Climate warming has the potential to disrupt plant-pollinator interactions or to increase competition of co-flowering plants for pollinators, due to species-specific phenological responses to ...temperature. However, studies focusing on the effect of temperature on solitary bee emergence and the flowering onset of their food plants under natural conditions are still rare. We studied the effect of temperature on the phenology of the two spring bees Osmia cornuta and Osmia bicornis, by placing bee cocoons on eleven grasslands differing in mean site temperature. On seven grasslands, we additionally studied the effect of temperature on the phenology of the red-list plant Pulsatilla vulgaris, which was the first flowering plant, and of co-flowering plants with later flowering. With a warming of 0.1°C, the abundance-weighted mean emergence of O. cornuta males advanced by 0.4 days. Females of both species did not shift their emergence. Warmer temperatures advanced the abundance-weighted mean flowering of P. vulgaris by 1.3 days per 0.1°C increase, but did not shift flowering onset of co-flowering plants. Competition for pollinators between P. vulgaris and co-flowering plants does not increase within the studied temperature range. We demonstrate that temperature advances plant flowering more strongly than bee emergence suggesting an increased risk of pollinator limitation for the first flowers of P. vulgaris.
Biodiversity is important for many ecosystem processes. Global declines in pollinator diversity and abundance have been recognized, raising concerns about a pollination crisis of crops and wild ...plants. However, experimental evidence for effects of pollinator species diversity on plant reproduction is extremely scarce. We established communities with 1-5 bee species to test how seed production of a plant community is determined by bee diversity. Higher bee diversity resulted in higher seed production, but the strongest difference was observed for one compared to more than one bee species. Functional complementarity among bee species had a far higher explanatory power than bee diversity, suggesting that additional bee species only benefit pollination when they increase coverage of functional niches. In our experiment, complementarity was driven by differences in flower and temperature preferences. Interspecific interactions among bee species contributed to realized functional complementarity, as bees reduced interspecific overlap by shifting to alternative flowers in the presence of other species. This increased the number of plant species visited by a bee community and demonstrates a new mechanism for a biodiversity-function relationship ("interactive complementarity"). In conclusion, our results highlight both the importance of bee functional diversity for the reproduction of plant communities and the need to identify complementarity traits for accurately predicting pollination services by different bee communities.
Pollination improves the yield of most crop species and contributes to one-third of global crop production, but comprehensive benefits including crop quality are still unknown. Hence, pollination is ...underestimated by international policies, which is particularly alarming in times of agricultural intensification and diminishing pollination services. In this study, exclusion experiments with strawberries showed bee pollination to improve fruit quality, quantity and market value compared with wind and self-pollination. Bee-pollinated fruits were heavier, had less malformations and reached higher commercial grades. They had increased redness and reduced sugar–acid–ratios and were firmer, thus improving the commercially important shelf life. Longer shelf life reduced fruit loss by at least 11%. This is accounting for 0.32 billion US$ of the 1.44 billion US$ provided by bee pollination to the total value of 2.90 billion US$ made with strawberry selling in the European Union 2009. The fruit quality and yield effects are driven by the pollination-mediated production of hormonal growth regulators, which occur in several pollination-dependent crops. Thus, our comprehensive findings should be transferable to a wide range of crops and demonstrate bee pollination to be a hitherto underestimated but vital and economically important determinant of fruit quality.
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.
Agricultural land use results in direct biodiversity decline through loss of natural habitat, but may also cause indirect cross-habitat effects on conservation areas. We conducted three ...landscape-scale field studies on 67 sites to test the hypothesis that mass flowering of oilseed rape (Brassica napus) results in a transient dilution of bees in crop fields, and in increased competition between crop plants and grassland plants for pollinators. Abundances of bumble-bees, which are the main pollinators of the grassland plant Primula veris, but also pollinate oilseed rape (OSR), decreased with increasing amount of OSR. This landscape-scale dilution affected bumble-bee abundances strongly in OSR fields and marginally in grasslands, where bumble-bee abundances were generally low at the time of Primula flowering. Seed set of Primula veris, which flowers during OSR bloom, was reduced by 20 per cent when the amount of OSR within 1 km radius increased from 0 to 15 per cent. Hence, the current expansion of bee-attractive biofuel crops results in transient dilution of crop pollinators, which means an increased competition for pollinators between crops and wild plants. In conclusion, mass-flowering crops potentially threaten fitness of concurrently flowering wild plants in conservation areas, despite the fact that, in the long run, mass-flowering crops can enhance abundances of generalist pollinators and their pollination service.
Mass-flowering crops may affect long-term population dynamics, but effects on pollinators have never been studied across several years. We monitored wild bees in oilseed rape fields in 16 landscapes ...in Germany in two consecutive years. Effects on bee densities of landscape oilseed rape cover in the years of monitoring and in the previous years were evaluated with landscape data from three consecutive years. We fit empirical data to a mechanistic model to provide estimates for oilseed rape attractiveness and its effect on bee productivity in comparison to the rest of the landscape, and we evaluated consequences for pollinator densities in consecutive years. Our results show that high oilseed rape cover in the previous year enhances current densities of wild bees (except for bumble bees). Moreover, we show a strong attractiveness of and dilution on (i.e., decreasing bee densities with increasing landscape oilseed rape cover) oilseed rape for bees during flowering in the current year, modifying the effect of the previous year's oilseed rape cover in the case of wild bees (excluding
Bombus
). As long as other factors such as nesting sites or natural enemies do not limit bee reproduction, our findings suggest long-term positive effects of mass-flowering crops on bee populations, at least for non-
Bombus
generalists, which possibly help to maintain crop pollination services even when crop area increases. Similar effects are conceivable for other organisms providing ecosystem services in annual crops and should be considered in future studies.
► Many landscapes are mosaics of human-managed and natural areas. ► Movement, or spillover, across habitat types may be an important ecological process. ► We review the evidence for spillover for ...five functional groups. ► There is much evidence for spillover from natural habitats to managed areas. ► Research is lacking for spillover from managed areas to natural habitats.
Land-use intensification has led to a landscape mosaic that juxtaposes human-managed and natural areas. In such human-dominated and heterogeneous landscapes, spillover across habitat types, especially in systems that differ in resource availability, may be an important ecological process structuring communities. While there is much evidence for spillover from natural habitats to managed areas, little attention has been given to flow in the opposite direction. This paper synthesizes studies published to date from five functionally important trophic groups, herbivores, pathogens, pollinators, predators, and seed dispersers, and discusses evidence for spillover from managed to natural systems in all five groups. For each of the five focal groups, studies in the natural to managed direction are common, often with multiple review articles on each subject which document dozens of examples. In contrast, the number of studies which examine movement in the managed to natural direction is generally less than five studies per trophic group. These findings suggest that spillover in the managed to natural direction has been largely underestimated. As habitat modification continues, resulting in increasingly fragmented landscapes, the likelihood and size of any spillover effect will only increase.
Understanding how landscape characteristics affect biodiversity patterns and ecological processes at local and landscape scales is critical for mitigating effects of global environmental change. In ...this review, we use knowledge gained from human-modified landscapes to suggest eight hypotheses, which we hope will encourage more systematic research on the role of landscape composition and configuration in determining the structure of ecological communities, ecosystem functioning and services. We organize the eight hypotheses under four overarching themes. Section A: ‘landscape moderation of biodiversity patterns' includes (1) the landscape species pool hypothesis—the size of the landscape-wide species pool moderates local (alpha) biodiversity, and (2) the dominance of beta diversity hypothesis—landscape-moderated dissimilarity of local communities determines landscape-wide biodiversity and overrides negative local effects of habitat fragmentation on biodiversity. Section B: ‘landscape moderation of population dynamics' includes (3) the cross-habitat spillover hypothesis—landscape-moderated spillover of energy, resources and organisms across habitats, including between managed and natural ecosystems, influences landscape-wide community structure and associated processes and (4) the landscape-moderated concentration and dilution hypothesis—spatial and temporal changes in landscape composition can cause transient concentration or dilution of populations with functional consequences. Section C: ‘landscape moderation of functional trait selection’ includes (5) the landscape-moderated functional trait selection hypothesis—landscape moderation of species trait selection shapes the functional role and trajectory of community assembly, and (6) the landscape-moderated insurance hypothesis—landscape complexity provides spatial and temporal insurance, i.e. high resilience and stability of ecological processes in changing environments. Section D: ‘landscape constraints on conservation management' includes (7) the intermediate landscape-complexity hypothesis—landscape-moderated effectiveness of local conservation management is highest in structurally simple, rather than in cleared (i.e. extremely simplified) or in complex landscapes, and (8) the landscape-moderated biodiversity versus ecosystem service management.
Growing evidence for declines in wild bees calls for the development and implementation of effective mitigation measures. Enhancing floral resources is a widely accepted measure for promoting bees in ...agricultural landscapes, but effectiveness varies considerably between landscapes and regions. We hypothesize that this variation is mainly driven by a combination of the direct effects of measures on local floral resources and the availability of floral resources in the surrounding landscape. To test this, we established wildflower strips in four European countries, using the same seed mixture of forage plants specifically targeted at bees. We used a before–after control–impact approach to analyse the impacts of wildflower strips on bumblebees, solitary bees and Red List species and examined to what extent effects were affected by local and landscape‐wide floral resource availability, land‐use intensity and landscape complexity. Wildflower strips generally enhanced local bee abundance and richness, including Red‐listed species. Effectiveness of the wildflower strips increased with the local contrast in flower richness created by the strips and furthermore depended on the availability of floral resources in the surrounding landscape, with different patterns for solitary bees and bumblebees. Effects on solitary bees appeared to decrease with increasing amount of late‐season alternative floral resources in the landscape, whereas effects on bumblebees increased with increasing early‐season landscape‐wide floral resource availability. Synthesis and applications. Our study shows that the effects of wildflower strips on bees are largely driven by the extent to which local flower richness is increased. The effectiveness of this measure could therefore be enhanced by maximizing the number of bee forage species in seed mixtures, and by management regimes that effectively maintain flower richness in the strips through the years. In addition, for bumblebees specifically, our study highlights the importance of a continuous supply of food resources throughout the season. Measures that enhance early‐season landscape‐wide floral resource availability, such as the cultivation of oilseed rape, can benefit bumblebees by providing the essential resources for colony establishment and growth in spring. Further research is required to determine whether, and under what conditions, wildflower strips result in actual population‐level effects.
Mass‐flowering crops (MFCs) are increasingly cultivated and might influence pollinator communities in MFC fields and nearby semi‐natural habitats (SNHs). Across six European regions and 2 years, we ...assessed how landscape‐scale cover of MFCs affected pollinator densities in 408 MFC fields and adjacent SNHs. In MFC fields, densities of bumblebees, solitary bees, managed honeybees and hoverflies were negatively related to the cover of MFCs in the landscape. In SNHs, densities of bumblebees declined with increasing cover of MFCs but densities of honeybees increased. The densities of all pollinators were generally unrelated to the cover of SNHs in the landscape. Although MFC fields apparently attracted pollinators from SNHs, in landscapes with large areas of MFCs they became diluted. The resulting lower densities might negatively affect yields of pollinator‐dependent crops and the reproductive success of wild plants. An expansion of MFCs needs to be accompanied by pollinator‐supporting practices in agricultural landscapes.
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