Foraging habitats and foraging distances of bumblebees, Bombus spp. (Hym., Apidae), in an agricultural landscape

Abstract: In selected foraging habitats of an agricultural landscape flower visits of bumblebees and community structure of foraging bumblebees were studied, with special regard to the role of crops as super-abundant resources. Most crops represent temporal foraging habitats with high abundance of bumblebees but mainly with low diversity in the bumblebee forage community, in contrast to permanent foraging habitats such as, for example, a hedgerow. The high numbers of bumblebees in the monoculture of crop plantations consisted mainly of short-tongued bumblebee species. The role of foraging distances for the visitation rate of foraging habitats was studied by performing capture–recapture experiments with natural nests of Bombus terrestris , Bombus lapidarius and Bombus muscorum . Differences were found on the species as well as the individual level. The foraging distances of B. muscorum were more restricted to the neighbourhood of the nesting habitat than the foraging activity of B. terrestris and B. lapidarius . High percentages of B. terrestris workers were recaptured while foraging on super-abundant resources in distances up to 1750 m from the nest. Isolated patches of highly rewarding forage crops, in agricultural landscapes, are probably only accessed by bumblebee species with large mean foraging distances, such as the short-tongued B. terrestris . Species like the rare, long-tongued B. muscorum depend on a close connection between nesting and foraging habitat. A restricted foraging radius might be one important factor of bumblebee species loss and potential pollinator limitation in modern agricultural landscapes. Furthermore, long-distance flights of bumblebee pollinators have to be considered in the present discussion on gene flow from transgenic plant species on a landscape scale.     

The effect of management practices on bumblebee densities in hedgerow and grassland habitats

Large-scale declines in pollinator species are a concern at present. Such declines have been attributed to a range of factors that act in tandem, rather than in isolation. Some of the most pervasive factors affecting pollinator populations are habitat loss and degradation, which results in the loss of floral resources, nesting sites and landscape connectivity. Intensification of agriculture and urbanisation are two major causes of such habitat alterations. Hedgerows and grasslands are two vital habitats for pollinators in European landscapes. When managed appropriately, these habitats may provide abundant floral resources and nesting opportunities, as well as connectivity between habitats in a fragmented landscape. This study examined the effects that management practices of hedgerows and grasslands may have on bumblebee species, an important group of wild pollinators. Bumblebee abundance was recorded using transect walks in managed and unmanaged sites, including both hedgerows and grasslands. Greater densities of bumblebees were found in unmanaged grasslands in comparison to managed grasslands. Unmanaged hedgerows were also found to have a greater density of bumblebees than managed hedgerows. These results indicate that sites which are less intensively managed provide a more suitable habitat for bumblebees. Therefore, our study underlines the importance of (a) enforcing restrictions on hedge-cutting, and (b) reducing the management intensity of grasslands to provide adequate habitat for pollinators.     

Spatial and temporal variations in floral resource availability affect bumblebee communities in heathlands

Modifications of landscape structure and composition can decrease the availability of floral resources, resulting in the decline of many pollinator species, including bumblebees. These declines may have significant ecological consequences, because bumblebees pollinate a large range of plant species. Our study was carried out in heathlands, open semi-natural habitats that have decreased considerably due to human activities. We analysed how floral resources affect bumblebee communities throughout the colony lifetime at three scales: plot scale, heathland patch scale, and landscape scale. Floral density at the plot scale and spruce plantations at the landscape scale influenced bumblebee communities. The abundance of bumblebees on ericaceous species was higher when the landscape included a substantial proportion of unsuitable foraging habitat (i.e., spruce plantations). Both life history traits and colony life cycle stage influenced bumblebee responses to the availability of floral resources. Bumblebees were more affected by floral resources during the colony development phase than during the nest-foundation or mating phases. Moreover, bumblebees of species that form large colonies needed larger quantities of favourable foraging habitat, compared with small-colony bees, and their proportion decreased in habitats dominated by spruce plantations. In conclusion, the conservation of plant–bumblebee interactions will require management at a larger spatial scale than the restricted protected habitats. Moreover, at the landscape scale, both quantity of favourable foraging patches and their ecological continuity are important to conserve both small- and large- colony species.     

Landscape composition modifies pollinator densities,foraging behavior and yield formation in faba beans

Wildlife-friendly management practices promote pollinators and pollination services in agricultural landscapes. Wild bee densities are driven by landscape composition, as they benefit from an increased availability of nesting and foraging resources at landscape scale. However, effects of landscape composition on bee foraging decisions and consequences for crop pollination have rarely been studied. We investigated, how landscape composition affects bee densities and foraging behavior in faba bean (Vicia faba L.) fields and how this impacts faba bean yield. We recorded densities and nectar robbing behavior of honeybees, long- tongued and short-tongued bumblebees in faba bean fields in eleven landscapes with varying landscape composition (e.g. land cover of oilseed rape, faba bean and semi-natural habitats). Moreover, we assessed yield components of faba beans via pollinator exclusion experiments. Increasing covers of faba bean and semi-natural habitats positively influenced bumblebee densities, while high oilseed rape covers negatively affected short-tongued bumblebee densities in bean fields. Increased faba bean covers enhanced the proportion of nectar-robbing short-tongued bumblebees. The number of beans per pod was increased by insect pollination, while the number of pods was decreased; these effects however depended on variety. Landscape composition interacted with bee densities in shaping yield components in V. faba. Our study emphasizes the importance of considering landscape management to maximize crop yields, as shown for the case of faba beans. The composition of agricultural landscape can modulate bee densities in crop fields, bees foraging behavior and pollination services.     

Mass flowering crops enhance pollinator densities at a landscape scale

To counteract the decline of pollinators in Europe, conservation strategies traditionally focus on enhancing the local availability of semi‐natural habitats, as supported by the European Union's Common Agriculture Policy. In contrast, we show that densities of bumblebees, an important pollinator group in agroecosystems, were not determined by the proportion of semi‐natural habitats in agricultural landscapes. Instead, bumblebee densities were positively related to the availability of highly rewarding mass flowering crops (i.e. oilseed rape) in the landscape. In addition, mass flowering crops were only effective determinants of bumblebee densities when grown extensively at the landscape scale, but not at smaller local scales. Therefore, future conservation measures should consider the importance of mass flowering crops and the need for management schemes at landscape level to sustain vital pollination services in agroecosystems.     

Density and seed set in a self-compatible forb, Penstemon digitalis (Plantaginaceae), with multiple pollinators

Seed production may be limited because flowers do not get enough suitable pollen or because plants lack the resources to make seeds. We used replicated plantings to test factors that influence effects of bumblebee behavior on pollen limitation, as measured by the difference in seed set between hand- and naturally pollinated flowers, of Penstemon digitalis in patches of four to 41 flowering individuals. Seed set per flower was 376% higher in the largest as compared with the smallest Penstemon patches. This positive density dependence reflects activity of long-tongued bees, which (1) have higher effective density as patch size increases, (2) visit greater proportions of plants as patch size increases, and (3) visit smaller proportions of flowers per visited plant as patch size increases. Our results suggest that economics of flight and maneuverability of large, long-tongued bumblebees lead them to transfer more pollen between than within Penstemon plants in large patches. Density of smaller, short-tongued bumblebees was not positively associated with Penstemon seed set, but these bees may be important pollinators at low plant densities. Our experimental system indicates a clear positive relationship between activity of effective pollinators and seed set in a species capable of pollinating itself.     

Mass-flowering red clover crops have positive effects on bumblebee richness and diversity after bloom

Floral resource quantity in agricultural landscapes plays a key role in the persistence of wild pollinators. An equally important, but less investigated factor is how variation in floral resource availability over time, e.g. floral resource pulses, affects pollinator abundances and diversity. Despite the potential importance of late-season resource pulses for bumblebee reproduction, few studies have evaluated the effects of late-season mass-flowering crops on bumblebee abundances and diversity during and after crop bloom. We assessed how bumblebee abundances, diversity and traits associated with species rarity were affected by cultivation of late-season mass-flowering red clover grown for seed production. Bumblebees were surveyed in red clover fields and flower-rich field borders across 20 landscapes with or without a red clover field during and after crop bloom in southern Sweden. Bumblebee worker abundances were higher in clover fields compared to flower-rich borders in the surrounding landscape. There was no relationship between presence of clover fields and the abundance of males of social bumblebees, but more male cuckoo bumblebees were found in flower-rich borders in landscapes with clover following crop bloom. Mass-flowering red clover also had a positive effect on bumblebee species richness and diversity after crop bloom. Overall, clover had positive and lasting effects on less common bumblebees thereby sustaining higher bumblebee species richness after bloom. Cultivation of red clover has the potential, in combination with the management of flower-rich habitats, to benefit less common bumblebee species in temperate agroecosystems.     

Florally rich habitats reduce insect pollination and the reproductive success of isolated plants

Landscape heterogeneity in floral communities has the potential to modify pollinator behavior. Pollinator foraging varies with the diversity, abundance, and spatial configuration of floral resources. However, the implications of this variation for pollen transfer and ultimately the reproductive success of insect pollinated plants remains unclear, especially for species which are rare or isolated in the landscape. We used a landscape‐scale experiment, coupled with microsatellite genotyping, to explore how the floral richness of habitats affected pollinator behavior and pollination effectiveness. Small arrays of the partially self‐compatible plant Californian poppy (Eschscholzia californica) were introduced across a landscape gradient to simulate rare, spatially isolated populations. The effects on pollinator activity, outcrossing, and plant reproduction were measured. In florally rich habitats, we found reduced pollen movement between plants, leading to fewer long‐distance pollination events, lower plant outcrossing, and a higher incidence of pollen limitation. This pattern indicates a potential reduction in per capita pollinator visitation, as suggested by the lower activity densities and richness of pollinators observed within florally rich habitats. In addition, seed production reduced by a factor of 1.8 in plants within florally rich habitats and progeny germination reduced by a factor of 1.2. We show this to be a consequence of self‐fertilization within the partially self‐compatible plant, E. californica. These findings indicate that locally rare plants are at a competitive disadvantage within florally rich habitats because neighboring plant species disrupt conspecific mating by co‐opting pollinators. Ultimately, this Allee effect may play an important role in determining the long‐term persistence of rarer plants in the landscape, both in terms of seed production and viability. Community context therefore requires consideration when designing and implementing conservation management for plants which are comparatively rare in the landscape.     

Pesticide exposure affects flight dynamics and reduces flight endurance in bumblebees

The emergence of agricultural land use change creates a number of challenges that insect pollinators, such as eusocial bees, must overcome. Resultant fragmentation and loss of suitable foraging habitats, combined with pesticide exposure, may increase demands on foraging, specifically the ability to collect or reach sufficient resources under such stress. Understanding effects that pesticides have on flight performance is therefore vital if we are to assess colony success in these changing landscapes. Neonicotinoids are one of the most widely used classes of pesticide across the globe, and exposure to bees has been associated with reduced foraging efficiency and homing ability. One explanation for these effects could be that elements of flight are being affected, but apart from a couple of studies on the honeybee (Apis mellifera), this has scarcely been tested. Here, we used flight mills to investigate how exposure to a field realistic (10 ppb) acute dose of imidacloprid affected flight performance of a wild insect pollinator—the bumblebee, Bombus terrestris audax. Intriguingly, observations showed exposed workers flew at a significantly higher velocity over the first ¾ km of flight. This apparent hyperactivity, however, may have a cost because exposed workers showed reduced flight distance and duration to around a third of what control workers were capable of achieving. Given that bumblebees are central place foragers, impairment to flight endurance could translate to a decline in potential forage area, decreasing the abundance, diversity, and nutritional quality of available food, while potentially diminishing pollination service capabilities.     

Environmentally-friendly and organic management practices enable complementary diversification of plant–bumblebee food webs

Plant and pollinator diversity have declined concurrently in Europe in the last half century. We studied plant–bumblebee food webs to understand the effects of two agri-environmental schemes (AES, organic farming and environmentally-friendly management practice) vs. conventional farming as control group, landscape structure (heterogeneous vs. homogeneous landscapes) and seasonality (June, July, and August) interactions using Estonian AES monitoring data. In the summer of 2014, we observed foraging bumblebees (20 species) on 64 farms that varied in agricultural management and landscape structure, yielding a total of 2303 flower visits on 76 plant species. We found that both management practice and landscape structure influenced the generality (redundancy in the use of flower resources) of food webs. In homogeneous landscapes, environmentally-friendly management practices, including restrictions on the application of glyphosates, enhancement of bumblebee habitats, such as permanent grassland field margins, the allocation of a minimum of 15% of arable land (including rotational grasslands) to legumes, contributed to a higher number of visited plant species (generality) in July, whereas organic farming did so in August. Therefore, both environmentally-friendly and organic management practices are needed to support plant–bumblebee food webs in agricultural landscapes. Food web generality and diversity (Shannon index) are affected by a significant interaction between landscape structure and seasonality: food web diversity varied in homogeneous landscapes between the three different survey months, whereas food webs were more diverse in heterogeneous landscapes. We did not find any significant interaction effect of management, landscape structure and seasonality on linkage density and vulnerability. A full list of the most visited plant species by bumblebees based on species-specific flower visitation was also assembled. In homogeneous landscapes, resource limitation is an issue for bumblebees in certain time periods. For supporting bumblebees in the agricultural landscapes, avoiding resource limitation is important and this can be secured with a combination of AES management practices.     

Population structures and individual performances of <Emphasis Type="Italic">Trillium grandiflorum</Emphasis> in hedgerow and forest habitats

In agricultural landscapes, linear habitats, such as hedgerows at field margins increase structural connectivity among forest patches, potentially providing dispersal corridors for forest herbs. The spatial structure of linear habitats, however, also results in edge effects and perturbations that can influence the individual and population performance of forest plants. This study compares the stage structure and components of growth and reproduction of 14 Trillium grandiflorum populations in hedgerows and forests. Hedgerow Trillium tended to grow faster and, when mature, produced more flowers and more ovules per flowers than forest Trillium, a pattern possibly associated to differences in nutrients and light availability between the two habitats. Seed production and germination rate, however, did not differ between hedgerows and forests. At the population level, seedlings and juveniles were proportionally less abundant in hedgerows than in forests. Although well-established plants can thrive in hedgerows, reduced recruitment may eventually limit the capacity to establish new populations and therefore hamper migration along hedgerow-corridors. Considering the strategies by which plants persist in linear habitats becomes particularly relevant at a time when species are expected to be much in need of dispersal corridors because of climatic stress.     

黄河中下游农业景观中景观简化对传粉昆虫多样性的影响——以巩义市为例

在农业景观中,传粉昆虫的生存繁衍与半自然生境的面积大小有关。集约化生产方式使半自然生境比例逐渐减少,农田比例不断增加,随着景观简化梯度的变化(农田比例逐渐增大),传粉昆虫群落多样性将会发生怎样的变化?选择黄河中下游典型农区巩义市为研究区域,采用诱捕盘法(Pan traps)进行农田、林地的传粉昆虫取样,以21个样点作为景观简化梯度(农田比例范围5%—86%)的呈现,基于每个样点的传粉昆虫多度和丰富度变化来探究景观简化对传粉昆虫多样性的影响。结果显示:区内累计捕获传粉昆虫39660头,优势类群包括双翅目(Diptera)、膜翅目(Hymenoptera)、鞘翅目(Coleoptera)等。采用逐步回归分析及线性拟合后发现景观简化程度与传粉昆虫多度和丰富度呈显著负相关(P0.05);景观简化对传粉昆虫类群间的影响也是有差异的,其中对鞘翅目多度的影响最为密切(R~2=0.27),同时对膜翅目和双翅目也有较大影响(R~2=0.14、R~2=0.11),景观简化与鳞翅目多度呈正相关;随景观简化的程度加深,农田生境中膜翅目多度呈显著下降趋势(P0.05),而林地中膜翅目多度变化不明显。在未来的景观规划中,应着重考虑传粉昆虫中鞘翅目类群的栖息地变化及食物资源状况。依据研究结果建议林地生境中应注重保护现有的自然植被群落,在人工林中可以种植一定面积的蜜粉源植物;农田生境内对杂草群落、半自然生境斑块进行合理规划的基础上,还可以种植线性景观植物作为传粉昆虫的食物源。     

Pollen-Foraging Dynamics of Subalpine Bumblebees (Bombus Latr.)

Abstract A 23-year study on Mount Rainier of nectar-foraging hummingbirds and pollen-foraging bumblebees (Bom-bus Latr.) indicated a constant relative frequency of Bombus species in the area and a wide occurrence of insect species on flowers plus a wide range of flower species pollinated by a single insect. Early-blooming plants had a high queen/worker pollinator ratio, while workers predominated on later-blooming plants. No altitudinal difference in distribution of Bombus species occurred. Analysis of 955 corbicular pollen loads from 1158 bumblebees collected yielded 49.5% monolecty for queens and 34.5% for workers with a wider range for queens than for workers. Colors of corollas photographed in visible and ultraviolet light corresponded to the visual spectra of animal pollinators. In the stable plant-pollinator community investigated, pollinating insect activity appears phenologically controlled by floral anthesis and behaviorally related to floral form and function. Bumblebee tongue length, however. Is not a valid determinant for foraging niche separation except in Castilleja with a deep corolla tube accommodating long tongues of hummingbirds and bumblebees. It is concluded that in a biotic community well-coordinated with the physical environment, limited flower constancy of pollen-foraging pollinators appears to contribute a necessary degree of adaptive versatility through sharing of insect and floral resources. Flower constancy is primarily a function of the circumstance in which a pollinator operates and not simply a characteristic of the pollinator itself.     

Insect and wind pollination of an alpine biennial Aconitum gymnandrum (Ranunculaceae)

Wind pollination can provide reproductive insurance for animal‐pollinated dioecious plants in the absence of available pollinators, but combinations of insect and wind pollination (ambophily) have rarely been studied in hermaphrodite herbs. We examined the stable occurrence of insect pollination and wind pollination over 4 years in a population of a biennial Aconitum species (A. gymnandrum) with actinomorphic and degenerate sepals. The total frequency of visits of two bumblebee species showed no distinct fluctuations in the studied population among the 4 years. However, seed production of netted flowers after emasculation indicated wind pollination had occurred. The seed number of bagged flowers with one visit by bumblebees was significantly less than that of netted flowers after one visit, or in control flowers. Both seed number and fruit set of netted flowers were significantly lower than in control flowers. These results suggest that wind pollination provides supplementary pollen to unvisited and/or once‐visited flowers, but accounts for only a small amount of seed production compared to bumblebee pollination in natural conditions. Such a combination of insect and wind pollination might play an important role in maintaining sexual reproduction of this biennial herb, allowing it to persist in arid habitats on the Qinghai‐Tibetan Plateau, especially during Quaternary glacial periods when pollinator populations oscillated extensively.     

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.     

Mass‐flowering crops dilute pollinator abundance in agricultural landscapes across Europe

Mass‐flowering crops (MFCs) are increasingly cultivated and might influence pollinator communities in MFC fields and nearby semi‐natural habitats (SNHs). Across six European regions and 2 years, we assessed how landscape‐scale cover of MFCs affected pollinator densities in 408 MFC fields and adjacent SNHs. In MFC fields, densities of bumblebees, solitary bees, managed honeybees and hoverflies were negatively related to the cover of MFCs in the landscape. In SNHs, densities of bumblebees declined with increasing cover of MFCs but densities of honeybees increased. The densities of all pollinators were generally unrelated to the cover of SNHs in the landscape. Although MFC fields apparently attracted pollinators from SNHs, in landscapes with large areas of MFCs they became diluted. The resulting lower densities might negatively affect yields of pollinator‐dependent crops and the reproductive success of wild plants. An expansion of MFCs needs to be accompanied by pollinator‐supporting practices in agricultural landscapes.     

Pollination and reproduction of a self-incompatible forest herb in hedgerow corridors and forest patches

Habitat-corridors are assumed to counteract the negative impacts of habitat loss and fragmentation, but their efficiency in doing so depends on the maintenance of ecological processes in corridor conditions. For plants dispersing in linear habitats, one of these critical processes is the maintenance of adequate pollen transfer to insure seed production within the corridor. This study focuses on a common, self-incompatible forest herb, Trillium grandiflorum, to assess plant–pollinator interactions and the influence of spatial processes on plant reproduction in hedgerow corridors compared to forests. First, using pollen supplementation experiments over 2 years, we quantified the extent of pollen limitation in both habitats, testing the prediction of greater limitation in small hedgerow populations than in forests. While pollen limitation of fruit and seed set was common, its magnitude did not differ between habitats. Variations among sites, however, suggested an influence of landscape context on pollination services. Second, we examined the effect of isolation on plant reproduction by monitoring fruit and seed production, as well as pollinator activity and assemblage, in small flower arrays transplanted in hedgerows at increasing distances from forest and from each other. We detected no difference in the proportion of flowers setting fruit or in pollinator activity with isolation, but we observed some differences in pollinator assemblages. Seed set, on the other hand, declined significantly with increasing isolation in the second year of the study, but not in the first year, suggesting altered pollen transfer with distance. Overall, plants in hedgerow corridors and forests benefited from similar pollination services. In this system, plant–pollinator interactions and reproduction seem to be influenced more by variations in resource distribution over years and landscapes than by local habitat conditions. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Pollinator sharing between mass-flowering oilseed rape and co-flowering wild plants: implications for wild plant pollination

Pollinating insects are not only important in wild plant pollination, but also in the production of a large number of crops. Oilseed rape production is increasing globally due to demands for biofuels which may have impacts on pollinating insects which visit the crop and on the pollination services delivered to co-flowering wild plants. In this study, we tested (1) the degree of pollinator sharing between oilseed rape and native wild plants in field margins and hedgerows and (2) the effects of oilseed rape on the quality of pollination service delivered to these wild plants. We found large overlap between flower visitors of wild plants and oilseed rape, but the composition of species overlap differed with respect to each wild plant species. Nearly all individual visitors caught on both the crop and foraging on wild species carried crop pollen, but more than half the insects also carried pollen from wild plants. However, very little oilseed rape pollen was deposited on wild plant stigmas. This shows that (1) oilseed rape overlaps in pollinator niche with most co-flowering wild plants, and (2) crop pollen deposition on wild plant stigmas is low which may indicate that it is unlikely to cause reductions in seed set of wild plants, although this was not measured here. Furthermore, wild plants in field margins and hedgerows are important sources of alternative forage for pollinating insects even when a crop is mass flowering, and we suggest maintenance and augmentation of field margins and hedgerows to provide alternative forage for pollinator conservation to continue provision of pollination services to both crops and wild plants.     

Plant traits and landscape simplification drive intraspecific trait diversity of Bombus terrestris in wildflower plantings

Wildflower plantings are an important mitigation tool within agri-environmental schemes to counter insect decline in resource-scarce agricultural landscapes. Effectiveness of wildflower plantings for insect conservation is typically studied at the community or species level. It is the individual, however, that is subject to changing abiotic and biotic conditions, not the species per se. Accordingly, functional traits of individuals, i.e., the intraspecific functional diversity within species, likely mediate responses to wildflower resources and landscape context. Here we focused on the ecologically and economically important wild insect pollinator Bombus terrestris to study its intraspecific functional diversity and plant-pollinator individual interactions in wildflower plantings. We found considerable trait variation among flower-visiting B. terrestris workers. Locally, this variation could be attributed to flowering plant traits, with larger workers visiting larger inflorescences and individuals with longer tongues preferentially feeding on zygomorphic but not radially symmetrical flowers. In addition, wildflower plantings with high floral abundance attracted individuals with larger pollen baskets. At the landscape scale, increasing proportion of arable land resulted in smaller B. terrestris individuals in wildflower plantings, and a decrease in the overall size diversity of workers. These findings highlight the so far little considered role of intraspecific variation in functional traits of wild pollinators, which can mediate the trait-matching between plants and pollinator individuals. Landscape simplification from agriculture threatens intraspecific pollinator diversity, with potential harmful effects for pollinator fitness and plant reproduction. Tailored wildflower plantings can thus serve as an important tool to increase intraspecific variation in simplified landscapes. When designing seed mixtures for these plantings, high complementarity in plant traits is key for promoting high intraspecific trait diversity of bumblebees and potentially of other associated insect species.     

The effects of landscape on bumblebees to ensure crop pollination in the highland agricultural ecosystems in China

Honey bees and wild bees provide critical pollination services to agricultural ecosystems; however, the relative contributions of different bee taxa are not well understood. The natural habitats surrounding farmland support food and nesting resources for wild bees and therefore play an important role in the maintenance of crop pollination. In this study, we selected Cucurbita pepo L. (squash) as a model crop to investigate the relative importance of honey bees and bumblebees in pollinating the crop. Thirteen fields, which were surrounded by a gradient of natural habitat, were investigated on the Yunnan‐Guizhou Plateau in China. We measured the visit densities of honey bees and bumblebees, the number of pollen grains deposited in a single visit by the two bee taxa, as well as the overall pollen grains deposited on stigmas during a flowering day, and then used Bayesian inference to decouple the pollen grains deposited by either the honey bees or the bumblebees. Compared with honey bees, bumblebees deposited a higher number of pollen grains on stigmas in a single visit, but had a lower visit density than honey bees. Meanwhile, the bumblebee visit density increased along the proportion of natural habitat, while the honey bee visit density was not affected by the surrounding natural habitat. Data simulations using Bayesian inference showed that on a flowering day, the number of pollen grains deposited by bumblebees increased with the proportion of natural habitat in the surrounding landscape, but the number of pollen grains deposited by honey bees did not. Moreover, the total numbers of pollen grains deposited by honey bees or bumblebees alone were all below 2000 (the critical level to satisfy the pollination requirement of this crop). Pollen calculations demonstrated that the number of pollen grains deposited by the two bee taxa was greater than 2000 in fields surrounded by more than 13% natural habitat (grasslands and forests). The results revealed that bumblebees ensured C. pepo pollination in combination with honey bees in the highland agricultural ecosystems.