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.     

Resource complementation and the response of an insect herbivore to habitat area and fragmentation

Few studies have disentangled the effects of the area and fragmentation of a focal habitat type on species that use multiple habitat types within a landscape. We experimentally investigated the effects of habitat area, habitat fragmentation, and matrix composition on the movement and distribution of Melanoplus femurrubrum. Adults of this grasshopper feed preferentially on grasses, but oviposit almost exclusively in soil dominated by forbs. We compared population densities among plots that were made to vary in the area and fragmentation of clover habitat and composition of the matrix (grass or bare ground) within which clover habitat was embedded. In addition, a mark-recapture survey was conducted to examine effects of habitat area, fragmentation, and matrix composition on loss of individuals from a plot’s clover habitat and movement between clover subplots within plots. Overall densities of adult M. femurrubrum (averaged over clover and matrix) were 2.2× higher in plots where the matrix was composed of grass as compared to bare ground, and 1.8× higher in plots with 64 compared to 16 m² of clover habitat. Overall densities of nymphs were also positively influenced by greater clover area, but were unaffected by matrix composition. Within focal clover habitat embedded in grass matrix, adult densities were 2.1× higher in small clover subplots than large clover subplots. We conclude that the grass matrix had a positive effect on adult densities, but not nymph densities, because grass and forb-dominated habitats likely provide complementary resources only for adults. The aggregation of adults on small clover subplots within grass matrix was mainly attributed to a greater rate of emigration loss per unit area. In addition, this study emphasizes that a species’ response to changes in the area of a focal habitat type can depend significantly on the availability of complementary resources in the surrounding landscape. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Pollinator community responses to the spatial population structure of wild plants: A pan-European approach

Land-use changes can alter the spatial population structure of plant species, which may in turn affect the attractiveness of flower aggregations to different groups of pollinators at different spatial scales. To assess how pollinators respond to spatial heterogeneity of plant distributions and whether honeybees affect visitation by other pollinators we used an extensive data set comprising ten plant species and their flower visitors from five European countries. In particular we tested the hypothesis that the composition of the flower visitor community in terms of visitation frequencies by different pollinator groups were affected by the spatial plant population structure, viz. area and density measures, at a within-population (‘patch’) and among-population (‘population’) scale. We found that patch area and population density were the spatial variables that best explained the variation in visitation frequencies within the pollinator community. Honeybees had higher visitation frequencies in larger patches, while bumblebees and hoverflies had higher visitation frequencies in sparser populations. Solitary bees had higher visitation frequencies in sparser populations and smaller patches. We also tested the hypothesis that honeybees affect the composition of the pollinator community by altering the visitation frequencies of other groups of pollinators. There was a positive relationship between visitation frequencies of honeybees and bumblebees, while the relationship with hoverflies and solitary bees varied (positive, negative and no relationship) depending on the plant species under study. The overall conclusion is that the spatial structure of plant populations affects different groups of pollinators in contrasting ways at both the local (‘patch’) and the larger (‘population’) scales and, that honeybees affect the flower visitation by other pollinator groups in various ways, depending on the plant species under study. These contrasting responses emphasize the need to investigate the entire pollinator community when the effects of landscape change on plant–pollinator interactions are studied.     

Spatial variability in a plant–pollinator community across a continuous habitat: high heterogeneity in the face of apparent uniformity

Large‐scale spatial variability in plant–pollinator communities (e.g. along geographic gradients, across different landscapes) is relatively well understood. However, we know much less about how these communities vary at small scales within a uniform landscape. Plants are sessile and highly sensitive to microhabitat conditions, whereas pollinators are highly mobile and, for the most part, display generalist feeding habits. Therefore, we expect plants to show greater spatial variability than pollinators. We analysed the spatial heterogeneity of a community of flowering plants and their pollinators in 40 plots across a 40‐km² area within an uninterrupted Mediterranean scrubland. We recorded 3577 pollinator visits to 49 plant species. The pollinator community (170 species) was strongly dominated by honey bees (71.8% of the visits recorded). Flower and pollinator communities showed similar beta‐diversity, indicating that spatial variability was similar in the two groups. We used path analysis to establish the direct and indirect effects of flower community distribution and honey bee visitation rate (a measure of the use of floral resources by this species) on the spatial distribution of the pollinator community. Wild pollinator abundance was positively related to flower abundance. Wild pollinator visitation rate was negatively related to flower abundance, suggesting that floral resources were not limiting. Pollinator and flower richness were positively related. Pollinator species composition was weakly related to flower species composition, reflecting the generalist nature of flower–pollinator interactions and the opportunistic nature of pollinator flower choices. Honey bee visitation rate did not affect the distribution of the wild pollinator community. Overall, we show that, in spite of the apparent physiognomic uniformity, both flowers and pollinators display high levels of heterogeneity, resulting in a mosaic of idiosyncratic local communities. Our results provide a measure of the background of intrinsic heterogeneity within a uniform habitat, with potential consequences on low‐scale ecosystem function and microevolutionary patterns.     

Direct and indirect effects of episodic frost on plant growth and reproduction in subalpine wildflowers

Frost is an important episodic event that damages plant tissues through the formation of ice crystals at or below freezing temperatures. In montane regions, where climate change is expected to cause earlier snow melt but may not change the last frost‐free day of the year, plants that bud earlier might be directly impacted by frost through damage to flower buds and reproductive structures. However, the indirect effects of frost mediated through changes in plant–pollinator interactions have rarely been explored. We examined the direct and pollinator‐mediated indirect effects of frost on three wildflower species in southwestern Colorado, USA, Delphinium barbeyi (Ranunculaceae), Erigeron speciosus (Asteraceae), and Polemonium foliosissimum (Polemoniaceae), by simulating moderate (?1 to ?5°C) frost events in early spring in plants in situ. Subsequently, we measured plant growth, and upon flowering measured flower morphology and phenology. Throughout the flowering season, we monitored pollinator visitation and collected seeds to measure plant reproduction. We found that frost had species‐specific direct and indirect effects. Frost had direct effects on two of the three species. Frost significantly reduced flower size, total flowers produced, and seed production of Erigeron. Furthermore, frost reduced aboveground plant survival and seed production for Polemonium. However, we found no direct effects of frost on Delphinium. When we considered the indirect impacts of frost mediated through changes in pollinator visitation, one species, Erigeron, incurred indirect, negative effects of frost on plant reproduction through changes in floral traits and pollinator visitation, along with direct effects. Overall, we found that flowering plants exhibited species‐specific direct and pollinator‐mediated indirect responses to frost, thus suggesting that frost may play an important role in affecting plant communities under climate change.     

The presence of co-flowering species facilitates reproductive success of Pedicularis monbeigiana (Orobanchaceae) through variation in bumble-bee foraging behaviour

Background and Aims The presence of co-flowering species can alter pollinator foraging behaviour and, in turn, positively or negatively affect the reproductive success of the focal species. Such interactions were investigated between a focal species, Pedicularis monbeigiana, and a co-flowering species, Vicia dichroantha, which was mediated by behaviour alteration of the shared bumble-bee pollinator. Methods Floral display size and floral colour change of P. monbeigiana were compared between pure (P. monbeigiana only) and mixed (P. monbeigiana and V. dichroantha) plots in two populations. Pollinator visitation rates, interspecific floral switching and successive within-plant pollinator visits were recorded. In addition, supplemental pollination at plant level was performed, and the fruit set and seed set were analysed in pure and mixed plots with different densities of P. monbeigiana. Key Results Pollinator visitation rates were dramatically higher in mixed plots than in pure plots. The higher pollinator visitation rates were recorded in both low- and high-density plots. In particular, successive flower visits within an individual plant were significantly lower in mixed plots. Supplemental pollination significantly increased fruit set and seed set of individuals in pure plots, while it only marginally increased seed set per fruit of plants in mixed plots. Conclusions The presence of V. dichroantha can facilitate pollination and increase female reproductive success of P. monbeigiana via both quantity (mitigating pollinator limitation) and quality (reducing geitonogamy) effects. This study suggests that successive pollinator movements among flowers within a plant, as well as pollinator visitation rates and interspecific flower switching, may be important determinants of the direction and mechanisms of interaction between species.     

The town Crepis and the country Crepis: How does fragmentation affect a plant–pollinator interaction?

In fragmented habitats, one cause of the decrease of plant diversity and abundance is the disruption of plant–animal interactions, and in particular plant–pollinator interactions. Since habitat fragmentation acts both on pollinator behaviour and plant reproduction, its consequences for the stability of such interactions are complex. An extreme case of habitat fragmentation occurs in urbanised areas where suitable habitat (in the present study small patches around ornamental trees) is embedded in a highly unsuitable environment (concrete matrix). Based on simple experiments, we ask whether pollinators can adapt their foraging behaviour in response to the amount of available resources (flowers) in the fragments and their isolation, as predicted by the optimal foraging theory. To do so we analysed the effect of fragmentation on the behaviour of pollinators visiting Crepis sancta (L.) Bornm. (Asteraceae), which forms large populations in the countryside and patchy populations in urban environments. More precisely we studied pollinator visitation rates, capitulum visit durations, capitulum search durations and capitulum size choice. Pollinators chose larger capitula in both types of populations and their foraging behaviour differed between the two population types in three ways: (1) pollinator visits were lower in urban fragmented populations, perhaps due to the lower accessibility of urban patches; (2) capitulum visit durations were longer in urban fragmented populations, a possible compensation of energy lost during flights among patches; and (3) capitulum search durations where longer in urban fragmented populations, which may represent an increase in capitulum prospecting effort. We discuss the possible impacts of such differences for plant population functioning in the two types of populations.     

Effects of flowering plant density on pollinator visitation, pollen receipt, and seed production in Delphinium barbeyi (Ranunculaceae)

Variation in flowering plant density can have conflicting effects on pollination and seed production. Dense flower patches may attract more pollinators, but flowers in those patches may also compete for pollinator visits and abiotic resources. We examined how natural and experimental conspecific flowering plant density affected pollen receipt and seed production in a protandrous, bumble bee-pollinated wildflower, Delphinium barbeyi (Ranunculaceae). We also compared floral sex ratios, pollinator visitation rates, and pollen limitation of seed set from early to late in the season to determine whether these factors mirrored seasonal changes in pollen receipt and seed production. Pollen receipt increased with natural flowering plant density, while seed production increased across lower densities and decreased across higher flower densities. Experimental manipulation of flowering plant density did not affect pollinator visitation rate, pollen receipt, or seed production. Although pollinator visitation rate increased 10-fold from early to late in the season, pollen receipt and seed set decreased over the season. Seed set was never pollen-limited. Thus, despite widespread effects of flowering plant density on plant reproduction in other species, the effects of conspecific flowering plant density on D. barbeyi pollination and seed production are minor.     

Contrasting effects of plant population size on florivory and pollination

Changes in plant population size, induced by various forms of habitat degradation, can affect the performance of plants by altering their interactions with other organisms such as pollinators and herbivores. However, studies on plant reproductive response to variation in population size that simultaneously consider different interactions are rare. In this study, we examined (1) how levels of pollinator visitation and florivory vary with population size of a self-incompatible herb, Verbascum nigrum (Scrophulariaceae), (2) the relative effect of these two interactions on host seed set, and (3) whether the intensity of florivory influences pollinator visitation rate. The intensity of florivory increased, whereas pollinator visitation rate decreased with host population size. Although average seed production was negatively affected by the intensity of florivory, seed production was independent of population size. The direct negative effect of florivores on plant seed set was indirectly enforced by their negative effect on pollinator visitation rate. Our results emphasize the complexity of responses of different plant–animal interactions to plant population size. However, interactions involving specialized organisms are likely to disrupt first as plant population size decreases.     

Effects of an exotic plant and habitat disturbance on pollinator visitation and reproduction in a boreal forest herb

The invasion of exotic species into natural habitats is considered to be a major threat to biodiversity, and many studies have examined how exotic plants directly affect native plant species through competitive interactions for abiotic resources. However, although exotics can have potentially great ecological and evolutionary consequences, very few researchers have studied the effect of exotics on the interactions between plants and their mutualistic partners, such as pollinators, and none have reported on such impacts in logged and undisturbed boreal forest ecosystems. Here we show how experimental introductions of an exotic plant species (Phacelia tanacetifolia Bentham) affect pollinator visitation and female reproductive success of a native plant (Melampyrum pratense L.) in recently disturbed (i.e., logged) and in undisturbed boreal forest habitats. The presence of Phacelia significantly increased the number of bumble bees entering plots in both habitat types. However, the exotic species had a strong negative impact on the visitation rate to the native species in both habitat types. Despite this negative impact on pollinator visitation, the exotic had no effect on female reproductive success of the native species in any habitat. Our results show that seed production may be more robust than pollinator visitation to exotic invasion, irrespective of habitat disturbance history.     

Precipitation and predation risk alter the diversity and behavior of pollinators and reduce plant fitness

Biotic and abiotic factors may individually or interactively disrupt plant–pollinator interactions, influencing plant fitness. Although variations in temperature and precipitation are expected to modify the overall impact of predators on plant–pollinator interactions, few empirical studies have assessed if these weather conditions influence anti-predator behaviors and how this context-dependent response may cascade down to plant fitness. To answer this question, we manipulated predation risk (using artificial spiders) in different years to investigate how natural variation in temperature and precipitation may affect diversity (richness and composition) and behavioral (visitation) responses of flower-visiting insects to predation risk, and how these effects influence plant fitness. Our findings indicate that predation risk and an increase in precipitation independently reduced plant fitness (i.e., seed set) by decreasing flower visitation. Predation risk reduced pollinator visitation and richness, and altered species composition of pollinators. Additionally, an increase in precipitation was associated with lower flower visitation and pollinator richness but did not alter pollinator species composition. However, maximum daily temperature did not affect any component of the pollinator assemblage or plant fitness. Our results indicate that biotic and abiotic drivers have different impacts on pollinator behavior and diversity with consequences for plant fitness components. Even small variation in precipitation conditions promotes complex and substantial cascading effects on plants by affecting both pollinator communities and the outcome of plant–pollinator interactions. Tropical communities are expected to be highly susceptible to climatic changes, and these changes may have drastic consequences for biotic interactions in the tropics.     

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.     

Insect herbivory in an experimental agroecosystem: the relative importance of habitat area, fragmentation, and the matrix

Habitat area, fragmentation, and the surrounding matrix influence levels of herbivory in various ecosystems, but the relative importance of these effects has rarely been assessed. We compared levels of herbivory and densities of dominant arthropod herbivores (the hemipteran insects Agallia constricta, Empoasca fabae, Therioaphis trifolii, Lygus lineolaris and Halticus bractatus ) among experimental plots that varied in the area and fragmentation of clover habitat and the composition of the matrix (bare ground or grass) surrounding clover habitat. To assess levels of herbivory, we compared clover biomass within herbivore exclosures to the biomass accessible to herbivores. The area and fragmentation of clover habitat, as well as matrix composition, significantly influenced the collective densities of herbivores, although each species exhibited unique responses to habitat structure. Herbivory was strongest in plots with large (64  m²) as compared to small (16  m²) amounts of clover habitat. The difference in clover biomass between the inside and outside of exclosures increased significantly with increasing density of Empoasca fabae but was unrelated to the densities of the other herbivores, suggesting that Empoasca fabae was an exceptionally important herbivore in this system. This study supports the view that herbivore densities and herbivory generally increase with increasing area of plant monocultures, but emphasizes that levels of herbivory may be driven primarily by one or a few key herbivore species.     

Does human-induced habitat transformation modify pollinator-mediated selection? A case study in Viola portalesia (Violaceae)

Pollinator-mediated selection is one of the most important factors driving adaptation in flowering plants. However, as ecological conditions change through habitat loss and fragmentation, the interactions among species may evolve in new and unexpected directions. Human-induced environmental variation is likely to affect selection regimes, but as yet no empirical examples have been reported. In the study reported here, we examined the influence of human-induced habitat transformation on the composition of pollinator assemblages and, hence, pollinator-mediated selection on the flower phenotype of Viola portalesia (Violaceae). Our results indicate that pollinator assemblages differed substantially in terms of species composition and visitation rate between nearby native and transformed habitats. Similarly, the insect species that contributed most to visitation rates differed between plant populations. While the magnitude and sign of pollinator-mediated selection on flower length and width did not differ between sites, selection for flower number lost significance in the transformed habitat, and a significant pattern of disruptive selection for flower shape, undetected in the native habitat, was present in the transformed one. Overall, the results of this study suggest that human-induced habitat change may not only modify the species composition of pollinator assemblages, relaxing the selection process on some flower characters, but they may also create new opportunities for fitness-trait covariation not present in pristine conditions.     

Direct versus indirect effects of habitat fragmentation on community patterns in experimental landscapes

Habitat area and fragmentation are confounded in many ecological studies investigating fragmentation effects. We thus devised an innovative experiment founded on fractal neutral landscape models to disentangle the relative effects of habitat area and fragmentation on arthropod community patterns in red clover (Trifolium pratense). The conventional approach in experimental fragmentation studies is to adjust patch size and isolation to create different landscape patterns. We instead use fractal distributions to adjust the overall amount and fragmentation of habitat independently at the scale of the entire landscape, producing different patch properties. Although habitat area ultimately had a greater effect on arthropod abundance and diversity in this system, we found that fragmentation had a significant effect in clover landscapes with ≤40 % habitat. Landscapes at these lower habitat levels were dominated by edge cells, which had fewer arthropods and lower richness than interior cells. Fragmentation per se did not have a direct effect on local-scale diversity, however, as demonstrated by the lack of a broader landscape effect (in terms of total habitat area and fragmentation) on arthropods within habitat cells. Fragmentation—through the creation of edge habitat—thus had a strong indirect effect on morphospecies richness and abundance at the local scale. Although it has been suggested that fragmentation should be important at low habitat levels (≤20–30 %), we show that fragmentation per se is significant only at intermediate (40 %) levels of habitat, where edge effects were neither too great (as at lower levels of habitat) nor too weak (as at higher levels of habitat).     

The reproductive assurance benefit of selfing: importance of flower size and population size

Autonomous selfing can provide reproductive assurance (RA) for flowering plants that are unattractive to pollinators or in environments that are pollen limited. Pollen limitation may result from the breakdown of once-continuous habitat into smaller, more isolated patches (habitat fragmentation) if fragmentation negatively impacts pollinator populations. Here we quantify the levels of pollen limitation and RA among large and small populations of Collinsia parviflora, a wildflower with inter-population variation in flower size. We found that none of the populations were pollen limited, as pollen-supplemented and intact flowers did not differ in seed production. There was a significant effect of flower size on RA; intact flowers (can self) produced significantly more seeds than emasculated flowers (require pollen delivery) in small-flowered plants but not large-flowered plants. Population size nested within flower size did not significantly affect RA, but there was a large difference between our two replicate populations for large-flowered, small populations and small-flowered, large populations that appears related to a more variable pollination environment under these conditions. In fact, levels of RA were strongly negatively correlated with rates of pollinator visitation, whereby infrequent visitation by pollinators yielded high levels of RA via autonomous selfing, but there was no benefit of autonomous selfing when visitation rates were high. These results suggest that autonomous selfing may be adaptive in fragmented habitats or other ecological circumstances that affect pollinator visitation rates.     

Delayed response in a plant–pollinator system to experimental grassland fragmentation

The fragmentation of natural habitat is considered to be a major threat to biodiversity. Decreasing habitat quality and quantity caused by fragmentation may lead to a disruption of plant–pollinator interactions and to a reduction in sexual reproduction in plant species. We conducted a 6-year field experiment to investigate the effects of small-scale fragmentation on plant–pollinator interactions and genetic diversity in the self-compatible Betonica officinalis. We examined the abundance and composition of pollinators, the foraging behaviour of bumblebees and the performance, outcrossing rate and genetic diversity of B. officinalis after 2 and 6 years in experimentally fragmented nutrient-poor, calcareous grassland in the northern Swiss Jura mountains. Fragments of different size (2.25 and 20.25 m²) were isolated by a 5-m-wide strip of frequently mown vegetation. Control plots of corresponding size were situated in adjacent undisturbed grassland. Experimental grassland fragmentation altered the composition of B. officinalis pollinators and reduced their flower visitation rate. Furthermore, the foraging behaviour of bumblebees was changed in the fragments. After 6 years of fragmentation seed weight was higher in fragments than in control plots. However, the densities of B. officinalis rosettes and inflorescences, plant height and inflorescence length were not affected by fragmentation. The outcrossing frequency of B. officinalis growing in fragments was reduced by 15% after 2 years and by 33% after 6 years of experimental fragmentation. This resulted in a significant reduction of the genetic diversity in seedlings emerging in fragments after 6 years. Our study shows that small-scale habitat fragmentation can disturb the interaction between B. officinalis and pollinators resulting in a reduced outcrossing frequency and genetic diversity in plants growing in fragments. However, the response to fragmentation was considerably delayed. This finding strengthens the claim for long-term field experiments with proper replications and controls to assess delayed effects of habitat fragmentation.     

Pollinator conservation at the local scale: flower density,diversity and community structure increase flower visiting insect activity to mixed floral stands

Insect pollinators play a keystone role in terrestrial ecosystems. The parallel declines in plant and pollinator communities emphasizes that plant-pollinator interactions at the community level are highly relevant for biodiversity conservation. Here we examine relationships between plants and flower-visiting insects (anthophiles) at the scale of local floral patches. We conducted a visitation survey during the spring flowering season, a peak time for pollinator activity in the threatened Cape Floristic Region, South Africa. We tested floral density, diversity and composition as predictors of anthophile diversity (measured at the family/family group level) and visitation rates in multispecies stands of flowers. Although different anthophile groups responded differently, generally anthophile visitation rates and diversity were positively affected by floral density, diversity and community structure. Anthophiles were more abundant and diverse in areas with a high density and diversity of flowers. Plant community structure affected both the likelihood of occurrence and activity of anthophiles in the plots. The two mass flowering species examined here, Relhania fruticosa and Salvia chamaeleagnea, were strong determinants of anthophile activity, greatly increasing visitation rates, even though there was, on average, lower floral density and diversity. Our results show that anthophile activity is affected by highly localised, small-scale factors, namely the density and diversity of flowers and community structure. Important among these factors are patches of high quality habitat, high in floral abundance and diversity, both of which should be included in landscape-level plans for pollinator conservation, providing stepping stones for these insects in transformed landscapes.     

Neighborhood‐contingent indirect interactions between native and exotic plants: multiple shared pollinators mediate reproductive success during invasions

Native and exotic plants can influence one another's fecundity through their influence on shared pollinators. Specifically, invasion may alter abundance and composition of local floral resources, affecting pollinator visitation and ultimately causing seedset of natives in more‐invaded and less‐invaded floral neighborhoods to differ. Such pollinator‐mediated effects of exotic plants on natives are common, but native and exotic plants often share multiple pollinators, which may differ in their responses to altered floral neighborhoods. We quantified pollinator‐mediated interactions between three common forbs of western Washington prairies (native Microseris laciniata and Eriophyllum lanatum and European Hypochaeris radicata) in three floral neighborhoods: 1) high native and low exotic floral density, 2) high exotic floral density and low native density, and 3) experimentally manipulated low exotic floral density. Pollinator visitation rates varied by floral neighborhood, plant species identity, and their interaction for all three plant species. Similarly, pollinator functional groups (eusocial bees, solitary bees, and syrphid flies) contributed differing proportions of total visitation to each species depending upon neighborhood context. Consequently, in exotic neighborhoods H. radicata competed with native M. laciniata, reducing seed set, while simultaneously facilitating visitation and seed set for native E. lanatum. Seed set of H. radicata was also highest in exotic neighborhoods (with high densities of conspecifics), raising the possibility of a positive feedback between exotic abundance and success. Our results suggest that the outcome of indirect interactions between native and exotic plants depends on the density and the composition of the floral neighborhood and of the pollinator fauna, and on context‐dependent pollinator foraging.     

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.