Effects of perennial wildflower strips and landscape structure on birds in intensively farmed agricultural landscapes

Farmland bird populations are in a deep crisis across Europe. Agri-environment schemes (AES) were implemented by the European Union to stop and reverse the general decline of biodiversity in agricultural landscapes. In Germany, flower strips are one of the most common AES. Establishing high-quality perennial wildflower strips (WFS) with species-rich native forb mixtures from regional seed propagation is a recent approach, for which the effectiveness for birds has not yet been sufficiently studied. We surveyed breeding birds and vegetation on 40 arable fields with WFS (20 with single and 20 with aggregated WFS) and 20 arable fields lacking WFS as controls across Saxony-Anhalt (Germany). Additionally, vegetation composition, WFS quantity and landscape structure (e.g. distance to nearest woody element) were considered in our analyses. All WFS were established with species-rich native seed mixtures (30 forbs) in agricultural practice as AES. Arable fields with WFS had a higher species richness and territory density of birds than controls, confirming the effectiveness of this AES. A forb-rich vegetation was the main driver promoting birds. Flower strip quantity at the landscape level had positive effects only on bird densities, but also single WFS achieved benefits. A short distance from WFS to woody elements increased total bird species richness. However, the density of farmland birds, which are target species of these AES, were negatively affected by the proximity and proportion of woody elements in the vicinity. The effect of the proportion of non-intensively used open habitats and overall habitat richness was unexpectedly low in the otherwise intensively farmed landscape. Species-rich perennial WFS significantly promoted breeding birds. Successful establishment of WFS, resulting in high-quality habitats, a high flower strip quantity as well as implementation in open landscapes were shown to maximise the effectiveness for restoring declining and AES target farmland birds.     

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.     

Spatial configuration and landscape context of wildflower areas determine their benefits to pollinator α- and β-diversity

Wildflower areas have become a staple tool within agro-environmental schemes (AES) to counteract pollinator declines. While their role in providing food resources to resident flower-visiting insects is unambiguous, the conservation effectiveness in a landscape context is less clear. Particularly, how multiple vs. single wildflower area utilization differs between simple and complex landscapes is understudied. We examined colonisation and community dynamics of wild bees and hoverflies in 33 newly established wildflower areas across a gradient of landscape complexity (amount of semi-natural habitat) and connectivity (presence of additional wildflower areas) for seven weeks during three consecutive years (one year during and two after establishment). We recorded more than 25% of the wild bee and hoverfly species of Hesse in an area of approx. 10 ha, substantiating the general benefit of wildflower areas to pollinators. While alpha-diversity increased with landscape complexity in isolated wildflower areas, the opposite pattern was observed for connected areas. The low alpha-diversity in complex landscapes indicated a dilution effect between connected sites. The inverse relationship between alpha and beta diversity among wildflower areas within landscapes suggests interspecific trade-offs between local resource conditions and landscape context. Accordingly, the establishment of multiple wildflower areas within AES is advisable to increase connectivity of suitable habitats in simple landscapes. Moreover, adjusting local conditions (plant diversity) to landscape context likely optimizes conservation effectiveness in modern agroecosystems.     

Redundancy in wildflower strip species helps support spatiotemporal variation in wild bee communities on diversified farms

Wildflower strips are a management practice increasingly used to provide floral resources to wild bees in agroecosystems. Yet, despite known spatiotemporal variation in wild bee communities, the degree to which different wildflower strip species consistently support wild bee communities is poorly understood. Additionally, whether such consistency is related to the functional roles wildflower species play (e.g., in supporting diverse, rare, or unique suites of bee species) has not been considered. Over three years and on four diversified farms, we evaluated spatiotemporal variation in wild bee communities and bee-flower interactions in wildflower strips to better understand the roles of flower strip species in supporting bees. We documented spatiotemporal variation in the abundance, richness, and composition of local wild bee communities. Certain wildflower species consistently supported the highest richness of wild bees across years. These wildflower species were regularly core members of the bee-flower interaction network, visited by both generalist and specialist bees. By contrast, wildflower species supporting the most unique suites of bees were variable in this role among farms. In order to select plant species for wildflower strips that consistently support a high diversity of wild bee communities within farm landscapes, it is useful to consider several different functional roles that plants may play. Whereas a handful of wildflower species may be visited by the majority of local wild bee species, achieving support for the remaining, and perhaps rarer, bee species will require planting additional flower species, which may appear redundant until the spatiotemporal variation in wild bee communities is more thoroughly considered. This functional approach to selecting wildflower species for bee conservation efforts is important for making practical recommendations to land managers and for guiding best management practices in different regions and with diverse management goals.     

Trait-Specific Responses of Wild Bee Communities to Landscape Composition,Configuration and Local Factors

Land-use intensification and loss of semi-natural habitats have induced a severe decline of bee diversity in agricultural landscapes. Semi-natural habitats like calcareous grasslands are among the most important bee habitats in central Europe, but they are threatened by decreasing habitat area and quality, and by homogenization of the surrounding landscape affecting both landscape composition and configuration. In this study we tested the importance of habitat area, quality and connectivity as well as landscape composition and configuration on wild bees in calcareous grasslands. We made detailed trait-specific analyses as bees with different traits might differ in their response to the tested factors. Species richness and abundance of wild bees were surveyed on 23 calcareous grassland patches in Southern Germany with independent gradients in local and landscape factors. Total wild bee richness was positively affected by complex landscape configuration, large habitat area and high habitat quality (i.e. steep slopes). Cuckoo bee richness was positively affected by complex landscape configuration and large habitat area whereas habitat specialists were only affected by the local factors habitat area and habitat quality. Small social generalists were positively influenced by habitat area whereas large social generalists (bumblebees) were positively affected by landscape composition (high percentage of semi-natural habitats). Our results emphasize a strong dependence of habitat specialists on local habitat characteristics, whereas cuckoo bees and bumblebees are more likely affected by the surrounding landscape. We conclude that a combination of large high-quality patches and heterogeneous landscapes maintains high bee species richness and communities with diverse trait composition. Such diverse communities might stabilize pollination services provided to crops and wild plants on local and landscape scales.     

Larger patches of diverse floral resources increase insect pollinator density,diversity, and their pollination of native wildflowers

Native wildflower plantings can be used to provide nutritional resources to support pollinating insects, yet the effects of planting size and bloom richness on the density, diversity, and function of these insects are not well understood. We established stands of twelve native flowering perennial plant species in replicated plots ranging in size from 1 to 100 m². These plots were sampled for insect pollinators, bloom richness, and seed production by three wildflower species. Honeybees, wild bees, and hoverflies all responded positively to increasing flower richness, whereas particular insect pollinator groups responded differently to the size of the flowering plant area. The density of honeybees and hoverflies was not affected by increasing flowering patch size, whereas in general, wild bees were observed at higher density and diversity in the 30 and 100 m² patches. Increasing wildflower patch size, and thus wild bee density, resulted in greater seed set in the sampled wildflowers. These results indicate that wild bees are sensitive to the area and richness of floral resources in patches, even at relatively small scales. Therefore, larger wildflower plantings with more diverse flower species mixes are more suitable for the conservation of wild pollinators and reproduction of sown species.     

Agricultural landscapes with organic crops support higher pollinator diversity

Pollinators are traditionally thought to perceive non-flowering crop fields as hostile landscape matrix. In this study, we show that landscapes composed of higher proportions of organic crop fields support more bee species at greater abundances in fallow strips. An increase in organic cropping in the surrounding landscape from 5% to 20% enhanced bee species richness in fallow strips by 50%, density of solitary bees by 60% and bumble bee density by 150%. Bee species richness and bumble bee density responded strongest to organic cropping in landscape sectors with 500 m radius, solitary bee density in landscape sectors with 250 m radius. The most likely source of these results is that crop and noncrop habitats are strongly connected via bee foraging at the landscape scale. It seems likely that bees depending on nesting sites in fallow strips benefited from the more abundant flower resources provided by broadleaved weeds in organic crop fields. We conclude that the incorporation of organic crop fields into conventionally managed agricultural landscapes can provide food resources needed to sustain greater pollinator species richness in noncrop habitats.     

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

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Farming for bees: annual variation in pollinator populations across agricultural landscapes

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Wild bees respond complementarily to ‘high-quality’ perennial and annual habitats of organic farms in a complex landscape

Agricultural intensification leads to large-scale loss of habitats offering food and nesting sites for bees. This has resulted in a severe decline of wild bee diversity and abundance during the past decades. There is an urgent need for cost-effective conservation measures to mitigate this decline. We analysed the impact of five different high-quality habitats on species richness and abundance of wild bees in a complex landscape of north-western Switzerland at six sites. The five habitat types included 45 plots situated on eight organic farms and were composed of 16 low-input meadows, six low-input pastures, seven herbaceous strips adjacent to hedges, five sown flower strips and eleven organic cereal fields. All of them are financially subsidised by the Swiss agri-environmental scheme. Wild bees were sampled between the end of April and end of August 2014 by using trio-pan traps and complementary sweep netting on these five habitat types. On 45 plots we recorded 3973 bee specimens, belonging to 91 species, 16 of which are red listed, revealing a high bee species richness in the study area. Wild bee species richness and abundance were best explained by habitat type, number of flowering plants and site. A strong relationship of increasing number of flowering plants and bee species richness and abundance was found. Grassland habitats, especially low-input meadows, harboured the highest species richness and abundances. Organic cereal fields showed a potential to conserve bee species relevant to nature conservation (harbouring exclusively two red list species and four rare species). Ordination analysis of the bee communities showed a relative dissimilarity between the habitat types and indicates their complementary effects to benefit the diversity of wild bees. Our results demonstrate that a matrix of low-input habitats are needed to sustain rich assemblages of wild bees in agroecosystems.     

Linear woodlots increase wild bee abundance by providing additional food sources in an agricultural landscape

1. Mid-field woodlots play an important role in maintaining biodiversity in agricultural landscapes. However, it is not clear whether non-linear or linear woodlots are most beneficial for wild bee conservation.
2. We assessed the attractiveness of two common types of woodlots in an agricultural landscape in northern Poland (non-linear and linear: 7 and 9 sites, respectively) in terms of wild bee abundance, species richness, and functional diversity.
3. Linear habitats had higher abundance of wild bees. However, woodlot type did not affect wild bee species composition or functional trait composition. Species composition responded significantly to measures of syntaxonomic heterogeneity and landscape heterogeneity. Woodlot area, landscape context (isolation and landscape heterogeneity), and syntaxonomic heterogeneity explained most of the differences among habitats (non-linear vs. linear) in wild bee abundance and species richness, regardless of the habitat type. The higher attractiveness of linear woodlots was due to increased food availability in the herbaceous layer in the spring–summer (June) and summer (July–August) periods.
4. Linear woodlots have the potential to be used as tools for integrating agricultural production with biodiversity conservation and ecosystem services.

     

Forested landscapes promote richness and abundance of native bees (Hymenoptera: Apoidea: Anthophila) in Wisconsin apple orchards

Wild bees provide vital pollination services for many native and agricultural plant species, yet the landscape conditions needed to support wild bee populations are not well understood or appreciated. We assessed the influence of landscape composition on bee abundance and species richness in apple (Malus spp.) orchards of northeastern Wisconsin during the spring flowering period. A diverse community of bee species occurs in these apple orchards, dominated by wild bees in the families Andrenidae and Halictidae and the honey bee, Apis mellifera L. Proportion of forest area in the surrounding landscape was a significant positive predictor of wild bee abundance in orchards, with strongest effects at a GIS (Geographic Information Systems) buffer distance of 1,000 m or greater. Forest area also was positively associated with species richness, showing strongest effects at a buffer distance of 2,000 m. Nonagricultural developed land (homes, lawns, etcetera) was significantly negatively associated with species richness at buffer distances >750 m and wild bee abundance in bowl traps at all distances. Other landscape variables statistically associated with species richness or abundance of wild bees included proportion area of pasture (positive) and proportion area of roads (negative). Forest area was not associated with honey bee abundance at any buffer distance. These results provide clear evidence that the landscape surrounding apple orchards, especially the proportion of forest area, affects richness and abundance of wild bees during the spring flowering period and should be a part of sustainable land management strategies in agro-ecosystems of northeastern Wisconsin and other apple growing regions.     

Stability of pollination services decreases with isolation from natural areas despite honey bee visits

Sustainable agricultural landscapes by definition provide high magnitude and stability of ecosystem services, biodiversity and crop productivity. However, few studies have considered landscape effects on the stability of ecosystem services. We tested whether isolation from florally diverse natural and semi-natural areas reduces the spatial and temporal stability of flower-visitor richness and pollination services in crop fields. We synthesised data from 29 studies with contrasting biomes, crop species and pollinator communities. Stability of flower-visitor richness, visitation rate (all insects except honey bees) and fruit set all decreased with distance from natural areas. At 1 km from adjacent natural areas, spatial stability decreased by 25, 16 and 9% for richness, visitation and fruit set, respectively, while temporal stability decreased by 39% for richness and 13% for visitation. Mean richness, visitation and fruit set also decreased with isolation, by 34, 27 and 16% at 1 km respectively. In contrast, honey bee visitation did not change with isolation and represented > 25% of crop visits in 21 studies. Therefore, wild pollinators are relevant for crop productivity and stability even when honey bees are abundant. Policies to preserve and restore natural areas in agricultural landscapes should enhance levels and reliability of pollination services.     

Resource overlap and possible competition between honey bees and wild bees in central Europe

Evidence for interspecific competition between honey bees and wild bees was studied on 15 calcareous grasslands with respect to: (1) foraging radius of honey bees, (2) overlap in resource use, and (3) possible honey bee effects on species richness and abundance of flower-visiting, ground-nesting and trap-nesting wild bees. The grasslands greatly differed in the number of honey bee colonies within a radius of 2 km and were surrounded by agricultural habitats. The number of flower-visiting honey bees on both potted mustard plants and small grassland patches declined with increasing distance from the nearest apiary and was almost zero at a distance of 1.5–2.0 km. Wild bees were observed visiting 57 plant species, whereas honey bees visited only 24 plant species. Percentage resource overlap between honey bees and wild bees was 45.5%, and Hurlbert’s index of niche overlap was 3.1. In total, 1849 wild bees from 98 species were recorded on the calcareous grasslands. Neither species richness nor abundance of wild bees were negatively correlated with the density of honey bee colonies (within a radius of 2 km) or the density of flower-visiting honey bees per site. Abundance of flower- visiting wild bees was correlated only with the percentage cover of flowering plants. In 240 trap nests, 1292 bee nests with 6066 brood cells were found. Neither the number of bee species nor the number of brood cells per grassland was significantly correlated with the density of honey bees. Significant correlations were found only between the number of brood cells and the percentage cover of shrubs. The number of nest entrances of ground-nesting bees per square metre was not correlated with the density of honey bees but was negatively correlated with the cover of vegetation. Interspecific competition by honey bees for food resources was not shown to be a significant factor determining abundance and species richness of wild bees. Received: 22 March 1999 / Accepted: 24 September 1999     

Scale dependent drivers of wild bee diversity in tropical heterogeneous agricultural landscapes

Factors associated with agricultural intensification, for example, loss of seminatural vegetation and pesticide use has been shown to adversely affect the bee community. These factors may impact the bee community differently at different landscape scales. The scale dependency is expected to be more pronounced in heterogeneous landscapes. However, the scale‐dependent response of the bee community to drivers of its decline is relatively understudied, especially in the tropics where the agricultural landscape is often heterogeneous. This study looked at effects of agricultural intensification on bee diversity at patch and landscape scales in a tropical agricultural landscape. Wild bees were sampled using 12 permanent pan trap stations. Patch and landscape characteristics were measured within a 100 m (patch scale) and a 500 m (landscape scale) radius of pan trap stations. Information on pesticide input was obtained from farmer surveys. Data on vegetation cover, productivity, and percentage of agricultural and fallow land (FL) were collected using satellite imagery. Intensive areas in a bee‐site network were less specialized in terms of resources to attract rare bee species while the less intensive areas, which supported more rare species, were more vulnerable to disturbance. A combination of patch quality and diversity as well as pesticide use regulates species diversity at the landscape scale (500 m), whereas pesticide quantity drove diversity at the patch scale (100 m). At the landscape scale, specialization of each site in terms of resources for bees increased with increasing patch diversity and FL while at the patch scale specialization declined with increased pesticide use. Bee functional groups responded differentially to landscape characteristics as well as pesticide use. Wood nesting bees were negatively affected by the number of pesticides used but other bee functional groups were not sensitive to pesticides. Synthesis and Applications: Different factors affect wild bee diversity at the scale of landscape and patch in heterogeneous tropical agricultural systems. The differential response of bee functional groups to agricultural intensification underpins the need for guild‐specific management strategies for wild bee conservation. Less intensively farmed areas support more rare species and are vulnerable to disturbance; consequently, these areas should be prioritized for conservation to maintain heterogeneity in the landscape. It is important to conserve and restore seminatural habitats to maintain complexity in the landscapes through participatory processes and to regulate synthetic chemical pesticides in farm operations to conserve the species and functional diversity of wild bees.     

Small wild bee abundance declines with distance into strawberry crops regardless of field margin habitat

The preservation of pollinator habitat on croplands in the form of hedgerows, wildflower strips, and natural and semi-natural areas can help maintain and enhance wild bee populations in agricultural landscapes. However, there have been few comparisons of the effectiveness of different types of field-margin pollinator habitat in maintaining bee diversity and pollination of the focal crops. We compared wild bee abundance, species richness and community composition between strawberry crops bordered by hedgerows, and those bordered by larger expanses of natural land (forests). Strawberry is an ideal crop in which to investigate pollinator export from field margins as the rows are covered with straw, which reduces habitat for ground-nesting bees within the crop; thus, most wild pollinators need to enter the crop from the margins. We sampled bees in six strawberry fields with hedgerow margins and six strawberry fields with forested margins of at least 200 m in length, using a paired design. We examined strawberry pollen deposition at regular intervals into the fields, and the magnitude of pollinator export from the field margins towards the centre of the crops. We found that bees as a group were no more species-rich or abundant in crops bordered by forests than in crops bordered by hedgerows, although large-bodied bees were more abundant in the former than the latter. Regardless of field-margin type, we found that small wild bee abundance declined significantly from the edge to the centre of the crop, but honey bee (Apis mellifera L.) and large-bodied bee abundance did not. Strawberry pollen deposition also did not decline with distance into the crop. Although previous work indicates that small wild bees are more effective (yield-increasing) pollinators of strawberry on a per-visit basis, their limited foraging ranges suggest they may only pollinate areas near the crop margins, given typical field sizes in our area.     

Attractiveness of sown wildflower strips to flower-visiting insects depends on seed mixture and establishment success

Establishing wildflower strips has been suggested as an effective measure to promote pollination services, pest control or general insect biodiversity, but little is known about the integration of these different objectives when selecting flower seed mixtures. In ten agricultural landscapes in the Netherlands, we established a wildflower strip (0.4 – 4.9 ha) with half of each strip sown with a mixture targeting longer-tongued pollinators and the other half sown with a mixture targeting shorter-tongued pollinators and natural enemies. We determined establishment success of sown wildflowers and evaluated the attractiveness of the established flower communities to multiple functional groups of flower visitors: bumblebees (long-tongued pollinators), hoverflies (short-tongued pollinators and natural enemies), and butterflies and total flower-visitor richness (indicators of wider biodiversity values). Bumblebees clearly preferred the pollinator-targeted seed mixture and were positively associated with cover of Fabaceae and negatively with Apiaceae. Hoverflies consistently preferred the natural enemy mixture and were positively associated with Apiaceae. The other target groups displayed no clear responses to seed mixture type but instead were associated with local flower richness within strips. Across sites, responses of flower-visitors to sown mixture types did not depend on wildflower strip size, proportion of surrounding semi-natural habitat, or flower variables. However, all flower-visitors except butterflies increased with increasing established cover or richness of (sown) flower species across sites. Our results suggest that, although species-rich wildflower strips may benefit several species groups, maximising different objectives involves trade-offs between functional groups that prefer short- or long-corolla flowers. Furthermore, our study suggests that sowing a wildflower mixture does not necessarily result in a vegetation with the same composition as the seed mixture as species may establish poorly or not at all. Selection of flower species for seed mixtures should therefore, in addition to insect target group, take the establishment characteristics of plant species into account.     

Importance of forest fragments as pollinator habitat varies with season and guild

Large areas of Western Europe are covered with intensively managed agricultural land. In these landscapes, wild pollinators depend on fragments of semi-natural habitat for foraging or reproduction. Small forest patches are often the most abundant type of semi-natural habitat in these agricultural landscapes. We investigated the role these patches play in conserving the pollinator community in intensively managed agricultural landscapes.Our survey of the pollinator community in 16 forest fragments showed that the pollinator community in the edges of small forest fragments is strongly influenced by forest and forest edge characteristics. Old forest fragments with a well-developed herb layer had more diverse bee communities than recent forests or old forests without a herb layer, but overall lower activity-abundances, while sun exposure of the forest edges had a strong positive effect on pollinater activity-abundance in general. The hoverfly community had higher activity-abundances in forest edges with a higher flower-index, while saproxylic hoverflies were caught in higher numbers in sites with a higher forest cover in the surrounding landscape.We also detected a strong seasonal effect. The effects of herb layer cover on bee species richness and activity-abundance were much stronger in spring than in summer, while bee species richness was also strongly positively correlated with forest age in spring. A strong positive correlation between pollinator species richness and sun exposure was found in summer, after canopy closure.While the sampled forest edges harbour a rich and diverse pollinator community, cavity-nesting bees were very scarce. This is probably caused by the low amount of dead wood in the studied forest fragments.We conclude that small forest fragments can play an important role in conserving the pollinator community, especially bees and saproxylic hoverflies. The importance of these forest fragments is strongest in spring, when the herb layer provides foraging resources.     

Landscape spatial configuration is a key driver of wild bee demographics

The majority of studies investigating the effects of landscape composition and configuration on bee populations have been conducted in regions of intensive agricultural production, ignoring regions which are dominated by seminatural habitats, such as the islands of the Aegean Archipelago. In addition, research so far has focused on the landscape impacts on bees sampled in cropped fields while the landscape effects on bees inhabiting seminatural habitats are understudied. Here, we investigate the impact of the landscape on wild bee assemblages in 66 phryganic (low scrubland) communities on 8 Aegean islands. We computed landscape metrics (total area and total perimeter–area ratio) in 4 concentric circles (250, 500, 750, and 1000 m) around the center of each bee sampling site including 3 habitat groups (namely phrygana, cultivated land, and natural forests). We further measured the local flower cover in 25 quadrats distributed randomly at the center of each sampling site. We found that the landscape scale is more important than the local scale in shaping abundance and species richness of bees. Furthermore, habitat configuration was more important than the total area of habitats, probably because it affects bees’ movement across the landscape. Phrygana and natural forests had a positive effect on bee demographics, while cultivated land had a negative effect. This demonstrates that phryganic specialists drive bee assemblages in these seminatural landscapes. This finding, together with the shown importance of landscape scale, should be considered for the management of wild bees with special emphasis placed on the spatial configuration of seminatural habitats.     

Enhancing flowering plant functional richness improves wild bee diversity in vineyard inter‐rows in different floral kingdoms

Wild bees are threatened by multiple interacting stressors, such as habitat loss, land use change, parasites, and pathogens. However, vineyards with vegetated inter‐rows can offer high floral resources within viticultural landscapes and provide foraging and nesting habitats for wild bees. Here, we assess how vineyard management regimes (organic vs. conventional; inter‐row vegetation management) and landscape composition determine the inter‐row plant and wild bee assemblages, as well as how these variables relate to functional traits in 24 Austrian and 10 South African vineyards. Vineyards had either permanent vegetation cover in untilled inter‐rows or temporary vegetation cover in infrequently tilled inter‐rows. Proportion of seminatural habitats (e.g., fallows, grassland, field margins) and woody structures (e.g., woodlots, single trees, tree rows) were used as proxies for landscape composition and mapped within 500‐m radius around the study vineyards. Organic vineyard management increased functional richness (FRic) of wild bees and flowering plants, with woody structures marginally increasing species richness and FRic of wild bees. Wild bee and floral traits were differently associated across the countries. In Austria, several bee traits (e.g., lecty, pollen collection type, proboscis length) were associated with flower color and symmetry, while in South African vineyards, only bees’ proboscis length was positively correlated with floral traits characteristic of Asteraceae flowers (e.g., ray–disk morphology, yellow colors). Solitary bee species in Austria benefitted from infrequent tillage, while ground nesting species preferred inter‐rows with undisturbed soils. Higher proportions of woody structures in surrounding landscapes resulted in less solitary and corbiculate bees in Austria, but more aboveground nesting species in South Africa. In both countries, associations between FRic of wild bees and flowering plants were positive both in organic and in conventional vineyards. We recommend the use of diverse cover crop seed mixtures to enhance plant flowering diversity in inter‐rows, to increase wild bee richness in viticultural landscapes.