Agricultural landscape homogenization has detrimental effects on biodiversity and key ecosystem services. Increasing agricultural landscape heterogeneity by increasing seminatural cover can help to mitigate biodiversity loss. However, the amount of seminatural cover is generally low and difficult to increase in many intensively managed agricultural landscapes. We hypothesized that increasing the heterogeneity of the crop mosaic itself (hereafter “crop heterogeneity”) can also have positive effects on biodiversity. In 8 contrasting regions of Europe and North America, we selected 435 landscapes along independent gradients of crop diversity and mean field size. Within each landscape, we selected 3 sampling sites in 1, 2, or 3 crop types. We sampled 7 taxa (plants, bees, butterflies, hoverflies, carabids, spiders, and birds) and calculated a synthetic index of multitrophic diversity at the landscape level. Increasing crop heterogeneity was more beneficial for multitrophic diversity than increasing seminatural cover. For instance, the effect of decreasing mean field size from 5 to 2.8 ha was as strong as the effect of increasing seminatural cover from 0.5 to 11%. Decreasing mean field size benefited multitrophic diversity even in the absence of seminatural vegetation between fields. Increasing the number of crop types sampled had a positive effect on landscape-level multitrophic diversity. However, the effect of increasing crop diversity in the landscape surrounding fields sampled depended on the amount of seminatural cover. Our study provides large-scale, multitrophic, cross-regional evidence that increasing crop heterogeneity can be an effective way to increase biodiversity in agricultural landscapes without taking land out of agricultural production.

Agricultural intensification is one of the main causes for the current biodiversity crisis. While reversing habitat loss on agricultural land is challenging, increasing the farmland configurational heterogeneity (higher field border density) and farmland compositional heterogeneity (higher crop diversity) has been proposed to counteract some habitat loss. Here, we tested whether increased farmland configurational and compositional heterogeneity promote wild pollinators and plant reproduction in 229 landscapes located in four major western European agricultural regions. High-field border density consistently increased wild bee abundance and seed set of radish (), probably through enhanced connectivity. In particular, we demonstrate the importance of crop-crop borders for pollinator movement as an additional experiment showed higher transfer of a pollen analogue along crop-crop borders than across fields or along semi-natural crop borders. By contrast, high crop diversity reduced bee abundance, probably due to an increase of crop types with particularly intensive management. This highlights the importance of crop identity when higher crop diversity is promoted. Our results show that small-scale agricultural systems can boost pollinators and plant reproduction. Agri-environmental policies should therefore aim to halt and reverse the current trend of increasing field sizes and to reduce the amount of crop types with particularly intensive management.

1. Bumblebees are important pollinators for a wide range of crops and wild plants.Performance of their colonies depends on pollen and nectar as food resources, but flowering plants are scarce in modern agricultural landscapes. It is well-known that semi-natural habitats can enhance floral resources and bumblebee abundance, but the impact of different crop types and their heterogeneity at the landscape scale remains unclear.2. We tested the effect of two different crop types (oilseed rape [OSR] and maize) and of configurational (field border density) and compositional heterogeneity (crop diversity) on weight gain of buff-tailed bumblebee colonies (Bombus terrestris) and the pollen diversity collected by them in 20 landscapes in Central Germany.3. We found that augmenting maize cover had a detrimental effect on pollen diversity collected by bumblebees, probably due to intensive management resulting in low plant diversity. This low pollen diversity translated into reduced colony growth, since colonies with high pollen diversity gained more weight than colonies with low pollen diversity. 4. In contrast, OSR cover and configurational and compositional heterogeneity did neither affect colony growth nor pollen diversity. However, for OSR, the timing of the flowering period was important. When OSR fields had a high flower cover at the end of the OSR blooming period, colonies showed increased growth rates. Synthesis and applications. Our results complement previous laboratory studies byshowing that high pollen diversity leads to better colony performance under field conditions. Therefore, the maintenance of floral diversity in agricultural landscapes is crucial to ensure that bumblebees can fulfil their nutritional needs.However, the heterogeneity of crops, at least under the currently very low levels of crop rotation, does not contribute to this aim. In contrast, crop identity and timing of mass-flowering crops turned out to be important factors, as maize reduced pollen resources, while late blooming oilseed rape (OSR) was beneficial to bumblebee colonies. Hence, maize cover per landscape should be reduced and strategies to enhance landscape wide flower diversity, especially towards and after the | 295Journal of Applied Ecology HASS et Al.

Increasing landscape heterogeneity by restoring semi‐natural elements to reverse farmland biodiversity declines is not always economically feasible or acceptable to farmers due to competition for land. We hypothesized that increasing the heterogeneity of the crop mosaic itself, hereafter referred to as crop heterogeneity, can have beneficial effects on within‐field plant diversity. Using a unique multi‐country dataset from a cross‐continent collaborative project covering 1,451 agricultural fields within 432 landscapes in Europe and Canada, we assessed the relative effects of compositional and configurational crop heterogeneity on within‐field plant diversity components. We also examined how these relationships were modulated by the position within the field. We found strong positive effects of configurational crop heterogeneity on within‐field plant alpha and gamma diversity in field interiors. These effects were as high as the effect of semi‐natural cover. In field borders, effects of crop heterogeneity were limited to alpha diversity. We suggest that a heterogeneous crop mosaic may overcome the high negative impact of management practices on plant diversity in field interiors, whereas in field borders, where plant diversity is already high, landscape effects are more limited. Synthesis and applications. Our study shows that increasing configurational crop heterogeneity is beneficial to within‐field plant diversity. It opens up a new effective and complementary way to promote farmland biodiversity without taking land out of agricultural production. We therefore recommend adopting manipulation of crop heterogeneity as a specific, effective management option in future policy measures, perhaps adding to agri‐environment schemes, to contribute to the conservation of farmland plant diversity.

Pollinators have experienced a dramatic decrease world‐wide due to agricultural intensification. In many countries, agri‐environment schemes (AES) have been introduced to counteract this current trend. However, until now, the relative importance of each AES for biodiversity and ecosystem services is still little understood and might change depending on landscape context. Complex landscape‐experiments are required to fill this knowledge gap, enabling the implementation of sustainable intensification of food production. In our study, we compared the effectiveness of the two most popular AES in Germany, organic farming and flower strips, in supporting pollinators and flower resources. We selected nine landscapes along a gradient of increasing field size, (configurational heterogeneity), each with a triplet of winter wheat fields: one organic, one conventional with flower strip and one conventional without flower strip as a control. We surveyed insect‐pollinated plants and pollinators (bumblebees, solitary bees and hoverflies). Additionally, we placed bumblebee colonies in the field edges to monitor their growth (colony weight gain) and reproduction (queen production). Flower strips stood out with the highest abundance and richness of pollinators. In contrast, bumblebee colony growth and plant richness benefited equally from organic and flower strip schemes. At the landscape scale, smaller fields had a positive effect on plant richness and bumblebee reproduction in flower strips. By contrast, bumblebee colonies in organic agriculture benefited most from large fields, as large organic fields provided much more flower resources than the narrow flower strips. Synthesis and applications. Our results showed that both local and landscape management shaped pollinator communities and their reproduction. Overall, organic farming and flower strips appeared to be effective tools to mitigate flower shortage in conventional cereal fields, with organic farming supporting the highest flowering plant cover per field. Flower strips enhanced local pollinator richness most, but increased bumblebee reproduction only when the surrounding landscapes had small fields with long field borders. Therefore, our results reveal that European Union policies need to take into account that the effectiveness of agri‐environment schemes depends on the structure of the surrounding landscape.