Pesticide use negatively affects bumble bees across European landscapes.

Sustainable agriculture requires balancing crop yields with the effects of pesticides on non-target organisms, such as bees and other crop pollinators. Field studies demonstrated that agricultural use of neonicotinoid insecticides can negatively affect wild bee species1,2, leading to restrictions on these compounds3. However, besides neonicotinoids, field-based evidence of the effects of landscape pesticide exposure on wild bees is lacking. Bees encounter many pesticides in agricultural landscapes4-9 and the effects of this landscape exposure on colony growth and development of any bee species remains unknown. Here we show that the many pesticides found in bumble bee-collected pollen are associated with reduced colony performance during crop bloom, especially in simplified landscapes with intensive agricultural practices. Our results from 316 Bombus terrestris colonies at 106 agricultural sites across eight European countries confirm that the regulatory system fails to sufficiently prevent pesticide-related impacts on non-target organisms, even for a eusocial pollinator species in which colony size may buffer against such impacts10,11. These findings support the need for postapproval monitoring of both pesticide exposure and effects to confirm that the regulatory process is sufficiently protective in limiting the collateral environmental damage of agricultural pesticide use.

Carbon-biodiversity relationships in a highly diverse subtropical forest.

Carbon-focused climate mitigation strategies are becoming increasingly important in forests. However, with ongoing biodiversity declines we require better knowledge of how much such strategies account for biodiversity. We particularly lack information across multiple trophic levels and on established forests, where the interplay between carbon stocks, stand age, and tree diversity might influence carbon-biodiversity relationships. Using a large dataset (>4600 heterotrophic species of 23 taxonomic groups) from secondary, subtropical forests, we tested how multitrophic diversity and diversity within trophic groups relate to aboveground, belowground, and total carbon stocks at different levels of tree species richness and stand age. Our study revealed that aboveground carbon, the key component of climate-based management, was largely unrelated to multitrophic diversity. By contrast, total carbon stocks-that is, including belowground carbon-emerged as a significant predictor of multitrophic diversity. Relationships were nonlinear and strongest for lower trophic levels, but nonsignificant for higher trophic level diversity. Tree species richness and stand age moderated these relationships, suggesting long-term regeneration of forests may be particularly effective in reconciling carbon and biodiversity targets. Our findings highlight that biodiversity benefits of climate-oriented management need to be evaluated carefully, and only maximizing aboveground carbon may fail to account for biodiversity conservation requirements.

Nutritional stress exacerbates impact of a novel insecticide on solitary bees' behaviour, reproduction and survival.

Pesticide exposure and food stress are major threats to bees, but their potential synergistic impacts under field-realistic conditions remain poorly understood and are not considered in current pesticide risk assessments. We conducted a semi-field experiment to examine the single and interactive effects of the novel insecticide flupyradifurone (FPF) and nutritional stress on fitness proxies in the solitary bee Osmia bicornis. Individually marked bees were released into flight cages with monocultures of buckwheat, wild mustard or purple tansy, which were assigned to an insecticide treatment (FPF or control) in a crossed design. Nutritional stress, which was high in bees foraging on buckwheat, intermediate on wild mustard and low on purple tansy, modulated the impact of insecticide exposure. Within the first day after application of FPF, mortality of bees feeding on buckwheat was 29 times higher compared with control treatments, while mortality of FPF exposed and control bees was similar in the other two plant species. Moreover, we found negative synergistic impacts of FPF and nutritional stress on offspring production, flight activity, flight duration and flower visitation frequency. These results reveal that environmental policies and risk assessment schemes that ignore interactions among anthropogenic stressors will fail to adequately protect bees and the pollination services they provide.

Land-use intensification causes multitrophic homogenization of grassland communities.

Land-use intensification is a major driver of biodiversity loss. Alongside reductions in local species diversity, biotic homogenization at larger spatial scales is of great concern for conservation. Biotic homogenization means a decrease in ß-diversity (the compositional dissimilarity between sites). Most studies have investigated losses in local (α)-diversity and neglected biodiversity loss at larger spatial scales. Studies addressing ß-diversity have focused on single or a few organism groups (for example, ref. 4), and it is thus unknown whether land-use intensification homogenizes communities at different trophic levels, above- and belowground. Here we show that even moderate increases in local land-use intensity (LUI) cause biotic homogenization across microbial, plant and animal groups, both above- and belowground, and that this is largely independent of changes in α-diversity. We analysed a unique grassland biodiversity dataset, with abundances of more than 4,000 species belonging to 12 trophic groups. LUI, and, in particular, high mowing intensity, had consistent effects on ß-diversity across groups, causing a homogenization of soil microbial, fungal pathogen, plant and arthropod communities. These effects were nonlinear and the strongest declines in ß-diversity occurred in the transition from extensively managed to intermediate intensity grassland. LUI tended to reduce local α-diversity in aboveground groups, whereas the α-diversity increased in belowground groups. Correlations between the ß-diversity of different groups, particularly between plants and their consumers, became weaker at high LUI. This suggests a loss of specialist species and is further evidence for biotic homogenization. The consistently negative effects of LUI on landscape-scale biodiversity underscore the high value of extensively managed grasslands for conserving multitrophic biodiversity and ecosystem service provision. Indeed, biotic homogenization rather than local diversity loss could prove to be the most substantial consequence of land-use intensification.

Multi-trophic communities re-establish with canopy cover and microclimate in a subtropical forest biodiversity experiment.

Plant diversity affects multi-trophic communities, but in young regrowth forests, where forest insects are in the process of re-establishment, other biotic and also abiotic factors might be more important. We studied cavity-nesting bees, wasps and their natural enemies along an experimental tree diversity gradient in subtropical South-East China. We compared insect communities of experimental young forests with communities of established natural forests nearby the experiment and tested for direct and indirect effects of tree diversity, tree basal area (a proxy of tree biomass), canopy cover and microclimate on bee and wasp community composition, abundance and species richness. Finally, we tested if the trophic levels of bees, herbivore-hunting wasps, spider-hunting wasps and their natural enemies respond similarly. Forest bee and wasp community composition re-established towards communities of the natural forest with increasing tree biomass and canopy cover. These factors directly and indirectly, via microclimatic conditions, increased the abundance of bees, wasps and their natural enemies. While bee and wasp species richness increased with abundance and both were not related to tree diversity, abundance increased directly with canopy cover, mediated by tree biomass. Abundance of natural enemies increased with host (bee and wasp) abundance irrespective of their trophic position. In conclusion, although maximizing tree diversity is an important goal of reforestation and forest conservation, rapid closure of canopies is also important for re-establishing communities of forest bees, wasps and their natural enemies.

Exotic garden plants partly substitute for native plants as resources for pollinators when native plants become seasonally scarce.

Urban green spaces such as gardens often consist of native and exotic plant species, which provide pollen and nectar for flower-visiting insects. Although some exotic plants are readily visited by pollinators, it is unknown if and at which time of the season exotic garden plants may supplement or substitute for flower resources provided by native plants. To investigate if seasonal changes in flower availability from native vs. exotic plants affect flower visits, diversity and particularly plant-pollinator interaction networks, we studied flower-visiting insects over a whole growing season in 20 urban residential gardens in Germany. Over the course of the season, visits to native plants decreased, the proportion of flower visits to exotics increased, and flower-visitor species richness decreased. Yet, the decline in flower-visitor richness over the season was slowed in gardens with a relatively higher proportion of flowering exotic plants. This compensation was more positively linked to the proportion of exotic plant species than to the proportion of exotic flower cover. Plant-pollinator interaction networks were moderately specialized. Interactions were more complex in high summer, but interaction diversity, linkage density, and specialisation were not influenced by the proportion of exotic species. Thus, later in the season when few native plants flowered, exotic garden plants partly substituted for native flower resources without apparent influence on plant-pollinator network structure. Late-flowering garden plants support pollinator diversity in cities. If appropriately managed, and risk of naturalisation is minimized, late-flowering exotic plants may provide floral resources to support native pollinators when native plants are scarce.

Effectiveness of agri-environmental management on pollinators is moderated more by ecological contrast than by landscape structure or land-use intensity.

Agri-environment management (AEM) started in the 1980s in Europe to mitigate biodiversity decline, but the effectiveness of AEM has been questioned. We hypothesize that this is caused by a lack of a large enough ecological contrast between AEM and non-treated control sites. The effectiveness of AEM may be moderated by landscape structure and land-use intensity. Here, we examined the influence of local ecological contrast, landscape structure and regional land-use intensity on AEM effectiveness in a meta-analysis of 62 European pollinator studies. We found that ecological contrast was most important in determining the effectiveness of AEM, but landscape structure and regional land-use intensity played also a role. In conclusion, the most successful way to enhance AEM effectiveness for pollinators is to implement measures that result in a large ecological improvement at a local scale, which exhibit a strong contrast to conventional practices in simple landscapes of intensive land-use regions.

Tree diversity increases robustness of multi-trophic interactions.

Multi-trophic interactions maintain critical ecosystem functions. Biodiversity is declining globally, while responses of trophic interactions to biodiversity change are largely unclear. Thus, studying responses of multi-trophic interaction robustness to biodiversity change is crucial for understanding ecosystem functioning and persistence. We investigate plant-Hemiptera (antagonism) and Hemiptera-ant (mutualism) interaction networks in response to experimental manipulation of tree diversity. We show increased diversity at both higher trophic levels (Hemiptera and ants) and increased robustness through redundancy of lower level species of multi-trophic interactions when tree diversity increased. Hemiptera and ant diversity increased with tree diversity through non-additive diversity effects. Network analyses identified that tree diversity also increased the number of tree and Hemiptera species used by Hemiptera and ant species, and decreased the specialization on lower trophic level species in both mutualistic and antagonist interactions. Our results demonstrate that bottom-up effects of tree diversity ascend through trophic levels regardless of interaction type. Thus, local tree diversity is a key driver of multi-trophic community diversity and interaction robustness in forests.

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.

Chronic dryness and wetness and especially pulsed drought threaten a generalist arthropod herbivore.

Under climate change, both wetter and drier conditions, as well as an increase in extreme events like floods or droughts are projected for many areas. So far, studies only investigate the impact of drier or wetter conditions at a single stress severity level but do not consider how different intensities and types of changes affect insect herbivores feeding on stressed plants. Further, how effects of acute stress pulses differ from milder, chronic soil moisture stress is unclear. We investigated how changing soil moisture conditions affect a generalist insect herbivore feeding on grassland plants. We transplanted multi-species sections of grassland into pots and subjected them to different intensities and durations of flooding and drying stress. We compared effects of short, extreme drought and flooding pulses against the effects of milder, but chronic stress. Constantly drier conditions decreased plant and herbivore performance at all levels of stress severity. Severe permanent wetness did not affect plant growth, but decreased pupal weight (- 23%) and survival of larvae (- 34%). Extreme pulsed drought exacerbated negative effects of chronic drying, as most larvae died before they could benefit from rewetting plants after the drought (94% mortality). Pulsed flooding did not affect plants or larval development more than chronic severe wetness. Our findings imply that plant stress negatively affects generalist chewing herbivores, even with mixed diets. Both drier and severely wet, but not mildly wetter conditions, will reduce survival of some species. Especially, extreme droughts appear to have strong negative effects on generalist grassland herbivores.

Ants at Plant Wounds: A Little-Known Trophic Interaction with Evolutionary Implications for Ant-Plant Interactions.

Extrafloral nectaries (EFNs) allow plants to engage in mutualisms with ants, preventing herbivory in exchange for food. EFNs occur scattered throughout the plant phylogeny and likely evolved independent from herbivore-created wounds subsequently visited by ants collecting leaked sap. Records of wound-feeding ants are, however, anecdotal. By surveying 38,000 trees from 40 species, we conducted the first quantitative ecological study of this overlooked behavior. Ant-wound interactions were widespread (0.5% of tree individuals) and occurred on 23 tree species. Interaction networks were opportunistic, closely resembling ant-EFN networks. Fagaceae, a family lacking EFNs, was strongly overrepresented. For Fagaceae, ant occurrence at wounds correlated with species-level leaf damage, potentially indicating that wounds may attract mutualistic ants, which supports the hypothesis of ant-tended wounds as precursors of ant-EFN mutualisms. Given that herbivore wounds are common, wound sap as a steadily available food source might further help to explain the overwhelming abundance of ants in (sub)tropical forest canopies.

Tree species richness attenuates the positive relationship between mutualistic ant-hemipteran interactions and leaf chewer herbivory.

Interactions across trophic levels influence plant diversity effects on ecosystem functions, but the complexity of these interactions remains poorly explored. For example, the interplay between different interactions (e.g. mutualism, predation) might be an important moderator of biodiversity-ecosystem function relationships. We tested for relationships between trophobioses (facultative ant-hemipteran mutualism) and leaf chewer herbivory in a subtropical forest biodiversity experiment. We analysed trophobiosis and herbivory data of more than 10 000 trees along a tree species richness gradient. Against expectations, chewing damage was higher on trees with trophobioses. However, the net positive relationship between trophobioses and overall herbivory depended on tree species richness, being most pronounced at low richness. Our results point to indirect, positive effects of ant-tended sap suckers on leaf chewers, potentially by altering plant defences. Direct antagonistic relationships of trophobiotic ants and leaf-chewing herbivores-frequently reported to drive community-wide effects of trophobioses in other ecosystems-seemed less relevant. However, antagonistic interactions likely contributed to the attenuating effect of tree species richness, because trophobiotic ant and herbivore communities changed from monocultures to species-rich mixtures. Our findings, therefore, suggest that biodiversity loss might lead to complex changes in higher trophic level effects on ecosystem functions, mediated by both trophic and non-trophic interactions.

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.

Interannual variation in land-use intensity enhances grassland multidiversity.

Although temporal heterogeneity is a well-accepted driver of biodiversity, effects of interannual variation in land-use intensity (LUI) have not been addressed yet. Additionally, responses to land use can differ greatly among different organisms; therefore, overall effects of land-use on total local biodiversity are hardly known. To test for effects of LUI (quantified as the combined intensity of fertilization, grazing, and mowing) and interannual variation in LUI (SD in LUI across time), we introduce a unique measure of whole-ecosystem biodiversity, multidiversity. This synthesizes individual diversity measures across up to 49 taxonomic groups of plants, animals, fungi, and bacteria from 150 grasslands. Multidiversity declined with increasing LUI among grasslands, particularly for rarer species and aboveground organisms, whereas common species and belowground groups were less sensitive. However, a high level of interannual variation in LUI increased overall multidiversity at low LUI and was even more beneficial for rarer species because it slowed the rate at which the multidiversity of rare species declined with increasing LUI. In more intensively managed grasslands, the diversity of rarer species was, on average, 18% of the maximum diversity across all grasslands when LUI was static over time but increased to 31% of the maximum when LUI changed maximally over time. In addition to decreasing overall LUI, we suggest varying LUI across years as a complementary strategy to promote biodiversity conservation.

First mass development of Aedes albopictus (Diptera: Culicidae)-its surveillance and control in Germany.

The Asian tiger mosquito Aedes albopictus has undergone a dramatic expansion of its range in the last few decades. Since its first detection in 2007 in Germany at the motorway A5 coming from Italy via Switzerland to Germany, it has been continuously introduced by vehicles, most probably from Italy. After a hint from an alert gardener in an allotment garden area in Freiburg, Southwest Germany, in 2015, a surveillance programme was started focusing on the garden area and adjacent areas as well as most of the cemeteries as potential infestation areas. The surveillance programme confirmed a high infestation of the allotment garden. The container index (CI) exceeded almost 30% in August 2015. In lethal gravid Aedes traps (GATs) and BG-Sentinel traps, 4038 adults were caught. It could be proven that the Aedes population is more or less still spatially restricted to the allotment garden area which is adjacent to a train station where trucks from Novara, Italy, arrive loaded on trains. Outside the garden area, only a few breeding sites with developmental stages and adults were found within a radius of approximately 600 m from the highly infested garden area. It is most likely that Ae. albopictus females are constantly introduced as 'blind passengers' to Freiburg via trucks from Italy to Freiburg, Germany. After the first detection of the mass development of Ae. albopictus immediate and comprehensive control measures were initiated to reduce or even eliminate the Aedes population. Citizen awareness, especially of the gardeners, was increased by providing thorough information about the biology and control of Ae. albopictus. Beside environmental management, tablets based on Bacillus thuringiensis israelensis (Bti) were applied. The success of the control activities by the gardeners is reflected by the data gained during monthly inspection of the garden plots. The number of gardens without any container increased from 17% in July to 22% in August and 35% in September, 2015, resulting in a successful reduction of the Ae. albopictus population. The study underlines the importance of a comprehensive surveillance programme to assess the population density of Ae. albopictus as a basis for integrated control activities.

Tree phylogenetic diversity promotes host-parasitoid interactions.

Evidence from grassland experiments suggests that a plant community's phylogenetic diversity (PD) is a strong predictor of ecosystem processes, even stronger than species richness per se This has, however, never been extended to species-rich forests and host-parasitoid interactions. We used cavity-nesting Hymenoptera and their parasitoids collected in a subtropical forest as a model system to test whether hosts, parasitoids, and their interactions are influenced by tree PD and a comprehensive set of environmental variables, including tree species richness. Parasitism rate and parasitoid abundance were positively correlated with tree PD. All variables describing parasitoids decreased with elevation, and were, except parasitism rate, dependent on host abundance. Quantitative descriptors of host-parasitoid networks were independent of the environment. Our study indicates that host-parasitoid interactions in species-rich forests are related to the PD of the tree community, which influences parasitism rates through parasitoid abundance. We show that effects of tree community PD are much stronger than effects of tree species richness, can cascade to high trophic levels, and promote trophic interactions. As during habitat modification phylogenetic information is usually lost non-randomly, even species-rich habitats may not be able to continuously provide the ecosystem process parasitism if the evolutionarily most distinct plant lineages vanish.

A Global Synthesis of Jatropha Cultivation: Insights into Land Use Change and Management Practices.

Despite setbacks, interest in Jatropha cultivation remains high. This study addressed the question to what extent Jatropha cultivation has replaced specific vegetation and land use types and how the existing areas are managed. Major forms of land use change and management practices were identified based on cluster analysis of data from 106 interviewee's responses to a comprehensive global survey. Of the 1.04 × 10(6) ha cultivated with Jatropha in 2011 40% were established on land that was cleared of vegetation as a result of logging activities unrelated to Jatropha cultivation, 34% was defined as unused, and the remainder was attributable to areas previously used for crops or animal husbandry. With the exception of croplands, these areas were dominated (90-98%) by a few internationally active companies whose cultivation models were almost exclusively based on outgrower schemes. Management practices were largely extensive in nature (low mechanical input and infrequent use of fertilizers, pesticides and herbicides), and also dominated by large projects. Broad surveys, such as this, are useful in identifying general trends in this emerging global industry, but detailed case studies, particularly of large projects, are needed in order to draw more informed conclusions about the site-specific impacts of Jatropha cultivation.

Bottom-up effects of plant diversity on multitrophic interactions in a biodiversity experiment.

Biodiversity is rapidly declining, and this may negatively affect ecosystem processes, including economically important ecosystem services. Previous studies have shown that biodiversity has positive effects on organisms and processes across trophic levels. However, only a few studies have so far incorporated an explicit food-web perspective. In an eight-year biodiversity experiment, we studied an unprecedented range of above- and below-ground organisms and multitrophic interactions. A multitrophic data set originating from a single long-term experiment allows mechanistic insights that would not be gained from meta-analysis of different experiments. Here we show that plant diversity effects dampen with increasing trophic level and degree of omnivory. This was true both for abundance and species richness of organisms. Furthermore, we present comprehensive above-ground/below-ground biodiversity food webs. Both above ground and below ground, herbivores responded more strongly to changes in plant diversity than did carnivores or omnivores. Density and richness of carnivorous taxa was independent of vegetation structure. Below-ground responses to plant diversity were consistently weaker than above-ground responses. Responses to increasing plant diversity were generally positive, but were negative for biological invasion, pathogen infestation and hyperparasitism. Our results suggest that plant diversity has strong bottom-up effects on multitrophic interaction networks, with particularly strong effects on lower trophic levels. Effects on higher trophic levels are indirectly mediated through bottom-up trophic cascades.

Global malnutrition overlaps with pollinator-dependent micronutrient production.

Pollinators contribute around 10% of the economic value of crop production globally, but the contribution of these pollinators to human nutrition is potentially much higher. Crops vary in the degree to which they benefit from pollinators, and many of the most pollinator-dependent crops are also among the richest in micronutrients essential to human health. This study examines regional differences in the pollinator dependence of crop micronutrient content and reveals overlaps between this dependency and the severity of micronutrient deficiency in people around the world. As much as 50% of the production of plant-derived sources of vitamin A requires pollination throughout much of Southeast Asia, whereas other essential micronutrients such as iron and folate have lower dependencies, scattered throughout Africa, Asia and Central America. Micronutrient deficiencies are three times as likely to occur in areas of highest pollination dependence for vitamin A and iron, suggesting that disruptions in pollination could have serious implications for the accessibility of micronutrients for public health. These regions of high nutritional vulnerability are understudied in the pollination literature, and should be priority areas for research related to ecosystem services and human well-being.

Distribution of infectious and parasitic agents among three sentinel bee species across European agricultural landscapes.

Infectious and parasitic agents (IPAs) and their associated diseases are major environmental stressors that jeopardize bee health, both alone and in interaction with other stressors. Their impact on pollinator communities can be assessed by studying multiple sentinel bee species. Here, we analysed the field exposure of three sentinel managed bee species (Apis mellifera, Bombus terrestris and Osmia bicornis) to 11 IPAs (six RNA viruses, two bacteria, three microsporidia). The sentinel bees were deployed at 128 sites in eight European countries adjacent to either oilseed rape fields or apple orchards during crop bloom. Adult bees of each species were sampled before their placement and after crop bloom. The IPAs were detected and quantified using a harmonised, high-throughput and semi-automatized qPCR workflow. We describe differences among bee species in IPA profiles (richness, diversity, detection frequencies, loads and their change upon field exposure, and exposure risk), with no clear patterns related to the country or focal crop. Our results suggest that the most frequent IPAs in adult bees are more appropriate for assessing the bees' IPA exposure risk. We also report positive correlations of IPA loads supporting the potential IPA transmission among sentinels, suggesting careful consideration should be taken when introducing managed pollinators in ecologically sensitive environments.