A global quantitative synthesis of local and landscape effects on wild bee pollinators in agroecosystems

Bees provide essential pollination services that are potentially affected both by local farm management and the surrounding landscape. To better understand these different factors, we modelled the relative effects of landscape composition (nesting and floral resources within foraging distances), landscape configuration (patch shape, interpatch connectivity and habitat aggregation) and farm management (organic vs. conventional and local‐scale field diversity), and their interactions, on wild bee abundance and richness for 39 crop systems globally. Bee abundance and richness were higher in diversified and organic fields and in landscapes comprising more high‐quality habitats; bee richness on conventional fields with low diversity benefited most from high‐quality surrounding land cover. Landscape configuration effects were weak. Bee responses varied slightly by biome. Our synthesis reveals that pollinator persistence will depend on both the maintenance of high‐quality habitats around farms and on local management practices that may offset impacts of intensive monoculture agriculture.     

Humans,bees, and pollination services in the city: the case of Chicago,IL (USA)

Despite the global trend in urbanization, little is known about patterns of biodiversity or provisioning of ecosystem services in urban areas. Bee communities and the pollination services they provide are important in cities, both for small-scale urban agriculture and native gardens. To better understand this important ecological issue, we examined bee communities, their response to novel floral resources, and their potential to provide pollination services in 25 neighborhoods across Chicago, IL (USA). In these neighborhoods, we evaluated how local floral resources, socioeconomic factors, and surrounding land cover affected abundance, richness, and community composition of bees active in summer. We also quantified species-specific body pollen loads and visitation frequencies to potted flowering purple coneflower plants (Echinacea purpurea) to estimate potential pollination services in each neighborhood. We documented 37 bee species and 79 flowering plant genera across all neighborhoods, with 8 bee species and 14 flowering plant genera observed on average along each neighborhood block. We found that both bee abundance and richness increased in neighborhoods with higher human population density, as did visitation to purple coneflower flower heads. In more densely populated neighborhoods, bee communities shifted to a suite of species that carry more pollen and are more active pollinators in this system, including the European honey bee (Apis mellifera) and native species such as Agapostemon virescens. More densely populated neighborhoods also had a greater diversity of flowering plants, suggesting that the positive relationship between people and bees was mediated by the effect of people on floral resources. Other environmental variables that were important for bee communities included the amount of grass/herbaceous cover and solar radiation in the surrounding area. Our results indicate that bee communities and pollination services can be maintained in dense urban neighborhoods with single-family and multi-family homes, as long as those neighborhoods contain diverse and abundant floral resources.     

Response of wild bee diversity,abundance, and functional traits to vineyard inter‐row management intensity and landscape diversity across Europe

Agricultural intensification is a major driver of wild bee decline. Vineyards may be inhabited by plant and animal species, especially when the inter‐row space is vegetated with spontaneous vegetation or cover crops. Wild bees depend on floral resources and suitable nesting sites which may be found in vineyard inter‐rows or in viticultural landscapes. Inter‐row vegetation is managed by mulching, tillage, and/or herbicide application and results in habitat degradation when applied intensively. Here, we hypothesize that lower vegetation management intensities, higher floral resources, and landscape diversity affect wild bee diversity and abundance dependent on their functional traits. We sampled wild bees semi‐quantitatively in 63 vineyards representing different vegetation management intensities across Europe in 2016. A proxy for floral resource availability was based on visual flower cover estimations. Management intensity was assessed by vegetation cover (%) twice a year per vineyard. The Shannon Landscape Diversity Index was used as a proxy for landscape diversity within a 750 m radius around each vineyard center point. Wild bee communities were clustered by country. At the country level, between 20 and 64 wild bee species were identified. Increased floral resource availability and extensive vegetation management both affected wild bee diversity and abundance in vineyards strongly positively. Increased landscape diversity had a small positive effect on wild bee diversity but compensated for the negative effect of low floral resource availability by increasing eusocial bee abundance. We conclude that wild bee diversity and abundance in vineyards is efficiently promoted by increasing floral resources and reducing vegetation management frequency. High landscape diversity further compensates for low floral resources in vineyards and increases pollinating insect abundance in viticulture landscapes.     

Contrasting responses of plants and pollinators to woodland disturbance

Preserving species diversity is critical to ensure ecosystem functioning; however, different components of diversity might respond to human disturbance in different ways. Similarly, trophic levels might have uncoupled responses to the same disturbance, thus ameliorating or aggravating the persistence of ecological communities. In this study, we analysed how the density, richness and evenness of flowers and pollinators respond to four levels of woodland thinning intensity (0, 30, 50 and 70% of woodland basal area removed) over 2 years in three contrasting sites. We found a mismatch in the response of flowers and pollinators to thinning. Flower density and richness had disparate responses, depending on the site and year, while evenness did not change with thinning. In contrast, pollinator density and richness, but not evenness, consistently increased with thinning among years and sites. These results suggest that thinning has a great influence on pollinators through changes in abiotic conditions and, perhaps, flower attractiveness rather than through small‐scale changes in flower density and richness. At the site where tree flowers were absent, bee pollinator community composition was impoverished, suggesting that trees provide important floral resources to pollinators. Our findings indicate that disturbance may diminish local plant abundance and richness, but pollinator abundance and richness are enhanced after intense thinning at small scales.     

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

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

Wild bee species abundance and richness across an urban–rural gradient

Long-term and widespread monitoring programs are essential to understanding the role of human-dominated landscapes in supporting wild bee populations. Urbanization results in increased impervious surfaces throughout the landscape, fragmentation of green space, and a loss of naturally occurring floral vegetation. All of these changes have a negative impact on pollinator diversity. The objective of this study was to assess the abundance and richness of wild bee species throughout a small city in northwest Pennsylvania and identify how management of land throughout the city may influence bee communities. Seventeen sites across a land use gradient, moving from areas with large open spaces and mainly permeable surfaces, to sites in the city center consisting of mainly impermeable surfaces, were sampled over a 2-year period. During this time, 106 known species were identified with four state records and 1 undescribed species. Bee species richness was greatest at sites with the largest amount of permeable surface and naturally-occurring, native vegetation. Richness decreased on the college campus and city center where landscapes were highly managed and impermeable surfaces were most abundant. While floral richness was not related to bee abundance and richness, the number of open blooms near traps did have a positive impact on bee species richness. Overall, this survey revealed considerable richness never before recorded for northwest Pennsylvania, suggesting the importance of conservation management in homeowner and community yard space.     

Influences on the structure of suburban ant (Hymenoptera: Formicidae) communities and the abundance of Tapinoma sessile

Urbanization can alter the organization of ant communities and affect populations of urban pest ants. In this study, we sampled ant communities in urban and suburban yards to understand the habitat factors that shape these communities and influence the abundance of a common pest species, Tapinoma sessile (Say). We used pitfall traps to sample ant communities and a combination of pitfall traps and baiting to collect T. sessile at 24 sites in Knoxville, TN. In total, we collected 46 ant species. Ant species richness ranged from seven to 24 species per yard. Ant species richness tended to be lowest near houses, whereas T. sessile abundance was highest near houses. The best predictors of ant species richness in yards were canopy cover and presence of leaf litter: ant species richness peaked at mid-levels of canopy cover and was negatively correlated with the presence of leaf litter. Tapinoma sessile abundance increased with presence of logs, boards, or landscaping timbers and leaf litter in yards. Our results indicate that ant communities and the abundance of particular pest species in these urban and suburban landscapes are shaped by many of the same factors that structure ant communities in less anthropogenically disturbed environments.     

Spatial variability in a plant–pollinator community across a continuous habitat: high heterogeneity in the face of apparent uniformity

Large‐scale spatial variability in plant–pollinator communities (e.g. along geographic gradients, across different landscapes) is relatively well understood. However, we know much less about how these communities vary at small scales within a uniform landscape. Plants are sessile and highly sensitive to microhabitat conditions, whereas pollinators are highly mobile and, for the most part, display generalist feeding habits. Therefore, we expect plants to show greater spatial variability than pollinators. We analysed the spatial heterogeneity of a community of flowering plants and their pollinators in 40 plots across a 40‐km² area within an uninterrupted Mediterranean scrubland. We recorded 3577 pollinator visits to 49 plant species. The pollinator community (170 species) was strongly dominated by honey bees (71.8% of the visits recorded). Flower and pollinator communities showed similar beta‐diversity, indicating that spatial variability was similar in the two groups. We used path analysis to establish the direct and indirect effects of flower community distribution and honey bee visitation rate (a measure of the use of floral resources by this species) on the spatial distribution of the pollinator community. Wild pollinator abundance was positively related to flower abundance. Wild pollinator visitation rate was negatively related to flower abundance, suggesting that floral resources were not limiting. Pollinator and flower richness were positively related. Pollinator species composition was weakly related to flower species composition, reflecting the generalist nature of flower–pollinator interactions and the opportunistic nature of pollinator flower choices. Honey bee visitation rate did not affect the distribution of the wild pollinator community. Overall, we show that, in spite of the apparent physiognomic uniformity, both flowers and pollinators display high levels of heterogeneity, resulting in a mosaic of idiosyncratic local communities. Our results provide a measure of the background of intrinsic heterogeneity within a uniform habitat, with potential consequences on low‐scale ecosystem function and microevolutionary patterns.     

Bee Preference for Native versus Exotic Plants in Restored Agricultural Hedgerows

Habitat restoration to promote wild pollinator populations is becoming increasingly common in agricultural lands. Yet, little is known about how wild bees, globally the most important wild pollinators, use resources in restored habitats. We compared bee use of native and exotic plants in two types of restored native plant hedgerows: mature hedgerows (>10 years from establishment) designed for natural enemy enhancement and new hedgerows (≤2 years from establishment) designed to enhance bee populations. Bees were collected from flowers using timed aerial netting and flowering plant cover was estimated by species using cover classes. At mature hedgerow sites, wild bee abundance, richness, and diversity were greater on native plants than exotic plants. At new sites, where native plants were small and had limited floral display, abundance of bees was greater on native plants than exotic plants; but, controlling for floral cover, there was no difference in bee diversity and richness between the two plant types. At both mature and new hedgerows, wild bees preferred to forage from native plants than exotic plants. Honey bees, which were from managed colonies, also preferred native plants at mature hedgerow sites but exhibited no preference at new sites. Our study shows that wild bees, and managed bees in some cases, prefer to forage on native plants in hedgerows over co‐occurring weedy, exotic plants. Semi‐quantitative ranking identified which native plants were most preferred. Hedgerow restoration with native plants may help enhance wild bee abundance and diversity, and maintain honey bee health, in agricultural areas.     

Interaction effects between local flower richness and distance to natural woodland on pest and beneficial insects in apple orchards

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Tree diversity drives diversity of arthropod herbivores,but successional stage mediates detritivores

The high tree diversity of subtropical forests is linked to the biodiversity of other trophic levels. Disentangling the effects of tree species richness and composition, forest age, and stand structure on higher trophic levels in a forest landscape is important for understanding the factors that promote biodiversity and ecosystem functioning. Using a plot network spanning gradients of tree diversity and secondary succession in subtropical forest, we tested the effects of tree community characteristics (species richness and composition) and forest succession (stand age) on arthropod community characteristics (morphotype diversity, abundance and composition) of four arthropod functional groups. We posit that these gradients differentially affect the arthropod functional groups, which mediates the diversity, composition, and abundance of arthropods in subtropical forests. We found that herbivore richness was positively related to tree species richness. Furthermore, the composition of herbivore communities was associated with tree species composition. In contrast, detritivore richness and composition was associated with stand age instead of tree diversity. Predator and pollinator richness and abundance were not strongly related to either gradient, although positive trends with tree species richness were found for predators. The weaker effect of tree diversity on predators suggests a cascading diversity effect from trees to herbivores to predators. Our results suggest that arthropod diversity in a subtropical forest reflects the net outcome of complex interactions among variables associated with tree diversity and stand age. Despite this complexity, there are clear linkages between the overall richness and composition of tree and arthropod communities, in particular herbivores, demonstrating that these trophic levels directly impact each other.     

Effects of landscape cover and local habitat characteristics on visiting bees in tropical orchards

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Canopy thinning,not agricultural history,determines early responses of wild bees to longleaf pine savanna restoration

Longleaf pine savannas are highly threatened, fire‐maintained ecosystems unique to the southeastern United States. Fire suppression and conversion to agriculture have strongly affected this ecosystem, altering overstory canopies, understory plant communities, and animal populations. Tree thinning to reinstate open canopies can benefit understory plant diversity, but effects on animal communities are less well understood. Moreover, agricultural land‐use legacies can have long‐lasting impacts on plant communities, but their effects on animal communities either alone or through interactions with restoration are unclear. Resolving these impacts is important due to the conservation potential of fire‐suppressed and post‐agricultural longleaf savannas. We evaluated how historical agricultural land use and canopy thinning affect the diversity and abundance of wild bees in longleaf pine savannas. We employed a replicated, large‐scale factorial block experiment in South Carolina, where canopy thinning was applied to longleaf pine savannas that were either post‐agricultural or remnant (no agricultural history). Bees were sampled using elevated bee bowls. In the second growing season after restoration, thinned plots supported a greater bee abundance and bee community richness. Additionally, restored plots had altered wild bee community composition when compared to unthinned plots, indicating that reduction of canopy cover by the thinning treatment best predicted wild bee diversity and composition. Conversely, we found little evidence for differences between sites with or without historical agricultural land use. Some abundant Lasioglossum species were the most sensitive to habitat changes. Our results highlight how restoration practices that reduce canopy cover in fire‐suppressed savannas can have rapid benefits for wild bee communities.     

Arid grassland bee communities: associated environmental variables and responses to restoration

In recent years restoration project efforts in arid grasslands of the Pacific Northwest have increased; however, little is known about the bee communities in these areas or how restoration affects them. Native bees provide an essential ecosystem service through pollination of crops and native plants and understanding their response to restoration is a high priority. To address this issue, we conducted a three‐year study in an arid bunchgrass prairie with three objectives: (1) describe the bee community of this unique grassland type and its temporal variability; (2) investigate environmental variables influencing the community; and (3) examine effects of restoration on the community. We identified 62 bee species and found strong seasonal and inter‐annual variation in bee abundance, richness, diversity, and species composition. Unexpectedly, these temporal trends did not correspond with patterns in floral resources; however, several variables were associated with variation in bee abundance, richness, and diversity among sites. Sites with high levels of litter cover had more bees, while sites with taller vegetation or more blooming flowers had greater species richness but lower diversity. We found no detectable effect of restoration on bee abundance, richness, diversity, or composition. Species composition at native sites differed from those in actively and passively restored sites, which did not differ from each other. Restored sites also had fewer flowers and differing floral composition relative to native sites. These results suggest that if grassland restoration is to benefit bees, efforts should focus on both expanding floral resources and enhancing variables that influence nesting habitat.     

Ecological outcomes of agroforests and restoration 15 years after planting

Large‐scale forest restoration relies on approaches that are cost‐effective and economically attractive to farmers, and in this context agroforestry systems may be a valuable option. Here, we compared ecological outcomes among (1) 12–15‐year‐old coffee agroforests established with several native shade trees, (2) 12–15‐year‐old high‐diversity restoration plantations, and (3) reference old‐growth forests, within a landscape restoration project in the Pontal do Paranapanema region, in the Atlantic Forest of southeastern Brazil. We compared the aboveground biomass, canopy cover, and abundance, richness, and composition of trees, and the regenerating saplings in the three forest types. In addition, we investigated the landscape drivers of natural regeneration in the restoration plantations and coffee agroforests. Reference forests had a higher abundance of trees and regenerating saplings, but had similar levels of species richness compared to coffee agroforests. High‐diversity agroforests and restoration plantations did not differ in tree abundance. However, compared to restoration plantations, agroforests showed higher abundance and species richness of regenerating saplings, a higher proportion of animal‐dispersed species, and higher canopy cover. The abundance of regenerating saplings declined with increasing density of coffee plants, thus indicating a potential trade‐off between productivity and ecological benefits. High‐diversity coffee agroforests provide a cost‐effective and ecologically viable alternative to high‐diversity native tree plantations for large‐scale forest restoration within agricultural landscapes managed by local communities, and should be included as part of the portfolio of reforestation options used to promote the global agenda on forest and landscape restoration.     

Urbanization increases floral specialization of pollinators

Understanding how urbanization alters functional interactions among pollinators and plants is critically important given increasing anthropogenic land use and declines in pollinator populations. Pollinators often exhibit short‐term specialization and visit plants of the same species during one foraging trip. This facilitates plant receipt of conspecific pollen—pollen on a pollinator that is the same species as the plant on which the pollinator was foraging. Conspecific pollen receipt facilitates plant reproductive success and is thus important to plant and pollinator persistence. We investigated how urbanization affects short‐term specialization of insect pollinators by examining pollen loads on insects’ bodies and identifying the number and species of pollen grains on insects caught in urban habitat fragments and natural areas. We assessed possible drivers of differences between urban and natural areas, including frequency dependence in foraging, species richness and diversity of the plant and pollinator communities, floral abundance, and the presence of invasive plant species. Pollinators were more specialized in urban fragments than in natural areas, despite no differences in the species richness of plant communities across site types. These differences were likely driven by higher specialization of common pollinators, which were more abundant in urban sites. In addition, pollinators preferred to forage on invasive plants at urban sites and native plants at natural sites. Our findings reveal indirect effects of urbanization on pollinator fidelity to individual plant species and have implications for the maintenance of plant species diversity in small habitat fragments. Higher preference of pollinators for invasive plants at urban sites suggests that native species may receive fewer visits by pollinators. Therefore, native plant species diversity may decline in urban sites without continued augmentation of urban flora or removal of invasive species.     

Tree diversity promotes predatory wasps and parasitoids but not pollinator bees in a subtropical experimental forest

From regional to global scales, anthropogenic environmental change is causing biodiversity loss and reducing ecosystem functionality. Previous studies have investigated the relationship between plant diversity and functional insect communities in temperate and also in tropical grasslands and forests. However, few studies have explored these dynamics in subtropical forests. Here, cavity-nesting Hymenoptera and associated parasitoids were collected across a controlled tree diversity experiment in subtropical China to test how predatory wasps, bees and parasitoids respond to tree species richness. Abundance and species richness of predatory wasps and parasitoids were positively correlated with tree species richness, while bee abundance and bee species richness were unrelated to tree species richness. Our results indicate that tree species richness increases the abundance and species richness of important communities such as predators and parasitoids. Moreover, the results highlight the importance of subtropical forests in maintaining abundance and species richness of key functional insect groups.     

Almond orchards with living ground cover host more wild insect pollinators

Wild pollinators are becoming more valuable to global agriculture as the commercial honeybee industry is increasingly affected by disease and other stressors. Perennial tree crops are particularly reliant on insect pollination, and are often pollen limited. Research on how different tree crop production systems influence the richness and abundance of wild pollinators is, however, limited. We investigated, for the first time, the richness and abundance of potential wild pollinators in commercial temperate almond orchards in Australia, and compared them to potential pollinator communities in proximate native vegetation. We quantified ground cover variables at each site and assessed the value of ground cover on the richness and abundance of potential wild pollinators in commercial almond systems focussing on three common taxa: bees, wasps and flies. More insects were caught in orchards with living ground cover than in native vegetation or orchards without ground cover, although overall species richness was highest in native vegetation. Percent ground cover was positively associated with wasp richness and abundance, and native bee richness, but flies showed no association with ground cover. The strongest positive relationship was between native bee abundance and the richness of ground cover plants. Our results suggest that maintaining living ground cover within commercial almond orchards could provide habitat and resources for potential wild pollinators, particularly native bees. These insects have the potential to provide a valuable ecosystem service to pollinator-dependent crops such as almond.     

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

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

Habitat type plays a greater role than livestock grazing in structuring shrubsteppe plant–pollinator communities

Livestock grazing is a widespread grassland disturbance and can negatively impact biodiversity. Pollinators constitute a vital component of grassland ecosystems, but the impact of grazing on pollinator diversity has seldom been evaluated in North America. We assessed vegetation structure, and pollinator and flowering plant abundance, richness, diversity, and community composition in four pairs of spring-grazed/ungrazed sites in south-central British Columbia, Canada. We also investigated whether pollinator or floral communities differed between the two threatened shrubsteppe habitat types we sampled—antelope-brush and big sagebrush shrubsteppe. Pan-trapping surveys captured 5907 bees, flies, beetles, wasps and butterflies constituting 253 species. We found that the percent cover of shrubs and bare soil increased with grazing, while the height of grasses and forbs decreased. In contrast, pollinator and flowering plant abundance, richness, diversity, and community composition were not significantly affected by grazing. Flowering plant and pollinator community composition did differ significantly between shrubsteppe habitats. Our results indicate that grasslands in North America, when managed responsibly, can maintain pollinator and flowering plant diversity under grazing pressure. The continued effort of land managers to balance ecological integrity and economic viability will be important for the conservation of grassland pollination systems.