Benefits and limitations of isolated floral patches in a pollinator restoration project in Arizona

This study examined invertebrate floral visitor responses to floral richness, floral abundance, and distance between floral patches within a newly planted pollinator restoration habitat in an arid ecosystem in central Arizona, United States. We created a pollinator habitat experiment consisting of a large central garden (11‐m diameter) surrounded by concentric rings of smaller habitat patches (1‐m diameter), separated from one another by 1, 8, 13, and 21 m, respectively, and including four flowering species. We observed plant and visitor interactions via structured 10‐minute flower visitation observations over a 3‐month period. Key findings included: (1) each plant species interacted with a variety of flower visitors, but flower visitor groups differed only marginally among the plant species; (2) floral patches outside the central garden exhibited reduced quantities of floral structures; and (3) number of floral structures per patch, but not isolation of floral patches within the habitat, affected the number of visitors and visitor taxa richness. For practitioners and land managers looking to restore pollination systems in arid ecosystems with low establishment via seeding, the results of this study suggest that installing species‐rich and florally abundant patches of flowering plant species within a habitat could efficiently support plant‐pollinator interactions.     

Local adaptation: Mechanical fit between floral ecotypes of Nerine humilis (Amaryllidaceae) and pollinator communities

Geographic variation in floral morphology is often assumed to reflect geographic variation in pollinator communities and associated divergence in selective pressures. We studied populations of Nerine humilis (Amaryllidaceae) to assess whether geographic variation in floral form is the result of local adaptation to different pollinator communities. We first tested for associations between floral traits and visitor communities, and found that populations with similar floral morphologies were visited by similar insect communities. Mean style length in each population was also closely associated with the mean body length of the local visitor community. A reciprocal translocation experiment demonstrated that native phenotypes set more seed than translocated phenotypes. Single visitation experiments showed that native flowers received more pollen, and set more seed per visit, than introduced phenotypes in both populations. This suggests that the effectiveness of pollinator visits is determined by the degree of mechanical fit between flowers and visitors. We provide strong evidence that the observed among‐population variation in floral traits is an adaptive response to geographic variation in the pollinator community.     

Selective pollination by fungus gnats potentially functions as an alternative reproductive isolation among five Arisaema species

Background and AimsInterspecific difference in pollinators (pollinator isolation) is important for reproductive isolation in flowering plants. Species-specific pollination by fungus gnats has been discovered in several plant taxa, suggesting that they can contribute to reproductive isolation. Nevertheless, their contribution has not been studied in detail, partly because they are too small for field observations during flower visitation. To quantify their flower visitation, we used the genus Arisaema (Araceae) because the pitcher-like spathe of Arisaema can trap all floral visitors.MethodsWe evaluated floral visitor assemblage in an altitudinal gradient including five Arisaema species. We also examined interspecific differences in altitudinal distribution (geographic isolation) and flowering phenology (phenological isolation). To exclude the effect of interspecific differences in altitudinal distribution on floral visitor assemblage, we established ten experimental plots including the five Arisaema species in high- and low-altitude areas and collected floral visitors. We also collected floral visitors in three additional sites. Finally, we estimated the strength and contribution of these three reproductive barriers using a unified formula for reproductive isolation.Key ResultsEach Arisaema species selectively attracted different fungus gnats in the altitudinal gradient, experimental plots and additional sites. Altitudinal distribution and flowering phenology differed among the five Arisaema species, whereas the strength of geographic and phenological isolations were distinctly weaker than those in pollinator isolation. Nevertheless, the absolute contribution of pollinator isolation to total reproductive isolation was weaker than geographic and phenological isolations, because pollinator isolation functions after the two early-acting barriers in plant life history.ConclusionsOur results suggest that selective pollination by fungus gnats potentially contributes to reproductive isolation. Since geographic and phenological isolations can be disrupted by habitat disturbance and interannual climate change, the strong and stable pollinator isolation might compensate for the weakened early-acting barriers as an alternative reproductive isolation among the five Arisaema species.     

Asynchronous range shifts drive alpine plant–pollinator interactions and reduce plant fitness

Climate change is driving species' range shifts, which are in turn disrupting species interactions due to species‐specific differences in their abilities to migrate in response to climate. We evaluated the consequences of asynchronous range shifts in an alpine plant–pollinator community by transplanting replicated alpine meadow turfs downslope along an elevational gradient thereby introducing them to warmer climates and novel plant and pollinator communities. We asked how these novel plant–pollinator interactions affect plant reproduction. We found that pollinator communities differed substantially across the elevation/temperature gradient, suggesting that these plants will likely interact with different pollinator communities with warming climate. Contrary to the expectation that floral visitation would increase monotonically with warmer temperatures at lower elevations, visitation rate to the transplanted communities peaked under intermediate warming at midelevation sites. In contrast, visitation rate generally increased with temperature for the local, lower elevation plant communities surrounding the experimental alpine turfs. For two of three focal plant species in the transplanted high‐elevation community, reproduction declined at warmer sites. For these species, reproduction appears to be dependent on pollinator identity such that reduced reproduction may be attributable to decreased visitation from key pollinator species, such as bumble bees, at warmer sites. Reproduction in the third focal species appears to be primarily driven by overall pollinator visitation rate, regardless of pollinator identity. Taken together, the results suggest climate warming can indirectly affect plant reproduction via changes in plant–pollinator interactions. More broadly, the experiment provides a case study for predicting the outcome of novel species interactions formed under changing climates.     

The use of floral resources by Visitors on Sparattosperma leucanthum (Vell.) K. Schum. (Bignoniaceae)

Aiming to estimate the rate of exploitation of the floral resources of Sparattosperma leucanthum (Vell.) K. Schum. as well as the interaction with their floral visitors in the pollination, the number of visits by flower was sampled, according to the type of visitation, the collected resource and the visitor's behavior during the forage for flowers. The floral visitors were grouped into seven guilds, organized in decreasing order of benefit to the S. leucanthum flower's pollination: effective pollinator, occasional pollinator, endogamic pollinator, generalist visitor, thievery visitor, thievery-pillager ant and pillager visitor. The total of 48.2 +/- 8.84 visits were recorded by flower. Nearly 50% of the visits resulted in nectar thief or pillage, which posed some problems to the reproduction of S. leucanthum, such as the drop in the attractiveness to pollinators and the harm to the flower's reproductive tissues. Trigona spinipes (Fabr.) (Hymenoptera: Apidae) was considered the most harmful species owing to the high frequency of pillage and forage. Bombus sp1, however, was probably the species that pollinated S. lecanthum flowers the most, making use of the crossed pollination.     

Drought and leaf herbivory influence floral volatiles and pollinator attraction

The effects of climate change on species interactions are poorly understood. Investigating the mechanisms by which species interactions may shift under altered environmental conditions will help form a more predictive understanding of such shifts. In particular, components of climate change have the potential to strongly influence floral volatile organic compounds (VOCs) and, in turn, plant–pollinator interactions. In this study, we experimentally manipulated drought and herbivory for four forb species to determine effects of these treatments and their interactions on (1) visual plant traits traditionally associated with pollinator attraction, (2) floral VOCs, and (3) the visitation rates and community composition of pollinators. For all forbs tested, experimental drought universally reduced flower size and floral display, but there were species‐specific effects of drought on volatile emissions per flower, the composition of compounds produced, and subsequent pollinator visitation rates. Moreover, the community of pollinating visitors was influenced by drought across forb species (i.e. some pollinator species were deterred by drought while others were attracted). Together, these results indicate that VOCs may provide more nuanced information to potential floral visitors and may be relatively more important than visual traits for pollinator attraction, particularly under shifting environmental conditions.     

Scale-dependent shifts in the species composition of flower visitors with changing floral density

Responses of flower-visiting animals to floral density can alter interactions between plants, influencing a variety of biological processes, including plant population dynamics and the evolution of flowering phenology. Many studies have found effects of floral or plant density on pollinator visitation rates at patch scales, but little is known about responses of flower visitors to floral densities at larger scales. Here, I present data from an observational field study in which I measured the effects of floral density on visitation to the annual composite Holocarpha virgata at both patch (4 m²) and site (12.6 ha) spatial scales. The species composition of flower visitors changed with floral density, and did so in different ways at the two scales. At the site scale, average floral density within patches of H. virgata or within patches of all summer-flowering species combined had a significant positive effect on per-flowerhead visitation by the long-horned bee Melissodes lupina and no significant effects on visitation by any other taxa. At the patch scale, per-flowerhead visitation by honeybees significantly increased whereas visitation by M. lupina often decreased with increasing floral density. For both species, responses to patch-scale floral density were strongest when site-scale floral density was high. The scale-dependence of flower visitor responses to floral density and the interactions between site- and patch-scale effects of floral density observed in this study underscore the importance of improving our understanding of pollinators’ responses to floral density at population scales.     

Initial floral visitor identity and foraging time strongly influence blueberry reproductive success

Priority effects occur when the order of species arrival affects subsequent ecological processes. The order that pollinator species visit flowers may affect pollination through a priority effect, whereby the first visitor reduces or modifies the contribution of subsequent visits. We observed floral visitation to blueberry flowers from honeybees, stingless bees or a mixture of both species and investigated how (i) initial visits differed in duration to later visits; and (ii) how visit sequences from different pollinator taxa influenced fruit weight. Stingless bees visited blueberry flowers for significantly longer than honeybees and maintained their floral visit duration, irrespective of the number of preceding visits. In contrast, honeybee visit duration declined significantly with an increasing number of preceding visits. Fruit weight was positively associated with longer floral visit duration by honeybees but not from stingless bee or mixed species visitation. Fruit from mixed species visits were heavier overall than single species visits, because of a strong priority effect. An initial visit by a stingless bee fully pollinated the flower, limiting the pollination contribution of future visitors. However, after an initial honeybee visit, flowers were not fully pollinated and additional visitation had an additive effect upon fruit weight. Blueberries from flowers visited first by stingless bees were 60% heavier than those visited first by honeybees when total floral visitation was short (∼1 min). However, when total visitation time was long (∼ 8 min), blueberry fruit were 24% heavier when initial visits were from honeybees. Our findings highlight that the initial floral visit can have a disproportionate effect on pollination outcomes. Considering priority effects alongside traditional measures of pollinator effectiveness will provide a greater mechanistic understanding of how pollinator communities influence plant reproductive success.     

Floral divergence, pollinator partitioning and the spatiotemporal pattern of plant–pollinator interactions in three sympatric Adenophora species

Floral divergence among congeners may relate to differential utilization of pollinators and contribute to reducing overlap in pollination niches. To investigate whether and how floral differences are associated with differential utilization of pollinators in three sympatric Adenophora species, we analyzed floral traits and evaluated the contribution of different visitors to pollination. We compared visitation rates of different pollinator categories in different years and sites. A suite of floral traits differed among the three Adenophora species, suggesting adaptation to diurnal versus nocturnal pollination and an intermediate condition. However, many visitor species were shared among the three plant species, suggesting that floral traits did not rigorously filter visitors. Effective pollinators were large bees and moths. The importance of large bees as pollinators decreased whereas that of moths increased along the gradient from typically bee-pollinated to moth-pollinated flowers. The intermediate species (A. khasiana) differed substantially from the other two species in pollinator species but not in pollinator categories. The principal pollinator category of each species was constant across years and sites except in the intermediate species where it differed between two sites. Overall, the three sympatric species of Adenophora partition pollinators by floral divergence and the principal pollinators coincide with the predictions based on floral syndromes.     

Facilitative interactions among co-flowering <Emphasis Type="Italic">Primula</Emphasis> species mediated by pollinator sharing

We investigated flowering phenology, pollinator visitation and visitor community composition in communities of self-incompatible sympatric Primula species in a high-elevation Himalayan ecosystem. Within the tight constraints imposed by short growing seasons in such ecosystems, interactions among co-occurring plants for pollinators may vary from competition to facilitation, depending on the specifics of the system. We found that pollinator community composition changed with elevation in this system: lepidopterans were the dominant visitors at lower elevations (2200–3000 masl), bees (other than bumblebees) dominated at mid-elevations (3000–3800 masl) and bumblebees dominated at higher elevations (3800–4600 masl). However, within an elevation zone, there were no significant differences in pollinators amongst co-occurring Primula species. At a focal study site where multiple Primula species co-occurred, our results showed that even while the overall flowering periods of these species broadly overlapped, the peak flowering periods of different Primula species were temporally segregated. Upon further inferring the nature of interaction amongst co-flowering Primula species, we found that plots with higher Primula diversity (≥?2 species) and density (80–100 individuals) experienced significantly higher pollinator visitation, compared with plots with single species and low flower densities (40–50 individuals). Our results suggest that in this community of sympatric, self-incompatible Primula species, a broadly aggregated, synchronous floral display of multiple species results in pollinator facilitation by attracting a greater number of pollinator visitors. Within this broadly synchronous display, the temporal segregation of peak flowering period of individual species may reduce competition for pollinators and limit heterospecific pollen transfer.     

The invasive <Emphasis Type="Italic">Lespedeza cuneata</Emphasis> attracts more insect pollinators than native congeners in tallgrass prairie with variable impacts

Invasive plant species can potentially exert competitive or facilitative effects on insect pollination services of native species. Factors that influence these effects include the degree of shared pollinator species, synchronous flowering phenology, similar flower morphology and color, relatedness of invasive and natives, and showiness and densities of flowers. We investigated such plant-pollinator dynamics by comparing the invasive Lespedeza cuneata and three native congeners, all sympatric with synchronous flowering, using in situ populations over 2 years during peak floral displays. Insect visitation rates of the invasive were significantly higher per plant in both years than on the native species. The invasive exerted a competitive effect on visitation of the two native species with fewer shared pollinators, and a facilitative effect on visitation of the native species with the highest degree of shared insect visitors. Positive correlations were found between floral density and visitation rate per plant in all the native species. Although no such correlation was found for the invasive, floral density in L. cuneata was at least twenty times higher than in the native species and likely saturated the response of the pollinator community. Analyses of insect visitor taxonomic data indicated the insect communities visiting each of the Lespedeza species were generally similar though with species-specific differences. The main exception was that the common honeybee, Apis mellifera, was a primary visitor to the invasive plant species, yet was never observed on the native Lespedeza species.     

Pollinator community structure and sources of spatial variation in plant–pollinator interactions in <Emphasis Type="Italic">Clarkia xantiana</Emphasis> ssp. <Emphasis Type="Italic">xantiana</Emphasis>

The structure of diverse floral visitor assemblages and the nature of spatial variation in plant–pollinator interactions have important consequences for floral evolution and reproductive interactions among pollinator-sharing plant species. In this study, I use surveys of floral visitor communities across the geographic range of Clarkia xantiana ssp. xantiana (hereafter C. x. xantiana) (Onagraceae) to examine the structure of visitor communities, the specificity of the pollination system, and the role of variation in the abiotic vs. biotic environment in contributing to spatial variation in pollinator abundance and community composition. Although the assemblage of bee visitors to C. x. xantiana is very diverse (49 species), few were regular visitors and likely to act as pollinators. Seventy-four percent of visitor species accounted for only 11% of total visitor abundance and 69% were collected in three or fewer plant populations (of ten). Of the few reliable visitors, Clarkia pollen specialist bees were the most frequent visitors, carried more Clarkia pollen compared to generalist foragers, and were less likely to harbor foreign pollen. Overall, the core group of pollinators was obscured by high numbers of incidental visitors that are unlikely to contribute to pollination. In a geographic context, the composition of specialist pollinator assemblages varied considerably along the abiotic gradient spanning the subspecies range. However, the overall abundance of specialist pollinators in plant populations was not influenced by the broad-scale abiotic gradient but strongly affected by local plant community associations. C. x. xantiana populations sympatric with pollinator-sharing congeners were visited twice as often by specialists compared to populations occurring alone. These positive indirect interactions among plant species may promote population persistence and species coexistence by enhancing individual reproductive success.Electronic Supplementary Material Supplementary material is available for this article at     

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.     

Diversity of floral visitors to sympatric Lithophragma species differing in floral morphology

Most coevolving relationships between pairs of species are embedded in a broader multispecific interaction network. The mutualistic interaction between Lithophragma parviflorum (Saxifragaceae) and its pollinating floral parasite Greya politella (Lepidoptera, Prodoxidae) occurs in some communities as a pairwise set apart from most other interactions in those communities. In other communities, however, this pair of species occurs with congeners and with other floral visitors to Lithophragma. We analyzed local and geographic differences in the network formed by interactions between Lithophragma plants and Greya moths in communities containing two Lithophragma species, two Greya species, and floral visitors other than Greya that visit Lithophragma flowers. Our goal was to evaluate if non-Greya visitors were common, if visitor assembly differs between Lithophragma species and populations and if these visitors act as effective pollinators. Sympatric populations of L. heterophyllum and L. parviflorum differ in floral traits that may affect assemblies of floral visitors. Visitation rates by non-Greya floral visitors were low, and the asymptotic number of visitor species was less than 20 species in all populations. Lithophragma species shared some of the visitors, with visitor assemblages differing between sites more for L. heterophyllum than for L. parviflorum. Pollination efficacy experiments showed that most visitors were poor pollinators. Single visits to flowers by this assemblage of species resulted in significantly higher seed set in Lithophragma heterophyllum (30.6 ± 3.9 SE) than in L. parviflorum (4.7 ± 3.4 SE). This difference was consistent between sites, suggesting that these visitors provide a better fit to the floral morphology of L. heterophyllum. Overall, none of the non-Greya visitors appears to be either sufficiently common or efficient as a pollinator to impose strong selection on any of these four Lithophragma populations in comparison with Greya, which occurs within almost all populations of these species throughout their geographic ranges.     

Diverse visitors with various pollinator importance and temporal change in the important pollinators of <Emphasis Type="Italic">Geranium thunbergii</Emphasis> (Geraniaceae)

Assessing pollinator importance of each floral visitor to a plant species is a key to understanding plant–pollinator interaction. The present study examined visitation frequency, pollination efficiency, and pollinator importance of the full range of floral visitors to Geranium thunbergii natural population, by measuring seed-set. During 2 years of observations, the flowers were visited by at least 45 insect species belonging to four orders. Among the main 22 visitor species, 11 species belonging to three orders (Hymenoptera, Diptera, and Lepidoptera) acted as the efficient pollinators. In both years, Hymenoptera, especially bees, was the most important pollinator to G.thunbergii. Thus, the flowers could be considered as bee-pollinated. However, the most important species were not constant between years. The study also documented that the efficient pollinators have larger body sizes. The dish-shaped floral morphology, taxonomically diverse pollinators, and temporal change in the most important pollinators indicate that G.thunbergii–pollinator interaction is a rather generalized system. The results suggest that casual observations of visitation, or even precise measurement of pollinator importance in a single season is insufficient to identify important pollinators.     

Breakdown of species boundaries in Mandevilla: floral morphological intermediacy,novel fragrances and asymmetric pollen flow

• Phenotypic intermediacy is an indicator of putative hybrid origin and has provided the main clues to discovering hybrid plants in nature. Mandevilla pentlandiana and M. laxa (Apocynaceae) are sister species with clear differences in floral phenotype and associated pollinator guilds: diurnal Hymenoptera and nocturnal hawkmoths, respectively. The presence of individuals with intermediate phenotypes in a wild population raises questions about the roles of visual and olfactory signals (i.e. corolla morphology and floral fragrances) as barriers to interbreeding, and how the breakdown of floral isolation occurs.
• We examined phenotypic variation in a mixed Mandevilla population, analysing the chemical composition of floral fragrances, characterising floral shape through geometric morphometrics and assessing individual grouping through taxonomically relevant traits and an unsupervised learning algorithm. We quantified the visitation frequencies of floral visitors and tracked their foraging movements using pollen analogues.
• The presence of morphologically intermediate individuals and pollen analogue movement suggested extensive hybridisation between M. laxa and M. pentlandiana, along with asymmetrical rates of backcrossing between these putative hybrids and M. laxa. Floral volatiles from putative hybrid individuals showed a transgressive phenotype, with additional compounds not emitted by either parental species.
• Our results suggest the presence of a hybrid swarm between sympatric M. pentlandiana and M. laxa and indicate that initial hybridisation events between these parental species are rare, but once they occur, visits between putative hybrids and M. laxa are common and facilitate continued introgression.

     

Comparison of pollen transfer dynamics by multiple floral visitors: experiments with pollen and fluorescent dye

BACKGROUND AND AIMS: Most plant species are visited by a diversity of floral visitors. Pollen transfer of the four most common pollinating bee species and one nectar-robbing bee of the distylous plant Gelsemium sempervirens were compared. METHODS: Naturally occurring pollen loads carried by the common floral visitor species of G. sempervirens were compared. In addition, dyed pollen donor flowers and sequences of four emasculated recipient flowers in field cages were used to estimate pollen transfer, and the utility of fluorescent dye powder as an analogue for pollen transfer was determined. KEY RESULTS: Xylocopa virginica, Osmia lignaria and Habropoda laboriosa carried the most G. sempervirens pollen on their bodies, followed by Bombus bimaculatus and Apis mellifera. However, B. bimaculatus, O. lignaria and H. laboriosa transferred significantly more pollen than A. mellifera. Nectar-robbing X. virginica transferred the least pollen, even when visiting legitimately. Dye particles were strongly correlated with pollen grains on a stigma, and therefore provide a good analogue for pollen in this system. The ratio of pollen : dye across stigmas was not affected by bee species or interactions between bee species and floral morphology. However, dye transfer was more sensitive than pollen transfer to differences in floral morphology. CONCLUSIONS: The results from this study add to a growing body of literature highlighting that floral visitors vary in pollination effectiveness, and that visitors carrying the most pollen on their bodies may not always be the most efficient at depositing pollen on stigmas. Understanding the magnitude of variability in pollinator quality is one important factor for predicting how different pollinator taxa may influence the evolution of floral traits.     

Genetics of floral traits influencing reproductive isolation between Aquilegia formosa and Aquilegia pubescens

Abstract: Reproductive isolation between Aquilegia formosa and Aquilegia pubescens is influenced by differences in their flowers through their effects on pollinator visitation and pollen transfer. Here, we investigate the genetic basis of floral characters differentiating these species. We found that in addition to the effects of flower orientation and the length of nectar spurs previously described, other characters such as flower color or odor affect hawkmoth visitation. Repeatability of measurements in an F2 population ranged from 0.53 to 0.83 among five floral traits, indicating that using the means of multiple measures per plant will substantially increase the power of a quantitative trait locus (QTL) analysis. Integration of floral traits was indicated by significant correlations among traits in an F2 population. In a separate F2 population we found that QTL for different floral traits were often closely associated, indicating that linkage or pleiotropy cause at least some of this integration. In addition, we found QTL for all floral traits examined. Because Aquilegia species are largely interfertile and vary extensively in both floral morphology and ecology, they offer the opportunity for QTL studies of a wide range of characters affecting reproductive isolation.     

Massive floral display affects insect visits but not pollinator‐mediated pollen transfer in Rhododendron ferrugineum

Fragmentation of natural vegetation creates one of the largest threats to plant–pollinator interactions. Although fragmentation impacts on plant populations have been explored in many, mainly herbaceous, species, the response of wild mass‐flowering species is poorly known. Here, we studied 28 heathland patches dominated by the mass‐flowering shrub Rhododendron ferrugineum, each presenting different R. ferrugineum floral display sizes (total inflorescence number per patch) and patch isolation (median distance to the three nearest patches). We assessed the impacts of these two factors on (i) heathland patch visitor assemblage (considering R. ferrugineum versus surrounding community) and (ii) R. ferrugineum flower visitation rate and pollen transfer limitation (comparing seed set from emasculated to pollen‐supplemented flowers). We found that diversity and abundance of bees visiting R. ferrugineum in heathland patches significantly decreased with decreasing R. ferrugineum floral display, while overall visitor density per patch and flower visitation rate increased. Moreover, a decrease in massive floral display and increase in patch isolation resulted in reduced visitor density in the surrounding community. Even in patches with few individuals, we found disproportionate visitor abundance in R. ferrugineum compared to the surrounding community. Finally, pollen transfer limitation in R. ferrugineum was neither affected by visitation rate nor by patch attributes. By disproportionally attracting pollinators from co‐flowering species, and probably promoting geitonogamous pollen transfer, the mass‐flowering trait appears adequate to compensate, in terms of conspecific pollen transfer, for the decrease in visitor diversity and abundance and in mate availability, which usually result from population fragmentation.     

Relationships between the density and diversity of floral resources and flower visitor activity in a temperate grassland community

Abstract 1. Does the diversity and abundance of one trophic level affect another? Several studies at the landscape level have found a positive relationship between the diversity of floral resources and the diversity and abundance of pollinators. However, little is known about the relationship between these trophic levels on a smaller spatial scale, and the importance of blossom density relative to plant species richness in predicting abundance and richness of different flower visitor groups. 2. This study used a small‐scale approach to investigate how, and if, the diversity and abundance of floral resources in study plots affected the visitation activity of different flower visitor groups. During 201 observation periods between late May and mid‐August 2003, 3682 visits were observed. Bumblebees (60%), muscoids (17%), syrphids (9%), and beetles (5%) were the most abundant flower visitors. 3. Regression analysis was used to investigate the relationship between blossom density and plant species richness with visitation activity, including the probability of presence in plots, the visits within plots, and the visitor richness of the most abundant pollinator groups. 4. The activity of beetles, bumblebees, and muscoids was positively predicted by the variation in blossom density, while syrphid activity was better predicted by plant species richness. Overall, the models for beetles and bumblebees explained much more of the variation in activity compared with the models for the dipterans, and blossom density was a better predictor of both flower visitor richness and activity than was plant species richness.