1. Continuous availability of food resources, such as pollen, is vital for many insects that provide pollination and pest control services to agriculture. However, there is a lack of knowledge about the shared or complementary use of floral resources by such species, which hampers more effective landscape management to simultaneously promote them in agroecosystems.2. Here, we simultaneously quantified pollen use by a bumblebee (Bombus terrestris) and a mason bee (Osmia bicornis), two bee species recognized as important crop pollinators, as well as a lacewing (Chrysoperla carnea) and a ladybeetle species (Harmonia axyridis), both common predators of crop aphids, throughout the season in 23 agricultural landscapes in Germany and Switzerland.3. Pollen diets were more diverse and similar among C. carnea and H. axyridis compared to the two bee species, but all four species shared key pollen types early in the season such as Acer, Quercus, Salix and Prunus. All species exhibited a pronounced shift in pollen sources from primarily woody plants (mainly trees) in spring to primarily herbaceous plants in summer. The majority of pollen (overall ≥64%) came from non-agricultural plants even in crop-dominated landscapes. Synthesis and applications.Our results highlight the importance of trees as pollen sources for many insect species, particularly early in the season. Our findings support incentives that promote heterogeneous agricultural landscapes including both woody and herbaceous semi-natural habitats, ensuring phenological complementarity of floral resources for insect species that can provide pollination and pest control services to agriculture. The identified key plant species can help to design and optimize agri-environment schemes to promote these functionally important insects. K E Y W O R D Sbumblebee, floral resources, foraging habitat, landscape resources, mason bee, pollen diet, resource specialization

The yield of animal-pollinated crops is threatened by bee declines, but its precise sensitivity is poorly known. We therefore determined the yield dependence of Hokkaido pumpkin in Germany on insect pollination by quantifying: (i) the relationship between pollen receipt and fruit set and (ii) the cumulative pollen deposition of each pollinator group. We found that approximately 2500 pollen grains per flower were needed to maximize fruit set. At the measured rates of flower visitation, we estimated that bumblebees (21 visits/flower lifetime, 864 grains/visit) or honeybees (123 visits, 260 grains) could individually achieve maximum crop yield, whereas halictid bees are ineffective (11 visits, 16 grains). The pollinator fauna was capable of delivering 20 times the necessary amount of pollen. We therefore estimate that pumpkin yield was not pollination-limited in our study region and that it is currently fairly resilient to single declines of honeybees or wild bumblebees.

Intensive agricultural landscapes can be hostile for bees due to a lack of floral and nesting resources, and due to management-related stress such as pesticide use and soil tillage. This threatens the pollination services that bees deliver to insect-pollinated crops. We studied the effects of farming intensity (organic vs. conventional, number of insecticide applications) and availability of semi-natural habitats at the field and landscape scale on pollinator visits and pollen delivery to pumpkin in Germany. We found that wild bumble bees were the key pollinators of pumpkin in terms of pollen delivery, despite fivefold higher visitation frequency of honey bees. Critically, we observed that the area of cropland had stronger effects on bees’ pollen deposition than the area of seminatural habitats. Specifically, a 10% increase of the proportion of cropland reduced pollen delivery by 7%. Pumpkin provides a striking example for a key role of wild pollinators in crop pollination even at high numerical dominance of honey bees. In addition, our findings suggest that habitat conversion to agricultural land is a driver of deteriorating pollination. This underlines the importance to maintain sufficient areas of non-crop habitats in agricultural landscapes.

Wild bumblebees are key pollinators of crops and wild plants that rely on the continuous availability of floral resources. A better understanding of the spatio-temporal availability and use of floral food resources may help to promote bumblebees and their pollination services in agricultural landscapes. We placed colonies of Bombus terrestris L. in 24 agricultural landscapes with various degrees of floral resource availability and assessed different parameters of colony growth and fitness. We estimated pollen availability during different periods of colony development based on detailed information of the bumblebee pollen diet and the spatial distribution of the visited plant species. Total pollen availability did not significantly explain colony growth or fitness. However, when using habitat maps, the weight gain of colonies, the number of queen cells, and colony survival decreased with increasing distance from the forest. The better explanation of bumblebee performance by forest proximity than by (plant-inferred) pollen availability indicates that other functions of forests than pollen provision were important. The conservation of forests next to agricultural land might help to sustain high populations of these important wild pollinators and enhance their crop pollination services. Combining different mapping approaches might help to further disentangle complex relationships between B. terrestris and their environment in agricultural landscapes.

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