion on visitors per flower for the 36 plant species flowering in both invaded and uninvaded fens. On average, the effect of invasion represented about a 20% reduction in visits per flower. We measured the influence of plant traits on vulnerability to L. salicaria invasion using meta-analysis. Bilaterally symmetrical flowers experienced stronger impacts on visitation, and similarity in flower color to L. salicaria weakly intensified competition with the invader for visitors. Finally, we assessed the reproductive consequences of competition with the invader in a dominant flowering shrub, Dasiphora fruticosa. Despite the negative effect of invasion on pollinator visitation in this species, pollen limitation of seed production was not stronger in invaded than in uninvaded sites, suggesting little impact of competition for pollinators on its population demography. Negative effects on pollination of native plants by this copiously flowering invader appeared to be mediated by increases in total flower density that were not matched by increases in pollinator density. The strength of impact was modulated across native species by their floral traits and reproductive ecology.