Climate, landscape composition, management practice, and wild bee pollination are all variables thought to play significant roles in commercial apple production. However, how these variables affect production efficiency under field-realistic conditions has not been investigated at large geographical scales. We combined intensive standardized field surveys (using netting and pan traps) with structural equation models to explore the relative impact of biotic and abiotic variables on bee diversity, apple yield and fruit quality, and their ability to represent reliable proxies of apple production. Here we show that apple yields are mainly driven by management practice, without evidence for a significantly superior contribution by managed honey bees. Total wild bee diversity, while negatively correlated with honey bee dominance, promoted apple quality by enhancing seed set number. Our study demonstrates that even across a broad geographical range there is potential to harness wild bee diversity as nature-based solution and as a substitute to an exclusive reliance on honey bees in the context of commercial apple production.

Differences in pollination effectiveness (PE) among pollinators have been widely documented. However, the morphological and behavioural traits underlying these differences have been less investigated. We used single-visit pollen deposition to apple flowers to explore the relationship between pollinator traits and PE. Our objectives were to determine which morphological and behavioural traits promote PE and to establish whether effective pollinators shared similar traits. For each flower visit we recorded duration, flower handling behaviour (top-working vs. side-working: flower visited from the top or from the side, respectively) and resource collected (pollen vs. nectar). For each pollinator species we measured body size, tongue length, hair density and hair length. PE depended on flower handling behaviour (top visitors were more effective than side visitors) and was positively correlated with body size and visit duration, but not with hairiness. We found large differences in PE among 11 pollinator species. The mean number of pollen grains deposited per visit ranged from 201.5 (Apis mellifera) to 833.7 (Andrena flavipes). In general, solitary bees (Andrena, Eucera, Osmia) were more effective than social bees (Bombus, Apis). Dipterans and beetles (Eristalis, muscoid flies, Tropinota, Oxythyrea) showed high to intermediate levels of PE. All effective pollinators were top-workers, but otherwise did not necessarily share similar traits. We conclude that there are different ways to be an effective pollinator. Our results provide a mechanism to explain the posi-tive relationship between pollinator diversity and pollination service and can be used to predict PE in bowl-shaped flowers. Our approach can be easily adapted to other flower-pollinator systems.

Functional traits mediate the response of communities to disturbances (response traits) and their contribution to ecosystem functions (effect traits). To predict how anthropogenic disturbances influence ecosystem services requires a dual approach including both trait concepts. Here, we used a response-effect trait conceptual framework to understand how local and landscape features affect pollinator functional diversity and pollination services in apple orchards. We worked in 110 apple orchards across four European regions. Orchards differed in management practices. Low-intensity (LI) orchards were certified organic or followed close-to-organic practices. High-intensity (HI) orchards followed integrated pest management practices. Within each management type, orchards encompassed a range of local (flower diversity, agri-environmental structures) and landscape features (orchard and pollinator-friendly habitat cover). We measured pollinator visitation rates and calculated trait composition metrics based on 10 pollinator traits. We used initial fruit set as a measure of pollination service. Some pollinator traits (body size and hairiness) were negatively related to orchard cover and positively affected by pollinator-friendly habitat cover. Bee functional diversity was lower in HI orchards and decreased with increased landscape orchard cover. Pollination service was not associated with any particular trait but increased with pollinator trait diversity in LI orchards. As a result, LI orchards with high pollinator trait diversity reached levels of pollination service similar to those of HI orchards. Synthesis and applications. Pollinator functional diversity enables pollinator communities to respond to agricultural intensification and to increase pollination function. Our results show that efforts to promote biodiversity provide greater returns in low-intensity than in high-intensity orchards. The fact that low-intensity orchards with high pollinator functional diversity reach levels of pollination services similar to those of high-intensity orchards provides a compelling argument for the conversion of high-intensity into low-intensity farms.

Pollinators face multiple pressures and there is evidence of populations in decline. As demand for insect-pollinated crops increases, crop production is threatened by shortfalls in pollination services. Understanding the extent of current yield deficits due to pollination and identifying opportunities to protect or improve crop yield and quality through pollination management is therefore of international importance. To explore the extent of "pollination deficits," where maximum yield is not being achieved due to insufficient pollination, we used an extensive dataset on a globally important crop, apples. We quantified how these deficits vary between orchards and countries and we compared "pollinator dependence" across different apple varieties. We found evidence of pollination deficits and, in some cases, risks of overpollination were even apparent for which fruit quality could be reduced by too much pollination. In almost all regions studied we found some orchards performing significantly better than others in terms of avoiding a pollination deficit and crop yield shortfalls due to suboptimal pollination. This represents an opportunity to improve production through better pollinator and crop management. Our findings also demonstrated that pollinator dependence varies considerably between apple varieties in terms of fruit number and fruit quality. We propose that assessments of pollination service and deficits in crops can be used to quantify supply and demand for pollinators and help to target local management to address deficits although crop variety has a strong influence on the role of pollinators.

Hairiness is a salient trait of insect pollinators that has been linked to thermoregulation, pollen uptake and transportation, and pollination success. Despite its potential importance in pollination ecology, hairiness is rarely included in pollinator trait analyses. This is likely due to the lack of standardized and efficient methods to measure hairiness. We describe a novel methodology that uses a stereomicroscope equipped with a live measurement module software to quantitatively measure two components of hairiness: hair density and hair length. We took measures of the two hairiness components in 109 insect pollinator species (including 52 bee species). We analyzed the relationship between hair density and length and between these two components and body size. We combined hair density and length measures to calculate a hairiness index and tested whether hairiness differed between major pollinator groups and bee genera. Body size was strongly and positively correlated to hair length and weakly and negatively correlated to hair density. The correlation between the two hairiness components was weak and negative. According to our hairiness index, butterflies and moths were the hairiest pollinator group, followed by bees, hoverflies, beetles, and other flies. Among bees, bumblebees (Bombus) and mason bees (Osmia) were the hairiest taxa, followed by digger bees (Anthophorinae), sand bees (Andrena), and sweat bees (Halictini). Our methodology provides an effective and standardized measure of the two components of hairiness (hair density and length), thus allowing for a meaningful interpretation of hairiness. We provide a detailed protocol of our methodology, which we hope will contribute to improve our understanding of pollination effectiveness, thermal biology, and responses to climate change in insects.

Local agri-environmental schemes, including hedgerows, flowering strips, organic management, and a landscape rich in semi-natural habitat patches, are assumed to enhance the presence of beneficial arthropods and their contribution to biological control in fruit crops. We studied the influence of local factors (orchard management and adjacent habitats) and of landscape composition on the abundance and community composition of predatory arthropods in apple orchards in three European countries. To elucidate how local and landscape factors influence natural enemy effectiveness in apple production systems, we calculated community energy use as a proxy for the communities' predation potential based on biomass and metabolic rates of predatory arthropods. Predator communities were assessed by standardised beating samples taken from apple trees in 86 orchards in Germany, Spain and Sweden. Orchard management included integrated production (IP; i.e. the reduced and targeted application of synthetic agrochemicals), and organic management practices in all three countries. Predator communities differed between management types and countries. Several groups, including beetles (Coleoptera), predatory bugs (Heteroptera), flies (Diptera) and spiders (Araneae) benefited from organic management depending on country. Woody habitat and IP supported harvestmen (Opiliones). In both IP and organic orchards we detected aversive influences of a high-quality surrounding landscape on some predator groups: for example, high covers of woody habitat reduced earwig abundances in German orchards but enhanced their abundance in Sweden, and high natural plant species richness tended to reduce predatory bug abundance in Sweden and IP orchards in Spain. We conclude that predatory arthropod communities and influences of local and landscape factors are strongly shaped by orchard management, and that the influence of management differs between countries. Our results indicate that organic management improves the living conditions for effective predator communities.

Apple is considered the most important fruit crop in temperate areas and profitable production depends on multiple ecosystem services, including the reduction of pest damage and the provision of sufficient pollination levels. Management approaches present an inherent trade‐off as each affects species differently. We quantified the direct and indirect effects of management (organic vs. integrated pest management, IPM) on species richness, ecosystem services, and fruit production in 85 apple orchards in three European countries. We also quantified how habit composition influenced these effects at three spatial scales: within orchards, adjacent to orchards, and in the surrounding landscape. Organic management resulted in 48% lower yield than IPM, and also that the variation between orchards was large with some organic orchards having a higher yield than the average yield of IPM orchards. The lower yield in organic orchards resulted directly from management practices, and from higher pest damage in organic orchards. These negative yield effects were partly offset by indirect positive effects from more natural enemies and higher flower visitation rates in organic orchards. Two factors other than management affected species richness and ecosystem services. Higher cover of flowering plants within and adjacent to the apple trees increased flower visitation rates by pollinating insects and a higher cover of apple orchards in the landscape decreased species richness of beneficial arthropods. The species richness of beneficial arthropods in orchards was uncorrelated with fruit production, suggesting that diversity can be increased without large yield loss. At the same time, organic orchards had 38% higher species richness than IPM orchards, an effect that is likely due to differences in pest management. Synthesis and applications. Our results indicate that organic management is more efficient than integrated pest management in developing environmentally friendly apple orchards with higher species richness. We also demonstrate that there is no inherent trade‐off between species richness and yield. Development of more environmentally friendly means for pest control, which do not negatively affect pollination services, needs to be a priority for sustainable apple production.

Organic management, connective woody habitats, and landscape complexity are supposed to enhance beneficial arthropods and biological pest control in agro-ecosystems. We studied earwigs (Dermaptera: Forficulidae) as generalist predators and aphids (Hemiptera: Aphididae) as key pests serving as earwig prey in a total of 58 commercial apple orchards differing in management (integrated production (IP) versus organic) in Germany and Spain. We focused on the effects of local agri-environmental structures, orchard management, and composition of the surrounding landscape on earwig populations and on tree infestation by the woolly apple aphid (WAA), Eriosoma lanigerum. Surprisingly, the common earwig, Forficula auricularia, did not benefit from organic management in either country, and we found even slightly higher earwig abundances in IP than in organic orchards in Germany. In Spain, we found a negative impact of IP compared to organic management on abundance of the earwig Forficula pubescens, whereas orchard management did not affect the abundance of F. auricularia. The presence of woody habitats adjacent to the orchard reduced the abundance of F. auricularia in IP but not in organic orchards in Germany. We did not study the effects of woody habitats in Spain, where these structures were very scarce. There was no effect of high plant species richness at the orchard boundary or compositional landscape heterogeneity on earwig abundance in either country. In Germany, WAA infestation was very low and driven by landscape characteristics rather than orchard management. In Spain, WAA infestation differed strongly between management types (higher in organic orchards). There were no strong, consistent correlations between earwig abundance and WAA infestation in either country. Our study shows that adjacent woody structures and orchard management may affect earwigs in perennial cropping systems. The consequences of orchard management, however, seem to strongly depend on earwig species. Our study suggests that woody elements may serve as sink habitats potentially attracting earwigs by providing alternative prey and shelter in IP (but not in organic) orchards.