Land-use change in the past 40 years explains shifts in arthropod community traits.

Understanding how anthropogenic activities induce changes in the functional traits of arthropod communities is critical to assessing their ecological consequences. However, we largely lack comprehensive assessments of the long-term impact of global-change drivers on the trait composition of arthropod communities across a large number of species and sites. This knowledge gap critically hampers our ability to predict human-driven impacts on communities and ecosystems. Here, we use a dataset of 1.73 million individuals from 877 species to study how four functionally important traits of carabid beetles and spiders (i.e. body size, duration of activity period, tolerance to drought, and dispersal capacity) have changed at the community level across ~40 years in different types of land use and as a consequence of land use changes (that is, urbanisation and loss of woody vegetation) at the landscape scale in Switzerland. The results show that the mean body size in carabid communities declined in all types of land use, with particularly stronger declines in croplands compared to forests. Furthermore, the length of the activity period and the tolerance to drought of spider communities decreased in most land use types. The average body size of carabid communities in landscapes with increased urbanisation in the last ~40 years tended to decrease. However, the length of the activity period, the tolerance to drought, and the dispersal capacity did not change significantly. Furthermore, urbanisation promoted increases in the average dispersal capacities of spider communities. Additionally, urbanisation favoured spider communities with larger body sizes and longer activity periods. The loss of woody areas at the landscape level was associated with trait shifts to carabid communities with larger body sizes, shorter activity periods, higher drought tolerances and strongly decreased dispersal capacities. Decreases in activity periods and dispersal capacities were also found in spider communities. Our study demonstrates that human-induced changes in land use alter key functional traits of carabid and spider communities in the long term. The detected trait shifts in arthropod communities likely have important consequences for their functional roles in ecosystems.

Quantifying potential trade-offs and win-wins between arthropod diversity and yield on cropland under agri-environment schemes-A meta-analysis.

In Europe, agri-environment schemes (AES) are a key instrument to combat the ongoing decline of farmland biodiversity. AES aim is to support biodiversity and maintain ecosystem services, such as pollination or pest control. To what extent AES affect crop yield is still poorly understood. We performed a systematic review, including hierarchical meta-analyses, to investigate potential trade-offs and win-wins between the effectiveness of AES for arthropod diversity and agricultural yield on European croplands. Altogether, we found 26 studies with a total of 125 data points that fulfilled our study inclusion criteria. From each study, we extracted data on biodiversity (arthropod species richness and abundance) and yield for fields with AES management and control fields without AES. The majority of the studies reported significantly higher species richness and abundance of arthropods (especially wild pollinators) in fields with AES (31 % increase), but yields were at the same time significantly lower on fields with AES compared to control fields (21 % decrease). Aside from the opportunity costs, AES that promote out-of-production elements (e.g. wildflower strips), supported biodiversity (29-32 % increase) without significantly compromising yield (2-5 % increase). Farmers can get an even higher yield in these situations than in current conventional agricultural production systems without AES. Thus, our study is useful to identify AES demonstrating benefits for arthropod biodiversity with negligible or relatively low costs regarding yield losses. Further optimization of the design and management of AES is needed to improve their effectiveness in promoting both biodiversity and minimizing crop yield losses.

Why has farming in Europe changed? A farmers' perspective on the development since the 1960s.

Farming in Europe has been the scene of several important socio-economic and environmental developments and crises throughout the last century. Therefore, an understanding of the historical driving forces of farm change helps identifying potentials for navigating future pathways of agricultural development. However, long-term driving forces have so far been studied, e.g. in anecdotal local case studies or in systematic literature reviews, which often lack context dependency. In this study, we bridged local and continental scales by conducting 123 oral history interviews (OHIs) with elderly farmers across 13 study sites in 10 European countries. We applied a driving forces framework to systematically analyse the OHIs. We find that the most prevalent driving forces were the introduction of new technologies, developments in agricultural markets that pushed farmers for farm size enlargement and technological optimisation, agricultural policies, but also cultural aspects such as cooperation and intergenerational arrangements. However, we find considerable heterogeneity in the specific influence of individual driving forces across the study sites, implying that generic assumptions about the dynamics and impacts of European agricultural change drivers hold limited explanatory power on the local scale. Our results suggest that site-specific factors and their historical development will need to be considered when addressing the future of agriculture in Europe in a scientific or policy context. Supplementary Information: The online version contains supplementary material available at 10.1007/s10113-023-02150-y.

Towards diverse agricultural land uses: socio-ecological implications of European agricultural pathways for a Swiss orchard region.

Diverse agricultural land uses are a typical feature of multifunctional landscapes. The uncertain change in the drivers of global land use, such as climate, market and policy technology and demography, challenges the long-term management of agricultural diversification. As these global drivers also affect smaller scales, it is important to capture the traits of regionally specific farm activities to facilitate adaptation to change. By downscaling European shared socioeconomic pathways (SSPs) for agricultural and food systems, combined with representative concentration pathways (RCP) to regionally specific, alternative socioeconomic and climate scenarios, the present study explores the major impacts of the drivers of global land use on regional agriculture by simulating farm-level decisions and identifies the socio-ecological implications for promoting diverse agricultural landscapes in 2050. A hilly orchard region in northern Switzerland was chosen as a case study to represent the multifunctional nature of Swiss agriculture. Results show that the different regionalised pathways lead to contrasting impacts on orchard meadows, production levels and biodiversity. Increased financial support for ecological measures, adequate farm labour supplies for more labour-intensive farming and consumer preferences that favour local farm produce can offset the negative impacts of climate change and commodity prices and contribute to agricultural diversification and farmland biodiversity. However, these conditions also caused a significant decline in farm production levels. This study suggests that considering a broader set of land use drivers beyond direct payments, while acknowledging potential trade-offs and diverse impacts across different farm types, is required to effectively manage and sustain diversified agricultural landscapes in the long run. Supplementary information: The online version contains supplementary material available at 10.1007/s10113-023-02092-5.

Validating Farmland Biodiversity Life Cycle Assessment at the Landscape Scale.

Life cycle assessment (LCA) aims at providing standardized evaluations of processes involving resource use, human health, and environmental consequences. Currently, spatial dependencies are most often neglected, though they are essential for impact categories like biodiversity. The "Swiss Agricultural Life Cycle Assessment for Biodiversity (SALCA-BD)" evaluates the impact of agricultural field management on 11 indicator species groups. We tested if its performance can be improved by accounting for the spatial context of the individual fields. We used high-resolution bird/butterfly point observations in two agricultural regions in Switzerland and built linear mixed models to compare SALCA-BD scores to the observed species richness at the field/landscape scale. We calculated a set of landscape metrics, tested their relationship with the landscape-model prediction errors, and then added all significant metrics as additional predictors to the landscape models. Our results show that field-scale SALCA-BD scores were significantly related to the observed field-scale richness for both indicator groups. However, the performance decreased when aggregated to the landscape scale, with high variability between regions. Adding specific landscape metrics improved the landscape model for birds but not for butterflies. Integrating the spatial context to LCA biodiversity assessments could provide moderate benefits, while its usefulness depends on the conditions of the respective assessment.

Different roles of concurring climate and regional land-use changes in past 40 years' insect trends.

Climate and land-use changes are main drivers of insect declines, but their combined effects have not yet been quantified over large spatiotemporal scales. We analysed changes in the distribution (mean occupancy of squares) of 390 insect species (butterflies, grasshoppers, dragonflies), using 1.45 million records from across bioclimatic gradients of Switzerland between 1980 and 2020. We found no overall decline, but strong increases and decreases in the distributions of different species. For species that showed strongest increases (25% quantile), the average proportion of occupied squares increased in 40 years by 0.128 (95% credible interval: 0.123-0.132), which equals an average increase in mean occupancy of 71.3% (95% CI: 67.4-75.1%) relative to their 40-year mean occupancy. For species that showed strongest declines (25% quantile), the average proportion decreased by 0.0660 (95% CI: 0.0613-0.0709), equalling an average decrease in mean occupancy of 58.3% (95% CI: 52.2-64.4%). Decreases were strongest for narrow-ranged, specialised, and cold-adapted species. Short-term distribution changes were associated to both climate changes and regional land-use changes. Moreover, interactive effects between climate and regional land-use changes confirm that the various drivers of global change can have even greater impacts on biodiversity in combination than alone. In contrast, 40-year distribution changes were not clearly related to regional land-use changes, potentially reflecting mixed changes in local land use after 1980. Climate warming however was strongly linked to 40-year changes, indicating its key role in driving insect trends of temperate regions in recent decades.

Farmer surveys in Europe suggest that specialized, intensive farms were more likely to perceive negative impacts from COVID-19.

It has been shown that the COVID-19 pandemic affected some agricultural systems more than others, and even within geographic regions, not all farms were affected to the same extent. To build resilience of agricultural systems to future shocks, it is key to understand which farms were affected and why. In this study, we examined farmers' perceived robustness to COVID-19, a key resilience capacity. We conducted standardized farmer interviews (n = 257) in 15 case study areas across Europe, covering a large range of socio-ecological contexts and farm types. Interviews targeted perceived livelihood impacts of the COVID-19 pandemic on productivity, sales, price, labor availability, and supply chains in 2020, as well as farm(er) characteristics and farm management. Our study corroborates earlier evidence that most farms were not or only slightly affected by the first wave(s) of the pandemic in 2020, and that impacts varied widely by study region. However, a significant minority of farmers across Europe reported that the pandemic was "the worst crisis in a lifetime" (3%) or "the worst crisis in a decade" (7%). Statistical analysis showed that more specialized and intensive farms were more likely to have perceived negative impacts. From a societal perspective, this suggests that highly specialized, intensive farms face higher vulnerability to shocks that affect regional to global supply chains. Supporting farmers in the diversification of their production systems while decreasing dependence on service suppliers and supply chain actors may increase their robustness to future disruptions. Supplementary Information: The online version contains supplementary material available at 10.1007/s13593-022-00820-5.

Effects of temporal floral resource availability and non-crop habitats on broad bean pollination.

Context: Flowering plants can enhance wild insect populations and their pollination services to crops in agricultural landscapes, especially when they flower before the focal crop. However, characterizing the temporal availability of specific floral resources is a challenge. Objectives: Developing an index for the availability of floral resources at the landscape scale according to the specific use by a pollinator. Investigating whether detailed and temporally-resolved floral resource maps predict pollination success of broad bean better than land cover maps. Methods: We mapped plant species used as pollen source by bumblebees in 24 agricultural landscapes and developed an index of floral resource availability for different times of the flowering season. To measure pollination success, patches of broad bean (Vicia faba), a plant typically pollinated by bumblebees, were exposed in the center of selected landscapes. Results: Higher floral resource availability before bean flowering led to enhanced seed set. Floral resource availability synchronous to broad bean flowering had no effect. Seed set was somewhat better explained by land cover maps than by floral resource availability, increasing with urban area and declining with the cover of arable land. Conclusions: The timing of alternative floral resource availability is important for crop pollination. The higher explanation of pollination success by land cover maps than by floral resource availability indicates that additional factors such as habitat disturbance and nesting sites play a role in pollination. Enhancing non-crop woody plants in agricultural landscapes as pollen sources may ensure higher levels of crop pollination by wild pollinators such as bumblebees. Supplementary information: The online version contains supplementary material available at 10.1007/s10980-022-01448-2.

Comparing floral resource maps and land cover maps to predict predators and aphid suppression on field bean.

CONTEXT: Predatory insects contribute to the natural control of agricultural pests, but also use plant pollen or nectar as supplementary food resources. Resource maps have been proposed as an alternative to land cover maps for prediction of beneficial insects. OBJECTIVES: We aimed at predicting the abundance of crop pest predating insects and the pest control service they provide with both, detailed flower resource maps and land cover maps. METHODS: We selected 19 landscapes of 500 m radius and mapped them with both approaches. In the centres of the landscapes, aphid predators - hoverflies (Diptera: Syrphidae), ladybeetles (Coleoptera: Coccinellidae) and lacewings (Neuroptera: Chrysopidae) - were surveyed in experimentally established faba bean phytometers (Vicia faba L. Var. Sutton Dwarf) and their control of introduced black bean aphids (Aphis fabae Scop.) was recorded. RESULTS: Landscapes with higher proportions of forest edge as derived from land cover maps supported higher abundance of aphid predators, and high densities of aphid predators reduced aphid infestation on faba bean. Floral resource maps did not significantly predict predator abundance or aphid control services. CONCLUSIONS: Land cover maps allowed to relate landscape composition with predator abundance, showing positive effects of forest edges. Floral resource maps may have failed to better predict predators because other resources such as overwintering sites or alternative prey potentially play a more important role than floral resources. More research is needed to further improve our understanding of resource requirements beyond floral resource estimations and our understanding of their role for aphid predators at the landscape scale. SUPPLEMENTARY INFORMATION: The online version contains supplementary material available at 10.1007/s10980-021-01361-0.

An approach for comparing agricultural development to societal visions.

There is broad agreement that agriculture has to become more sustainable in order to provide enough affordable, healthy food at minimal environmental and social costs. But what is "more sustainable"? More often than not, different stakeholders have opposing opinions on what a more sustainable future should look like. This normative dimension is rarely explicitly addressed in sustainability assessments. In this study, we present an approach to assess the sustainability of agricultural development that explicitly accounts for the normative dimension by comparing observed development with various societal visions. We illustrate the approach by analyzing farm- and landscape-scale development as well as sustainability outcomes in a Swiss case study landscape. Observed changes were juxtaposed with desired changes by Avenir Suisse, a liberal think tank representing free-market interests; the Swiss Farmers Association, representing a conservative force; and Landwirtschaft mit Zukunft, an exponent of the Swiss agroecological movement. Overall, the observed developments aligned most closely with desired developments of the liberal think-tank (72%). Farmer interviews revealed that in the case study area farms increased in size (+ 57%) and became more specialized and more productive (+ 223%) over the past 20 years. In addition, interpretation of aerial photographs indicated that farming became more rationalized at the landscape level, with increasing field sizes (+ 34%) and removal of solitary field trees (- 18%). The case study example highlights the varying degrees to which current developments in agriculture align with societal visions. By using societal visions as benchmarks to track the progress of agricultural development, while explicitly addressing their narratives and respective systems of values and norms, this approach offers opportunities to inform also the wider public on the extent to which current developments are consistent with different visions. This could help identify mismatches between desired and actual development and pave the way for designing new policies. SUPPLEMENTARY INFORMATION: The online version contains supplementary material available at 10.1007/s13593-021-00739-3.

Dependence of the frequency distribution around a sphere on the voxel orientation.

Microscopically small magnetic field inhomogeneities within an external static magnetic field cause a free induction decay in magnetic resonance imaging that generally exhibits two transverse components that are usually summarized to a complex entity. The Fourier transform of the complex-valued free induction decay is the purely real and positive-valued frequency distribution which allows an easy interpretation of the underlying dephasing mechanism. Typically, the frequency distribution inside a cubic voxel as caused by a spherical magnetic field inhomogeneity is determined by a histogram technique in terms of subdivision of the whole voxel into smaller subvoxels. A faster and more accurate computation is achieved by analytical expressions for the frequency distribution that are derived in this work. In contrast to the usually assumed simplified case of a spherical voxel, we also consider the tilt angles of the cubic voxel to the external magnetic field. The typical asymmetric form of the frequency distribution is reproduced and analyzed for the more realistic case of a cubic voxel. We observe a splitting of frequency distribution peaks for increasing tilt of the cubic voxel against the direction of the external magnetic field in analogy to the case for dephasing around cylindrical, vessel-like objects inside cubic voxels. These results are of value, e.g., for the analysis of susceptibility-weighted images or in quantitative susceptibility imaging since the reconstruction of these images is performed in cubic-shaped voxels.

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.

Insights into aphid prey consumption by ladybirds: Optimising field sampling methods and primer design for high throughput sequencing.

Elucidating the diets of insect predators is important in basic and applied ecology, such as for improving the effectiveness of conservation biological control measures to promote natural enemies of crop pests. Here, we investigated the aphid diet of two common aphid predators in Central European agroecosystems, the native Coccinella septempunctata (Linnaeus) and the invasive Harmonia axyridis (Pallas; Coleoptera: Coccinellidae) by means of high throughput sequencing (HTS). For acquiring insights into diets of mobile flying insects at landscape scale minimizing trapping bias is important, which imposes methodological challenges for HTS. We therefore assessed the suitability of three field sampling methods (sticky traps, pan traps and hand-collection) as well as new aphid primers for identifying aphid prey consumption by coccinellids through HTS. The new aphid primers facilitate identification to species level in 75% of the European aphid genera investigated. Aphid primer specificity was high in silico and in vitro but low in environmental samples with the methods used, although this could be improved in future studies. For insect trapping we conclude that sticky traps are a suitable method in terms of minimizing sampling bias, contamination risk and trapping success, but compromise on DNA-recovery rate. The aphid diets of both field-captured ladybird species were dominated by Microlophium carnosum, the common nettle aphid. Another common prey was Sitobion avenae (cereal aphid), which got more often detected in C. septempunctata compared to H. axyridis. Around one third of the recovered aphid taxa were common crop pests. We conclude that sampling methodologies need constant revision but that our improved aphid primers offer currently one of the best solutions for broad screenings of coccinellid predation on aphids.

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.

Characterization factors for land use impacts on biodiversity in life cycle assessment based on direct measures of plant species richness in European farmland in the 'Temperate Broadleaf and Mixed Forest' biome.

Life Cycle Assessment (LCA) is a widely used tool to assess environmental sustainability of products. The LCA should optimally cover the most important environmental impact categories such as climate change, eutrophication and biodiversity. However, impacts on biodiversity are seldom included in LCAs due to methodological limitations and lack of appropriate characterization factors. When assessing organic agricultural products the omission of biodiversity in LCA is problematic, because organic systems are characterized by higher species richness at field level compared to the conventional systems. Thus, there is a need for characterization factors to estimate land use impacts on biodiversity in life cycle assessment that are able to distinguish between organic and conventional agricultural land use that can be used to supplement and validate the few currently suggested characterization factors. Based on a unique dataset derived from field recording of plant species diversity in farmland across six European countries, the present study provides new midpoint occupation Characterization Factors (CF) expressing the Potentially Disappeared Fraction (PDF) to estimate land use impacts on biodiversity in the 'Temperate Broadleaf and Mixed Forest' biome in Europe. The method is based on calculation of plant species on randomly selected test sites in the biome and enables the calculation of characterization factors that are sensitive to particular types of management. While species richness differs between countries, the calculated CFs are able to distinguish between different land use types (pastures (monocotyledons or mixed), arable land and hedges) and management practices (organic or conventional production systems) across countries. The new occupation CFs can be used to supplement or validate the few current CF's and can be applied in LCAs of agricultural products to assess land use impacts on species richness in the 'Temperate Broadleaf and Mixed Forest' biome.

Farmland biodiversity and agricultural management on 237 farms in 13 European and two African regions.

Farmland is a major land cover type in Europe and Africa and provides habitat for numerous species. The severe decline in farmland biodiversity of the last decades has been attributed to changes in farming practices, and organic and low-input farming are assumed to mitigate detrimental effects of agricultural intensification on biodiversity. Since the farm enterprise is the primary unit of agricultural decision making, management-related effects at the field scale need to be assessed at the farm level. Therefore, in this study, data were collected on habitat characteristics, vascular plant, earthworm, spider, and bee communities and on the corresponding agricultural management in 237 farms in 13 European and two African regions. In 15 environmental and agricultural homogeneous regions, 6-20 farms with the same farm type (e.g., arable crops, grassland, or specific permanent crops) were selected. If available, an equal number of organic and non-organic farms were randomly selected. Alternatively, farms were sampled along a gradient of management intensity. For all selected farms, the entire farmed area was mapped, which resulted in total in the mapping of 11 338 units attributed to 194 standardized habitat types, provided together with additional descriptors. On each farm, one site per available habitat type was randomly selected for species diversity investigations. Species were sampled on 2115 sites and identified to the species level by expert taxonomists. Species lists and abundance estimates are provided for each site and sampling date (one date for plants and earthworms, three dates for spiders and bees). In addition, farmers provided information about their management practices in face-to-face interviews following a standardized questionnaire. Farm management indicators for each farm are available (e.g., nitrogen input, pesticide applications, or energy input). Analyses revealed a positive effect of unproductive areas and a negative effect of intensive management on biodiversity. Communities of the four taxonomic groups strongly differed in their response to habitat characteristics, agricultural management, and regional circumstances. The data has potential for further insights into interactions of farmland biodiversity and agricultural management at site, farm, and regional scale.

Predicting bee community responses to land-use changes: Effects of geographic and taxonomic biases.

Land-use change and intensification threaten bee populations worldwide, imperilling pollination services. Global models are needed to better characterise, project, and mitigate bees' responses to these human impacts. The available data are, however, geographically and taxonomically unrepresentative; most data are from North America and Western Europe, overrepresenting bumblebees and raising concerns that model results may not be generalizable to other regions and taxa. To assess whether the geographic and taxonomic biases of data could undermine effectiveness of models for conservation policy, we have collated from the published literature a global dataset of bee diversity at sites facing land-use change and intensification, and assess whether bee responses to these pressures vary across 11 regions (Western, Northern, Eastern and Southern Europe; North, Central and South America; Australia and New Zealand; South East Asia; Middle and Southern Africa) and between bumblebees and other bees. Our analyses highlight strong regionally-based responses of total abundance, species richness and Simpson's diversity to land use, caused by variation in the sensitivity of species and potentially in the nature of threats. These results suggest that global extrapolation of models based on geographically and taxonomically restricted data may underestimate the true uncertainty, increasing the risk of ecological surprises.

State-of-the-art practices in farmland biodiversity monitoring for North America and Europe.

Policy makers and farmers need to know the status of farmland biodiversity in order to meet conservation goals and evaluate management options. Based on a review of 11 monitoring programs in Europe and North America and on related literature, we identify the design choices or attributes of a program that balance monitoring costs and usefulness for stakeholders. A useful program monitors habitats, vascular plants, and possibly faunal groups (ecosystem service providers, charismatic species) using a stratified random sample of the agricultural landscape, including marginal and intensive regions. The size of landscape samples varies with the grain of the agricultural landscape; for example, samples are smaller in Europe and larger in North America. Raw data are collected in a rolling survey, which distributes sampling over several years. Sufficient practical experience is now available to implement broad monitoring schemes on both continents. Technological developments in remote sensing, metagenomics, and social media may offer new opportunities for affordable farmland biodiversity monitoring and help to lower the overall costs of monitoring programs.

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.