Pollinator guilds respond contrastingly at different scales to landscape parameters of land‐use intensity

Land‐use intensification is the main factor for the catastrophic decline of insect pollinators. However, land‐use intensification includes multiple processes that act across various scales and should affect pollinator guilds differently depending on their ecology. We aimed to reveal how two main pollinator guilds, wild bees and hoverflies, respond to different land‐use intensification measures, that is, arable field cover (AFC), landscape heterogeneity (LH), and functional flower composition of local plant communities as a measure of habitat quality. We sampled wild bees and hoverflies on 22 dry grassland sites within a highly intensified landscape (NE Germany) within three campaigns using pan traps. We estimated AFC and LH on consecutive radii (60–3000 m) around the dry grassland sites and estimated the local functional flower composition. Wild bee species richness and abundance was positively affected by LH and negatively by AFC at small scales (140–400 m). In contrast, hoverflies were positively affected by AFC and negatively by LH at larger scales (500–3000 m), where both landscape parameters were negatively correlated to each other. At small spatial scales, though, LH had a positive effect on hoverfly abundance. Functional flower diversity had no positive effect on pollinators, but conspicuous flowers seem to attract abundance of hoverflies. In conclusion, landscape parameters contrarily affect two pollinator guilds at different scales. The correlation of landscape parameters may influence the observed relationships between landscape parameters and pollinators. Hence, effects of land‐use intensification seem to be highly landscape‐specific.     

Electromagnetic radiation of mobile telecommunication antennas affects the abundance and composition of wild pollinators

The exponential increase of mobile telephony has led to a pronounced increase in electromagnetic fields in the environment that may affect pollinator communities and threaten pollination as a key ecosystem service. Previous studies conducted on model species under laboratory conditions have shown negative effects of electromagnetic radiation (EMR) on reproductive success, development, and navigation of insects. However, the potential effects that widespread mobile telecommunication antennas have on wild pollinator communities outside the laboratory microcosm are still unknown. Here we studied the effects of EMR from telecommunication antennas on key wild pollinator groups (wild bees, hoverflies, bee flies, remaining flies, beetles, butterflies, and wasps). We measured EMR at 4 distances (50, 100, 200 and 400 m) from 10 antennas (5 on Limnos Island and 5 on Lesvos Island, eastern Mediterranean, Greece), and correlated EMR values with insect abundance and richness (the latter only for wild bees and hoverflies). All pollinator groups except butterflies were affected by EMR. In both islands, beetle, wasp, and hoverfly abundance decreased with EMR, whereas the abundance of underground-nesting wild bees and bee flies unexpectedly increased with EMR. The effect of EMR on the abundance of remaining flies differed between islands. With respect to species richness, EMR only tended to have a negative effect on hoverflies in Limnos. As EMR affected the abundance of several insect guilds negatively, and changed the composition of wild pollinators in natural habitats, it might also have additional ecological and economic impacts on the maintenance of wild plant diversity, crop production and human welfare.     

Time since establishment drives bee and hoverfly diversity,abundance of crop-pollinating bees and aphidophagous hoverflies in perennial wildflower strips

Wildflower strips (WFS) are amongst the most commonly applied measures to promote pollinators and natural enemies of crop pests in agroecosystems. Their potential to enhance these functionally important insect groups may vary substantially with time since establishment of WFS. However, knowledge on their temporal dynamics remains scarce, hampering recommendations for optimized design and management. We therefore examined temporal dynamics of taxonomic and functional groups of bees and hoverflies in perennial WFS ranging from one to ≥6 years since sowing with a standardized species-rich seed mixture of flowering plants in 18 agricultural landscapes in Switzerland. The abundance of wild bees, honeybees and hoverflies declined after the second year by 89%, 62% and 72%, respectively. Declines in bee abundance and hoverfly species richness were linear and those of aphidophagous hoverflies exponential, while wild bee species richness peaked in the third year. Declines over time generally paralleled decreases in flower abundance (-83%) and flowering species richness (-61%) and an increase in grass cover (+70%) in WFS. Flowering plant species richness showed strong positive relationships with dominant crop-visiting wild bees and aphidophagous hoverflies. Furthermore, dominant crop-visiting wild bees, but not aphidophagous hoverflies, were positively related to the proportion of (semi-)open semi-natural habitat in the surrounding landscape (500 m radius), but negatively with forest. We conclude that the effectiveness of perennial WFS to promote pollinator diversity, crop-pollinating bees and aphidophagous hoverflies through foraging resources decreases after the first two to three years, probably due to a decline of diverse and abundant floral resources. Although older perennial WFS may still provide valuable nesting and overwintering opportunities for pollinators and natural enemies, our findings indicate that regular re-sowing of perennial WFS may be necessary to maintain adequate floral resource provisioning for effective pollinator conservation and promotion of crop pollination and natural pest control services in agricultural landscapes.     

Effects of grazing intensity on pollinator abundance and diversity,and on pollination services

1. Pollinating insects provide important ecosystem services and are influenced by the intensity of grazing. Based on the Intermediate Disturbance Hypothesis (IDH), pollinator diversity is expected to peak at intermediate grazing intensities. However, this hump‐shaped relationship is rarely found. 2. The effect of grazing intensity was tested on flower cover, on the abundance and richness of bees, hoverflies and bee flies, and on pollination services to early‐flowering bee‐pollinated Asphodelus ramosus L. For that, we used data on 11 plant–pollinator phryganic communities from Lesvos Island (Greece) widely differing in grazing intensities. 3. Flower abundance and richness showed hump‐shaped relationships with grazing intensity. Grazing affected the abundance and richness of bees and hoverflies directly and also indirectly, through changes in the flower community. Grazing influenced directly the richness but not the abundance of bee flies. Overall, pollinator abundance and richness showed hump‐shaped relationships with grazing intensity, but variations in strength (hoverfly abundance) and direction (bee community) of the effect appeared along the season. Early in the season, grazing increased bee abundance but decreased richness, resulting in increased pollen limitation in A. ramosus. 4. The effects of grazing on pollinators vary with the intensity of the disturbance, generally supporting the IDH, and the timing of land‐use activities may influence pollination services. Management strategies should include moderate grazing levels to preserve overall diversity in this area, however, the conservation of particular early bee or bee‐pollinated species may benefit from reduced grazing in early spring.     

Bee Species Diversity Enhances Productivity and Stability in a Perennial Crop

Wild bees provide important pollination services to agroecoystems, but the mechanisms which underlie their contribution to ecosystem functioning—and, therefore, their importance in maintaining and enhancing these services—remain unclear. We evaluated several mechanisms through which wild bees contribute to crop productivity, the stability of pollinator visitation, and the efficiency of individual pollinators in a highly bee-pollination dependent plant, highbush blueberry. We surveyed the bee community (through transect sampling and pan trapping) and measured pollination of both open- and singly-visited flowers. We found that the abundance of managed honey bees, Apis mellifera, and wild-bee richness were equally important in describing resulting open pollination. Wild-bee richness was a better predictor of pollination than wild-bee abundance. We also found evidence suggesting pollinator visitation (and subsequent pollination) are stabilized through the differential response of bee taxa to weather (i.e., response diversity). Variation in the individual visit efficiency of A. mellifera and the southeastern blueberry bee, Habropoda laboriosa, a wild specialist, was not associated with changes in the pollinator community. Our findings add to a growing literature that diverse pollinator communities provide more stable and productive ecosystem services.     

Managed and unmanaged floral margins for the conservation of bee communities in intensive agricultural areas

Floral resources on crop field margins are a well-accepted measure to increase bee abundance in agricultural landscapes. However, studies have mainly focused on managed margins, while studies on the effect of unmanaged floral margins for the conservation of bees are still scarce. This work aims to test and compare the effects of three types of floral margins (managed herbaceous, managed shrubby, and unmanaged herbaceous) on the abundance and diversity of bees in order to propose a management strategy for the conservation of pollinating insects. Bee abundance was recorded by visual samplings in plots of 2 × 2 square meters over two years in the three margin types in four localities in southern Spain. The diversity of plant species and the flowers they supported were measured to explain the associated bee communities. Differences in the relative abundances of bee families and the number and abundance of bee genera were observed between margin types. Andrenidae was generally more abundant in the herbaceous margins regardless of whether these were managed or not. With the exception of the Halictidae, the majority of bees families (wild Apidae, Apis mellifera, Colletidae and Megachilidae) were more abundant in the managed than unmanaged margins. Moreover, the number of bee genera was higher in managed than in unmanaged margins. In addition, here we show that managed margins supported at least 30% more rare bee genera than unmanaged margins, highlighting the importance of floral margins management for the enhancement and conservation of bee communities, restoring habitat and food resources for pollinators across the Mediterranean agricultural landscape.     

Negative effects of pesticides on wild bee communities can be buffered by landscape context

Wild bee communities provide underappreciated but critical agricultural pollination services. Given predicted global shortages in pollination services, managing agroecosystems to support thriving wild bee communities is, therefore, central to ensuring sustainable food production. Benefits of natural (including semi-natural) habitat for wild bee abundance and diversity on farms are well documented. By contrast, few studies have examined toxicity of pesticides on wild bees, let alone effects of farm-level pesticide exposure on entire bee communities. Whether beneficial natural areas could mediate effects of harmful pesticides on wild bees is also unknown. Here, we assess the effect of conventional pesticide use on the wild bee community visiting apple (Malus domestica) within a gradient of percentage natural area in the landscape. Wild bee community abundance and species richness decreased linearly with increasing pesticide use in orchards one year after application; however, pesticide effects on wild bees were buffered by increasing proportion of natural habitat in the surrounding landscape. A significant contribution of fungicides to observed pesticide effects suggests deleterious properties of a class of pesticides that was, until recently, considered benign to bees. Our results demonstrate extended benefits of natural areas for wild pollinators and highlight the importance of considering the landscape context when weighing up the costs of pest management on crop pollination services.     

Pollination efficiency of wild bees and hoverflies provided to oilseed rape

• 1 Declining numbers in honeybees and various wild bee species pose a threat to global pollination services. The identification and quantification of the pollination service provided by different taxa within the pollinator guild is a prerequisite for the successful establishment of nature conservation and crop management regimes.
• 2 Wild bees and hoverflies are considered to be valuable pollinators in agricultural and natural systems. Although some information on pollination efficiency of individual pollinator species is available, comparative studies of both taxa at different densities are rare. In the present study, the efficiency of the solitary mason bee Osmia rufa and two hoverfly species (Eristalis tenax and Episyrphus balteatus) as pollinators of oilseed rape Brassica napus was examined in a standardized caged plant breeding regime. Honeybee Apis mellifera colonies were used as a reference pollinator taxon.
• 3 Yield parameters responded differently to pollinator density and identity. Fruit set and number of seeds per pod increased with increasing pollinator density, although these were stronger in the mason bee than the hoverfly treatment. Weight per 1000 seeds did not respond to any pollinator treatment, indicating that seed quality was not affected. Oilseed rape yield in the highest tested densities of both pollinator taxa resulted in yield values close to the efficiency of small honeybee colonies.
• 4 Hoverflies required approximately five‐fold densities of the red mason bees to reach a similar fruit set and yield. Thus, mason bees are more efficient in plant breeding and managed pollination systems. Both natural pollinator taxa, however, are of potential value in open and closed crop production systems.

     

Railway Embankments as New Habitat for Pollinators in an Agricultural Landscape

Pollinating insect populations, essential for maintaining wild plant diversity and agricultural productivity, rely on (semi)natural habitats. An increasing human population is encroaching upon and deteriorating pollinator habitats. Thus the population persistence of pollinating insects and their associated ecosystem services may depend upon on man-made novel habitats; however, their importance for ecosystem services is barely understood. We tested if man-made infrastructure (railway embankments) in an agricultural landscape establishes novel habitats that support large populations of pollinators (bees, butterflies, hoverflies) when compared to typical habitats for these insects, i.e., semi-natural grasslands. We also identified key environmental factors affecting the species richness and abundance of pollinators on embankments. Species richness and abundance of bees and butterflies were higher for railway embankments than for grasslands. The occurrence of bare (non-vegetated) ground on embankments positively affected bee species richness and abundance, but negatively affected butterfly populations. Species richness and abundance of butterflies positively depended on species richness of native plants on embankments, whereas bee species richness was positively affected by species richness of non-native flowering plants. The density of shrubs on embankments negatively affected the number of bee species and their abundance. Bee and hoverfly species richness were positively related to wood cover in a landscape surrounding embankments. This is the first study showing that railway embankments constitute valuable habitat for the conservation of pollinators in farmland. Specific conservation strategies involving embankments should focus on preventing habitat deterioration due to encroachment of dense shrubs and maintaining grassland vegetation with patches of bare ground.     

Importance of forest fragments as pollinator habitat varies with season and guild

Large areas of Western Europe are covered with intensively managed agricultural land. In these landscapes, wild pollinators depend on fragments of semi-natural habitat for foraging or reproduction. Small forest patches are often the most abundant type of semi-natural habitat in these agricultural landscapes. We investigated the role these patches play in conserving the pollinator community in intensively managed agricultural landscapes.Our survey of the pollinator community in 16 forest fragments showed that the pollinator community in the edges of small forest fragments is strongly influenced by forest and forest edge characteristics. Old forest fragments with a well-developed herb layer had more diverse bee communities than recent forests or old forests without a herb layer, but overall lower activity-abundances, while sun exposure of the forest edges had a strong positive effect on pollinater activity-abundance in general. The hoverfly community had higher activity-abundances in forest edges with a higher flower-index, while saproxylic hoverflies were caught in higher numbers in sites with a higher forest cover in the surrounding landscape.We also detected a strong seasonal effect. The effects of herb layer cover on bee species richness and activity-abundance were much stronger in spring than in summer, while bee species richness was also strongly positively correlated with forest age in spring. A strong positive correlation between pollinator species richness and sun exposure was found in summer, after canopy closure.While the sampled forest edges harbour a rich and diverse pollinator community, cavity-nesting bees were very scarce. This is probably caused by the low amount of dead wood in the studied forest fragments.We conclude that small forest fragments can play an important role in conserving the pollinator community, especially bees and saproxylic hoverflies. The importance of these forest fragments is strongest in spring, when the herb layer provides foraging resources.     

Species Distribution Models for Crop Pollination: A Modelling Framework Applied to Great Britain

Insect pollination benefits over three quarters of the world''s major crops. There is growing concern that observed declines in pollinators may impact on production and revenues from animal pollinated crops. Knowing the distribution of pollinators is therefore crucial for estimating their availability to pollinate crops; however, in general, we have an incomplete knowledge of where these pollinators occur. We propose a method to predict geographical patterns of pollination service to crops, novel in two elements: the use of pollinator records rather than expert knowledge to predict pollinator occurrence, and the inclusion of the managed pollinator supply. We integrated a maximum entropy species distribution model (SDM) with an existing pollination service model (PSM) to derive the availability of pollinators for crop pollination. We used nation-wide records of wild and managed pollinators (honey bees) as well as agricultural data from Great Britain. We first calibrated the SDM on a representative sample of bee and hoverfly crop pollinator species, evaluating the effects of different settings on model performance and on its capacity to identify the most important predictors. The importance of the different predictors was better resolved by SDM derived from simpler functions, with consistent results for bees and hoverflies. We then used the species distributions from the calibrated model to predict pollination service of wild and managed pollinators, using field beans as a test case. The PSM allowed us to spatially characterize the contribution of wild and managed pollinators and also identify areas potentially vulnerable to low pollination service provision, which can help direct local scale interventions. This approach can be extended to investigate geographical mismatches between crop pollination demand and the availability of pollinators, resulting from environmental change or policy scenarios.     

Drivers of Collembola assemblages along an altitudinal gradient in northeast China

Altitudinal changes in the diversity of plants and animals have been well documented; however, soil animals received little attention in this context and it is unclear whether their diversity follows general altitudinal distribution patterns. Changbai Mountain is one of few well‐conserved mountain regions comprising natural ecosystems on the Eurasian continent. Here, we present a comprehensive analysis of the diversity and community composition of Collembola along ten altitudinal sites representing five vegetation types from forest to alpine tundra. Among 7834 Collembola individuals, 84 morphospecies were identified. Species richness varied marginally significant with altitude and generally followed a unimodal relationship with altitude. By contrast, the density of Collembola did not change in a consistent way with altitude. Collembola communities changed gradually with altitude, with local habitat‐related factors (soil and litter carbon‐to‐nitrogen ratio, litter carbon content, and soil pH) and climatic variables (precipitation seasonality) identified as major drivers of changes in Collembola community composition. Notably, local habitat‐related factors explained more variation in Collembola assemblages than climatic variables. The results suggest that local habitat‐related factors including precipitation and temperature are the main drivers of changes in Collembola communities with altitude. Specifically, soil and litter carbon‐to‐nitrogen ratio correlated positively with Collembola communities at high altitudes, whereas soil pH correlated positively at low altitudes. This documents that altitudinal gradients provide unique opportunities for identifying factors driving the community composition of not only above‐ but also belowground invertebrates.     

The relation between oilseed rape and pollination of later flowering plants varies across plant species and landscape contexts

Increasing cultivation of oilseed rape may have consequences for pollinators and wild plant pollination. By providing pollinating insects with pollen and nectar, oilseed rape benefits short-tongued, generalist insect species. Long-tongued bumble bee species, specialized to other flower types, may instead be negatively affected by increased competition from the generalists (e.g. due to nectar-robbing of long-tubed flowers) after oilseed rape flowering has ceased. We expected that the increased abundance of short-tongued pollinators and reduced abundance of long-tongued bumble bees in landscapes with a high proportion of oilseed rape would impact the pollination of later flowering wild plant species. In addition, we expected contrasting effects on plants pollinated by short-tongued pollinators and those pollinated by long-tongued bumble bees. We predicted that semi-natural grasslands, which provide insects with alternative floral resources, would reduce both negative and positive effects on pollination by mitigating competition between pollinators.In 16 semi-natural grasslands, surrounded by agricultural landscapes, with a variation in both the proportion of oilseed rape and the proportion of semi-natural grassland within 1 km, we studied reproductive output in two species of potted plants with different pollination strategies: the woodland strawberry (Fragaria vesca) and red clover (Trifolium pratense). The first species is mainly pollinated by short-tongued pollinators, e.g. hoverflies and solitary bees, and the latter by long-tongued bumble bees. Both species flowered after oilseed rape.Strawberry weight was higher in landscapes with a high proportion of oilseed rape, but only in landscapes with a low proportion of semi-natural grassland. The proportion of developed achenes was also positively related to the proportion of oilseed rape, but only during the latest flowering period. In contrast, red clover seed set was unrelated to the proportion of oilseed rape. Whereas the discrepancy between the two strawberry measurements calls for further research, this study suggests that oilseed rape can affect later flowering plants and that the impact differs among species.     

A contemporary survey of bumble bee diversity across the state of California

Bumble bees (genus Bombus) are important pollinators with more than 260 species found worldwide, many of which are in decline. Twenty‐five species occur in California with the highest species abundance and diversity found in coastal, northern, and montane regions. No recent studies have examined California bumble bee diversity across large spatial scales nor explored contemporary community composition patterns across the state. To fill these gaps, we collected 1740 bumble bee individuals, representing 17 species from 17 sites (~100 bees per site) in California, using an assemblage monitoring framework. This framework is intended to provide an accurate estimate of relative abundance of more common species without negatively impacting populations through overcollection. Our sites were distributed across six ecoregions, with an emphasis on those that historically hosted high bumble bee diversity. We compared bumble bee composition among these sites to provide a snapshot of California bumble bee biodiversity in a single year. Overall, the assemblage monitoring framework that we employed successfully captured estimated relative abundance of species for most sites, but not all. This shortcoming suggests that bumble bee biodiversity monitoring in California might require multiple monitoring approaches, including greater depth of sampling in some regions, given the variable patterns in bumble bee abundance and richness throughout the state. Our study sheds light on the current status of bumble bee diversity in California, identifies some areas where greater sampling effort and conservation action should be focused in the future, and performs the first assessment of an assembly monitoring framework for bumble bee communities in the state.     

Farming practices influence wild pollinator populations on squash and pumpkin

Recent declines in managed honey bee, Apis mellifera L., colonies have increased interest in the current and potential contribution of wild bee populations to the pollination of agricultural crops. Because wild bees often live in agricultural fields, their population density and contribution to crop pollination may be influenced by farming practices, especially those used to reduce the populations of other insects. We took a census of pollinators of squash and pumpkin at 25 farms in Virginia, West Virginia, and Maryland to see whether pollinator abundance was related to farming practices. The main pollinators were Peponapis pruinosa Say; honey bees, and bumble bees (Bombus spp.). The squash bee was the most abundant pollinator on squash and pumpkin, occurring at 23 of 25 farms in population densities that were commonly several times higher than that of other pollinators. Squash bee density was related to tillage practices: no-tillage farms hosted three times as great a density of squash bees as tilled farms. Pollinator density was not related to pesticide use. Honey bee density on squash and pumpkin was not related to the presence of managed honey bee colonies on farms. Farms with colonies did not have more honey bees per flower than farms that did not keep honey bees, probably reflecting the lack of affinity of honey bees for these crops. Future research should examine the economic impacts of managing farms in ways that promote pollinators, particularly pollinators of crops that are not well served by managed honey bee colonies.     

Interacting effects of landscape context and habitat quality on flower visiting insects in agricultural landscapes

Landscape context and habitat quality may have pronounced effects on the diversity of flower visiting insects. We investigated whether the effects of landscape context and habitat quality on flower visiting insects interact in agricultural landscapes in the Netherlands. Landscape context was expressed as the area of semi-natural habitats or the density of linear landscape features, and was quantified at spatial scales ranging from 250 to 2000 m. Habitat quality was determined as flower abundance. Species richness and abundance of hoverflies and bees were determined along 16 stream banks experiencing similar environmental conditions but situated in areas with contrasting landscape context. Only flower abundance and the area of semi-natural habitats within 500–1000 m were significantly related to species richness of hoverflies and bees and these factors had interacting effects on both species groups. Our results suggest that the regional area of semi-natural habitats had a positive effect on hoverfly species richness when flower abundance was relatively high, but not when flower abundance was low. Moreover, flower abundance had positive effects on hoverfly species richness only in areas with relatively many semi-natural habitats. Contrastingly, flower abundance had a more positive effect on bee species richness in landscapes with few semi-natural habitats compared to landscapes with more semi-natural habitats. Our results suggest that the importance of landscape context for the species richness of flower visiting insects depends upon the quality of the habitat patches.     

Managed bumble bees increase flower visitation but not fruit weight in polytunnel strawberry crops

Animal-mediated pollination is essential for the production and quality of fruits and seeds of many crops consumed by humans. However, crop pollination services might be compromised when wild pollinators are scarce. Managed pollinators are commonly used in crops to supplement such services with the assumption that they will enhance crop yield. However, information on the spatiotemporal pollinator-dependence of crops is still limited. We assessed the contribution of commercial bumble bee colonies compared to the available pollinator community on strawberry (‘Fortuna’ variety) flower visitation and strawberry quality across a landscape gradient of agricultural intensification (i.e. polytunnel berry crop cover). We used colonies of bumble bees in winter and in spring, i.e. when few and most wild pollinators are in their flight period, respectively. The placement of colonies increased visits of bumble bees to strawberry flowers, especially in winter. The use of bumble bee colonies did not affect flower visitation by other insects, mainly honey bees, hoverflies and other Diptera. Flower visitation by both honey bees and wild insects did not vary between seasons and was unrelated to the landscape gradient of berry crop cover. Strawberries were of the highest quality (i.e. weight) when insect-mediated pollination was allowed, and their quality was positively related to wild flower visitors in winter but not in spring. However, increased visits to strawberry flowers by managed bumble bees and honey bees had no effect on strawberry weight. Our results suggest that the pollination services producing high quality strawberry fruits are provided by the flower visitor community present in the study region without the need to use managed bumble bees.     

Landscape context and elevation affect pollinator communities in intensive apple orchards

Although an extensive research has been done on the contribution of wild insects to apple pollination, most of these studies did not evaluate the effect of the surrounding landscape context on local pollinator communities. Our aim was to compare communities of wild bees in 31 equally managed apple orchards located in three contrasting landscape types (either dominated by apple, forest, or grasslands) and along an elevation gradient and to test a potential interaction between landscape context and elevation. The study was carried out in 2009 in Trentino (NE Italy), one of the major apple growing areas of Europe with ～12,000 ha of commercial orchards distributed between 150 and 950 m a.s.l. We found that apple-dominated landscapes drastically reduced wild bee species richness and abundance in the orchard compared to landscapes dominated by either grassland or forest. Forest-dominated landscapes benefited local species richness more than grassland-dominated landscapes, while abundance did not differ between grassland and forest. Total species richness and abundance further declined with increasing elevation, while no interactive effect was found between temperature and landscape context. The abundance of Apis mellifera in the apple-dominated landscapes was two to four times higher than in the landscapes dominated by forest and grasslands, respectively. Measures to restore natural pollinator communities by providing suitable habitats around the orchard would not only benefit conservation of general biodiversity, but would probably also contribute to reduce the dependence of apple pollination on managed honey bees.     

Patch utilization and flower visitations by wild bees in a honey bee‐dominated,grassland landscape

Understanding habitat needs and patch utilization of wild and managed bees has been identified as a national research priority in the United States. We used occupancy models to investigate patterns of bee use across 1030 transects spanning a gradient of floral resource abundance and richness and distance from apiaries in the Prairie Pothole Region (PPR) of the United States. Estimates of transect use by honey bees were nearly 1.0 during our 3.5‐month sampling period, suggesting honey bees were nearly ubiquitous across transects. Wild bees more frequently used transects with higher flower richness and more abundant flowers; however, the effect size of the native flower abundance covariate (β^native = 3.90 ± 0.65 [1SE]) was four times greater than the non‐native flower covariate (β^non‐native = 0.99 ± 0.17). We found some evidence that wild bee use was lower at transects near commercial apiaries, but the effect size was imprecise (β^distance = 1.4 ± 0.81). Honey bees were more frequently detected during sampling events with more non‐native flowers and higher species richness but showed an uncertain relationship with native flower abundance. Of the 4039 honey bee and flower interactions, 85% occurred on non‐native flowers, while only 43% of the 738 wild bee observations occurred on non‐native flowers. Our study suggests wild bees and honey bees routinely use the same resource patches in the PPR but often visit different flowering plants. The greatest potential for resource overlap between honey bees and wild bees appears to be for non‐native flowers in the PPR. Our results are valuable to natural resource managers tasked with supporting habitat for managed and wild pollinators in agroecosystems.     

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

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