Functional redundancy of weed seed predation is reduced by intensified agriculture.

Intensified agriculture, a driver of biodiversity loss, can diminish ecosystem functions and their stability. Biodiversity can increase functional redundancy and is expected to stabilize ecosystem functions. Few studies, however, have explored how agricultural intensity affects functional redundancy and its link with ecosystem function stability. Here, within a continental-wide study, we assess how functional redundancy of seed predation is affected by agricultural intensity and landscape simplification. By combining carabid abundances with molecular gut content data, functional redundancy of seed predation was quantified for 65 weed genera across 60 fields in four European countries. Across weed genera, functional redundancy was reduced with high field management intensity and simplified crop rotations. Moreover, functional redundancy increased the spatial stability of weed seed predation at the field scale. We found that ecosystem functions are vulnerable to disturbances in intensively managed agroecosystems, providing empirical evidence of the importance of biodiversity for stable ecosystem functions across space.

Crop rotational diversity can mitigate climate-induced grain yield losses.

Diversified crop rotations have been suggested to reduce grain yield losses from the adverse climatic conditions increasingly common under climate change. Nevertheless, the potential for climate change adaptation of different crop rotational diversity (CRD) remains undetermined. We quantified how climatic conditions affect small grain and maize yields under different CRDs in 32 long-term (10-63 years) field experiments across Europe and North America. Species-diverse and functionally rich rotations more than compensated yield losses from anomalous warm conditions, long and warm dry spells, as well as from anomalous wet (for small grains) or dry (for maize) conditions. Adding a single functional group or crop species to monocultures counteracted yield losses from substantial changes in climatic conditions. The benefits of a further increase in CRD are comparable with those of improved climatic conditions. For instance, the maize yield benefits of adding three crop species to monocultures under detrimental climatic conditions exceeded the average yield of monocultures by up to 553 kg/ha under non-detrimental climatic conditions. Increased crop functional richness improved yields under high temperature, irrespective of precipitation. Conversely, yield benefits peaked at between two and four crop species in the rotation, depending on climatic conditions and crop, and declined at higher species diversity. Thus, crop species diversity could be adjusted to maximize yield benefits. Diversifying rotations with functionally distinct crops is an adaptation of cropping systems to global warming and changes in precipitation.

From a local descriptive to a generic predictive model of cereal aphid regulation by predators.

The temporal dynamics of insect populations in agroecosystems are influenced by numerous biotic and abiotic interactions, including trophic interactions in complex food webs. Predicting the regulation of herbivorous insect pests by arthropod predators and parasitoids would allow for rendering crop production less dependent on chemical pesticides. Curtsdotter et al. (2019) developed a food-web model simulating the influences of naturally occurring arthropod predators on aphid population dynamics in cereal crop fields. The use of an allometric hypothesis based on the relative body masses of the prey and various predator guilds reduced the number of estimated parameters to just five, albeit field-specific. Here, we extend this model and test its applicability and predictive capacity. We first parameterized the original model with a dataset with the dynamic arthropod community compositions in 54 fields in six regions in France. We then integrated three additional biological functions to the model: parasitism, aphid carrying capacity and suboptimal high temperatures that reduce aphid growth rates. We developed a multi-field calibration approach to estimate a single set of generic allometric parameters for a given group of fields, which would increase model generality needed for predictions. The original and revised models, when using field-specific parameterization, achieved quantitatively good fits to observed aphid population dynamics for 59% and 53% of the fields, respectively, with pseudo-R2 up to 0.99. But the multi-field calibration showed that increased model generality came at the cost of reduced model reliability (goodness-of-fit). Our study highlights the need to further improve our understanding of how body size and other traits affect trophic interactions in food webs. It also points up the need to acquire high-resolution data to use this type of modelling approach. We propose that a hypothesis-driven strategy of model improvement based on the integration of additional biological functions and additional functional traits beyond body size (e.g., predator space search or prey defences) into the food-web matrix can improve model reliability.

Cold winters drive consistent and spatially synchronous 8-year population cycles of cabbage stem flea beetle.

Population cycles have been observed in mammals as well as insects, but consistent population cycling has rarely been documented in agroecosystems and never for a beetle. We analysed the long-term population patterns of the cabbage stem flea beetle Psylliodes chrysocephala in winter oilseed rape over 50 years. Psylliodes chrysocephala larval density from 3045 winter oilseed rape fields in southern Sweden showed strong 8-year population cycles in regional mean density. Fluctuations in larval density were synchronous over time across five subregional populations. Subregional mean environmental variables explained 90.6% of the synchrony in P. chrysocephala populations at the 7-11 year time-scale. The number of days below -10°C showed strong anti-phase coherence with larval densities in the 7-11 year time-scale, such that more cold days resulted in low larval densities. High levels of the North Atlantic Oscillation weather system are coherent and anti-phase with cold weather in Scania, Sweden. At the field-scale, later crop planting date and more cold winter days were associated with decreased overwintering larval density. Warmer autumn temperatures, resulting in greater larval accumulated degree days early in the season, increased overwintering larval density. Despite variation in environmental conditions and crop management, 8-year cycles persisted for cabbage stem flea beetle throughout the 50 years of data collection. Moran effects, influenced by the North Atlantic Oscillation weather patterns, are the primary drivers of this cycle and synchronicity. Insect pest data collected in commercial agriculture fields is an abundant source of long-term data. We show that an agricultural pest can have the same periodic population cycles observed in perennial and unmanaged ecosystems. This unexpected finding has implications for sustainable pest management in agriculture and shows the value of long-term pest monitoring projects as an additional source of time-series data to untangle the drivers of population cycles.

The coordination of green-brown food webs and their disruption by anthropogenic nutrient inputs.

Aim: Our goal was to quantify nitrogen flows and stocks in green-brown food webs in different ecosystems, how they differ across ecosystems and how they respond to nutrient enrichment. Location: Global. Time period: Contemporary. Major taxa studied: Plants, phytoplankton, macroalgae, invertebrates, vertebrates and zooplankton. Methods: Data from >500 studies were combined to estimate nitrogen stocks and fluxes in green-brown food webs in forests, grasslands, brackish environments, seagrass meadows, lakes and oceans. We compared the stocks, fluxes and metabolic rates of different functional groups within each food web. We also used these estimates to build a dynamical model to test the response of the ecosystems to nutrient enrichment. Results: We found surprising symmetries between the green and brown channels across ecosystems, in their stocks, fluxes and consumption coefficients and mortality rates. We also found that nitrogen enrichment, either organic or inorganic, can disrupt this balance between the green and brown channels. Main conclusions: Linking green and brown food webs reveals a previously hidden symmetry between herbivory and detritivory, which appears to be a widespread property of natural ecosystems but can be disrupted by anthropogenic nitrogen additions.

Water stress and insect herbivory interactively reduce crop yield while the insect pollination benefit is conserved.

Climate change is predicted to hamper crop production due to precipitation deficits and warmer temperatures inducing both water stress and increasing herbivory due to more abundant insect pests. Consequently, crop yields will be impacted simultaneously by abiotic and biotic stressors. Extensive yield losses due to such climate change stressors might, however, be mitigated by ecosystem services such as insect pollination. We examined the single and combined effects of water stress, insect herbivory and insect pollination on faba bean yield components and above- and belowground plant biomass under realistic field conditions. We used rainout shelters to simulate a scenario in line with climate change projections, with adequate water supply at sowing followed by a long period without precipitation. This induced a gradually increasing water stress, culminating around crop flowering and yield formation. We found that gradually increasing water stress combined with insect herbivory by aphids interactively shaped yield in faba beans. Individually, aphid herbivory reduced yield by 79% and water stress reduced yield by 52%. However, the combined effect of water stress and aphid herbivory reduced yield less (84%) than the sum of the individual stressor effects. In contrast, insect pollination increased yield by 68% independently of water availability and insect herbivory. Our results suggest that yield losses can be greatly reduced when both water stress and insect herbivory are reduced simultaneously. In contrast, reducing only one stressor has negligible benefits on yield as long as the crop is suffering from the other stressor. We call for further exploration of interactions among ecosystem services and biotic and abiotic stressors that simulate realistic conditions under climate change.

Land-use intensity affects the potential for apparent competition within and between habitats.

Arthropod communities dwelling in adjacent habitats are able to impact one another via shared natural enemies. In agricultural landscapes, drastic differences in resource availability between crop and non-crop habitats cause variation in insect herbivore densities over short distances, potentially driving inter-habitat effects. Moreover, the composition of the landscape in which the habitats are embedded likely affects realised attack rates from natural enemies via impacts on local arthropod community structure. Here, we examine indirect effects between herbivore species within and between habitat types by calculating the potential for apparent competition between multiple populations. Firstly, we aim to determine how disparities in resource availability impact the strength of the potential for apparent competition occurring between habitats, secondly to examine the impact of landscape composition upon these effects, and finally to couch these observations in reality by investigating the link between the potential for apparent competition and realised attack rates. We used DNA metabarcoding to characterise host-parasitoid interactions within two habitat types (with divergent nutrient inputs) at 11 locations with variable landscape composition within an agroecosystem context. We then used these interaction networks to estimate the potential for apparent competition between each host pair and to compare expected versus realised attack rates across the system. Shared natural enemies were found to structure host herbivore communities within and across habitat boundaries. The size of this effect was related to the resource availability of habitats, such that the habitat with high nutrient input exerted a stronger effect. The overall potential for apparent competition declined with increasing land-use intensity in the surrounding landscape and exhibited a discernible impact on realised attack rates upon herbivore species. Thus, our results suggest that increasing the proportion of perennial habitat in agroecosystems could increase the prevalence of indirect effects such as apparent competition among insect herbivore communities, potentially leading to enhanced population regulation via increased attack rates from natural enemies like parasitoid wasps.

Seed coating with a neonicotinoid insecticide negatively affects wild bees.

Understanding the effects of neonicotinoid insecticides on bees is vital because of reported declines in bee diversity and distribution and the crucial role bees have as pollinators in ecosystems and agriculture. Neonicotinoids are suspected to pose an unacceptable risk to bees, partly because of their systemic uptake in plants, and the European Union has therefore introduced a moratorium on three neonicotinoids as seed coatings in flowering crops that attract bees. The moratorium has been criticized for being based on weak evidence, particularly because effects have mostly been measured on bees that have been artificially fed neonicotinoids. Thus, the key question is how neonicotinoids influence bees, and wild bees in particular, in real-world agricultural landscapes. Here we show that a commonly used insecticide seed coating in a flowering crop can have serious consequences for wild bees. In a study with replicated and matched landscapes, we found that seed coating with Elado, an insecticide containing a combination of the neonicotinoid clothianidin and the non-systemic pyrethroid ß-cyfluthrin, applied to oilseed rape seeds, reduced wild bee density, solitary bee nesting, and bumblebee colony growth and reproduction under field conditions. Hence, such insecticidal use can pose a substantial risk to wild bees in agricultural landscapes, and the contribution of pesticides to the global decline of wild bees may have been underestimated. The lack of a significant response in honeybee colonies suggests that reported pesticide effects on honeybees cannot always be extrapolated to wild bees.

Crop pests and predators exhibit inconsistent responses to surrounding landscape composition.

The idea that noncrop habitat enhances pest control and represents a win-win opportunity to conserve biodiversity and bolster yields has emerged as an agroecological paradigm. However, while noncrop habitat in landscapes surrounding farms sometimes benefits pest predators, natural enemy responses remain heterogeneous across studies and effects on pests are inconclusive. The observed heterogeneity in species responses to noncrop habitat may be biological in origin or could result from variation in how habitat and biocontrol are measured. Here, we use a pest-control database encompassing 132 studies and 6,759 sites worldwide to model natural enemy and pest abundances, predation rates, and crop damage as a function of landscape composition. Our results showed that although landscape composition explained significant variation within studies, pest and enemy abundances, predation rates, crop damage, and yields each exhibited different responses across studies, sometimes increasing and sometimes decreasing in landscapes with more noncrop habitat but overall showing no consistent trend. Thus, models that used landscape-composition variables to predict pest-control dynamics demonstrated little potential to explain variation across studies, though prediction did improve when comparing studies with similar crop and landscape features. Overall, our work shows that surrounding noncrop habitat does not consistently improve pest management, meaning habitat conservation may bolster production in some systems and depress yields in others. Future efforts to develop tools that inform farmers when habitat conservation truly represents a win-win would benefit from increased understanding of how landscape effects are modulated by local farm management and the biology of pests and their enemies.

The effectiveness of flower strips and hedgerows on pest control, pollination services and crop yield: a quantitative synthesis.

Floral plantings are promoted to foster ecological intensification of agriculture through provisioning of ecosystem services. However, a comprehensive assessment of the effectiveness of different floral plantings, their characteristics and consequences for crop yield is lacking. Here we quantified the impacts of flower strips and hedgerows on pest control (18 studies) and pollination services (17 studies) in adjacent crops in North America, Europe and New Zealand. Flower strips, but not hedgerows, enhanced pest control services in adjacent fields by 16% on average. However, effects on crop pollination and yield were more variable. Our synthesis identifies several important drivers of variability in effectiveness of plantings: pollination services declined exponentially with distance from plantings, and perennial and older flower strips with higher flowering plant diversity enhanced pollination more effectively. These findings provide promising pathways to optimise floral plantings to more effectively contribute to ecosystem service delivery and ecological intensification of agriculture in the future.

Species traits elucidate crop pest response to landscape composition: a global analysis.

Recent synthesis studies have shown inconsistent responses of crop pests to landscape composition, imposing a fundamental limit to our capacity to design sustainable crop protection strategies to reduce yield losses caused by insect pests. Using a global dataset composed of 5242 observations encompassing 48 agricultural pest species and 26 crop species, we tested the role of pest traits (exotic status, host breadth and habitat breadth) and environmental context (crop type, range in landscape gradient and climate) in modifying the pest response to increasing semi-natural habitats in the surrounding landscape. For natives, increasing semi-natural habitats decreased the abundance of pests that exploit only crop habitats or that are highly polyphagous. On the contrary, populations of exotic pests increased with an increasing cover of semi-natural habitats. These effects might be related to changes in host plants and other resources across the landscapes and/or to modified top-down control by natural enemies. The range of the landscape gradient explored and climate did not affect pests, while crop type modified the response of pests to landscape composition. Although species traits and environmental context helped in explaining some of the variability in pest response to landscape composition, the observed large interspecific differences suggest that a portfolio of strategies must be considered and implemented for the effective control of rapidly changing communities of crop pests in agroecosystems.

The interplay of landscape composition and configuration: new pathways to manage functional biodiversity and agroecosystem services across Europe.

Managing agricultural landscapes to support biodiversity and ecosystem services is a key aim of a sustainable agriculture. However, how the spatial arrangement of crop fields and other habitats in landscapes impacts arthropods and their functions is poorly known. Synthesising data from 49 studies (1515 landscapes) across Europe, we examined effects of landscape composition (% habitats) and configuration (edge density) on arthropods in fields and their margins, pest control, pollination and yields. Configuration effects interacted with the proportions of crop and non-crop habitats, and species' dietary, dispersal and overwintering traits led to contrasting responses to landscape variables. Overall, however, in landscapes with high edge density, 70% of pollinator and 44% of natural enemy species reached highest abundances and pollination and pest control improved 1.7- and 1.4-fold respectively. Arable-dominated landscapes with high edge densities achieved high yields. This suggests that enhancing edge density in European agroecosystems can promote functional biodiversity and yield-enhancing ecosystem services.

Ecosystem function in predator-prey food webs-confronting dynamic models with empirical data.

Most ecosystem functions and related services involve species interactions across trophic levels, for example, pollination and biological pest control. Despite this, our understanding of ecosystem function in multitrophic communities is poor, and research has been limited to either manipulation in small communities or statistical descriptions in larger ones. Recent advances in food web ecology may allow us to overcome the trade-off between mechanistic insight and ecological realism. Molecular tools now simplify the detection of feeding interactions, and trait-based approaches allow the application of dynamic food web models to real ecosystems. We performed the first test of an allometric food web model's ability to replicate temporally nonaggregated abundance data from the field and to provide mechanistic insight into the function of predation. We aimed to reproduce and explore the drivers of the population dynamics of the aphid herbivore Rhopalosiphum padi observed in ten Swedish barley fields. We used a dynamic food web model, taking observed interactions and abundances of predators and alternative prey as input data, allowing us to examine the role of predation in aphid population control. The inverse problem methods were used for simultaneous model fit optimization and model parameterization. The model captured >70% of the variation in aphid abundance in five of ten fields, supporting the model-embodied hypothesis that body size can be an important determinant of predation in the arthropod community. We further demonstrate how in-depth model analysis can disentangle the likely drivers of function, such as the community's abundance and trait composition. Analysing the variability in model performance revealed knowledge gaps, such as the source of episodic aphid mortality, and general method development needs that, if addressed, would further increase model success and enable stronger inference about ecosystem function. The results demonstrate that confronting dynamic food web models with abundance data from the field is a viable approach to evaluate ecological theory and to aid our understanding of function in real ecosystems. However, to realize the full potential of food web models, in ecosystem function research and beyond, trait-based parameterization must be refined and extended to include more traits than body size.

Non-bee insects are important contributors to global crop pollination.

Wild and managed bees are well documented as effective pollinators of global crops of economic importance. However, the contributions by pollinators other than bees have been little explored despite their potential to contribute to crop production and stability in the face of environmental change. Non-bee pollinators include flies, beetles, moths, butterflies, wasps, ants, birds, and bats, among others. Here we focus on non-bee insects and synthesize 39 field studies from five continents that directly measured the crop pollination services provided by non-bees, honey bees, and other bees to compare the relative contributions of these taxa. Non-bees performed 25-50% of the total number of flower visits. Although non-bees were less effective pollinators than bees per flower visit, they made more visits; thus these two factors compensated for each other, resulting in pollination services rendered by non-bees that were similar to those provided by bees. In the subset of studies that measured fruit set, fruit set increased with non-bee insect visits independently of bee visitation rates, indicating that non-bee insects provide a unique benefit that is not provided by bees. We also show that non-bee insects are not as reliant as bees on the presence of remnant natural or seminatural habitat in the surrounding landscape. These results strongly suggest that non-bee insect pollinators play a significant role in global crop production and respond differently than bees to landscape structure, probably making their crop pollination services more robust to changes in land use. Non-bee insects provide a valuable service and provide potential insurance against bee population declines.

Predictive power of food web models based on body size decreases with trophic complexity.

Food web models parameterised using body size show promise to predict trophic interaction strengths (IS) and abundance dynamics. However, this remains to be rigorously tested in food webs beyond simple trophic modules, where indirect and intraguild interactions could be important and driven by traits other than body size. We systematically varied predator body size, guild composition and richness in microcosm insect webs and compared experimental outcomes with predictions of IS from models with allometrically scaled parameters. Body size was a strong predictor of IS in simple modules (r2  = 0.92), but with increasing complexity the predictive power decreased, with model IS being consistently overestimated. We quantify the strength of observed trophic interaction modifications, partition this into density-mediated vs. behaviour-mediated indirect effects and show that model shortcomings in predicting IS is related to the size of behaviour-mediated effects. Our findings encourage development of dynamical food web models explicitly including and exploring indirect mechanisms.

Combined effects of agrochemicals and ecosystem services on crop yield across Europe.

Simultaneously enhancing ecosystem services provided by biodiversity below and above ground is recommended to reduce dependence on chemical pesticides and mineral fertilisers in agriculture. However, consequences for crop yield have been poorly evaluated. Above ground, increased landscape complexity is assumed to enhance biological pest control, whereas below ground, soil organic carbon is a proxy for several yield-supporting services. In a field experiment replicated in 114 fields across Europe, we found that fertilisation had the strongest positive effect on yield, but hindered simultaneous harnessing of below- and above-ground ecosystem services. We furthermore show that enhancing natural enemies and pest control through increasing landscape complexity can prove disappointing in fields with low soil services or in intensively cropped regions. Thus, understanding ecological interdependences between land use, ecosystem services and yield is necessary to promote more environmentally friendly farming by identifying situations where ecosystem services are maximised and agrochemical inputs can be reduced.

Above- and belowground insect herbivory modifies the response of a grassland plant community to nitrogen eutrophication.

Understanding the role that species interactions play in determining the rate and direction of ecosystem change due to nitrogen (N) eutrophication is important for predicting the consequences of global change. Insects might play a major role in this context. They consume substantial amounts of plant biomass and can alter competitive interactions among plants, indirectly shaping plant community composition. Nitrogen eutrophication affects plant communities globally, but there is limited experimental evidence of how insect herbivory modifies plant community response to raised N levels. Even less is known about the roles of above- and belowground herbivory in shaping plant communities, and how the interaction between the two might modify a plant community's response to N eutrophication. We conducted a 3-yr field experiment where grassland plant communities were subjected to above- and belowground insect herbivory with and without N addition, in a full-factorial design. We found that herbivory modified plant community responses to N addition. Aboveground herbivory decreased aboveground plant community biomass by 21%, but only at elevated N. When combined, above- and belowground herbivory had a stronger negative effect on plant community biomass at ambient N (11% decrease) than at elevated N (4% decrease). In addition, herbivory shifted the functional composition of the plant community, and the magnitude of the shifts depended on the N level. The N and herbivory treatments synergistically conferred a competitive advantage to forbs, which benefited when both herbivory types were present at elevated N. Evenness among the plant species groups increased when aboveground herbivory was present, but N addition attenuated this increase. Our results demonstrate that a deeper understanding of how plant-herbivore interactions above and below ground shape the composition of a plant community is crucial for making reliable predictions about the ecological consequences of global change.

Landscape simplification weakens the association between terrestrial producer and consumer diversity in Europe.

Land-use change is one of the primary drivers of species loss, yet little is known about its effect on other components of biodiversity that may be at risk. Here, we ask whether, and to what extent, landscape simplification, measured as the percentage of arable land in the landscape, disrupts the functional and phylogenetic association between primary producers and consumers. Across seven European regions, we inferred the potential associations (functional and phylogenetic) between host plants and butterflies in 561 seminatural grasslands. Local plant diversity showed a strong bottom-up effect on butterfly diversity in the most complex landscapes, but this effect disappeared in simple landscapes. The functional associations between plant and butterflies are, therefore, the results of processes that act not only locally but are also dependent on the surrounding landscape context. Similarly, landscape simplification reduced the phylogenetic congruence among host plants and butterflies indicating that closely related butterflies become more generalist in the resources used. These processes occurred without any detectable change in species richness of plants or butterflies along the gradient of arable land. The structural properties of ecosystems are experiencing substantial erosion, with potentially pervasive effects on ecosystem functions and future evolutionary trajectories. Loss of interacting species might trigger cascading extinction events and reduce the stability of trophic interactions, as well as influence the longer term resilience of ecosystem functions. This underscores a growing realization that species richness is a crude and insensitive metric and that both functional and phylogenetic associations, measured across multiple trophic levels, are likely to provide additional and deeper insights into the resilience of ecosystems and the functions they provide.

A global synthesis of the effects of diversified farming systems on arthropod diversity within fields and across agricultural landscapes.

Agricultural intensification is a leading cause of global biodiversity loss, which can reduce the provisioning of ecosystem services in managed ecosystems. Organic farming and plant diversification are farm management schemes that may mitigate potential ecological harm by increasing species richness and boosting related ecosystem services to agroecosystems. What remains unclear is the extent to which farm management schemes affect biodiversity components other than species richness, and whether impacts differ across spatial scales and landscape contexts. Using a global metadataset, we quantified the effects of organic farming and plant diversification on abundance, local diversity (communities within fields), and regional diversity (communities across fields) of arthropod pollinators, predators, herbivores, and detritivores. Both organic farming and higher in-field plant diversity enhanced arthropod abundance, particularly for rare taxa. This resulted in increased richness but decreased evenness. While these responses were stronger at local relative to regional scales, richness and abundance increased at both scales, and richness on farms embedded in complex relative to simple landscapes. Overall, both organic farming and in-field plant diversification exerted the strongest effects on pollinators and predators, suggesting these management schemes can facilitate ecosystem service providers without augmenting herbivore (pest) populations. Our results suggest that organic farming and plant diversification promote diverse arthropod metacommunities that may provide temporal and spatial stability of ecosystem service provisioning. Conserving diverse plant and arthropod communities in farming systems therefore requires sustainable practices that operate both within fields and across landscapes.

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.