Abstract In an agricultural landscape, production and conservation ideally go hand in hand. In a win‐win scenario, conservation measures, such as hedges and flower strips, provide support for biodiversity and crop production for example by increased pollination and pollinator diversity. However, these conservation measures may also decrease pollination by attracting pollinators and competing with crop flower visits. Here, we studied plant–pollinator interactions from two different perspectives. First, we looked at the production perspective investigating whether plant–pollinator networks differed between apple orchards with and without adjacent flower strips and hedges. With help of the Bayes factor, we investigated similarity and conclude that there are no differences between pollination networks with or without adjacent flower strips and hedges. Second, we looked at the conservation perspective and analyzed the impact of hedges and flower strips on pollinators and their interactions with plants before and after apple bloom in April. We showed that apple pollinators used more flower resources in flower strips and hedges across the whole season compared to isolated orchards. In orchards with flower strips and hedges, interactions were more constant over time. We conclude that flower strips and hedges are beneficial for conservation of apple pollinators without being harmful for apple flower pollination.

Plant-pollinator interactions are ecologically and economically important, and, as a result, their prediction is a crucial theoretical and applied goal for ecologists. Although various analytical methods are available, we still have a limited ability to predict plant-pollinator interactions. The predictive ability of different plant- pollinator interaction models depends on the specific definitions used to conceptualize and quantify species attributes (e.g., morphological traits), sampling effects (e.g., detection probabilities), and data resolution and availability. Progress in the study of plant-pollinator interactions requires conceptual and methodological advances concerning the mechanisms and species attributes governing interactions as well as improved modeling approaches to predict interactions. Current methods to predict plant-pollinator interactions present ample opportunities for improvement and spark new horizons for basic and applied research.

Tree diversity can promote both predator abundance and diversity. However, whether this translates into increased predation and top-down control of herbivores across predator taxonomic groups and contrasting environmental conditions remains unresolved. We used a global network of tree diversity experiments (TreeDivNet) spread across three continents and three biomes to test the effects of tree species richness on predation across varying climatic conditions of temperature and precipitation. We recorded bird and arthropod predation attempts on plasticine caterpillars in monocultures and tree species mixtures. Both tree species richness and temperature increased predation by birds but not by arthropods. Furthermore, the effects of tree species richness on predation were consistent across the studied climatic gradient. Our findings provide evidence that tree diversity strengthens top-down control of insect herbivores by birds, underscoring the need to implement conservation strategies that safeguard tree diversity to sustain ecosystem services provided by natural enemies in forests.

It is commonly expected that exotic plants experience reduced herbivory, but experimental evidence for such enemy release is still controversial. One reason for conflicting results might be that community context has rarely been accounted for, although the surrounding plant diversity may moderate enemy release. Here, we tested the effects of focal tree origin and surrounding tree diversity on herbivore abundance and leaf damage in a cross-Atlantic tree-diversity experiment in Canada and Germany. We evaluated six European tree species paired with six North American congeners in both their native and exotic range, expecting lower herbivory for the exotic tree species in each pair at each site. Such reciprocal experiments have long been called for, but have not been realized thus far. In addition to a thorough evaluation of overall enemy release effects, we tested whether enemy release effects changed with the surrounding tree diversity. Herbivore abundance was indeed consistently lower on exotics across all six tree genera (12 comparisons). This effect of exotic status was independent of the continent, phylogenetic relatedness, and surrounding tree diversity. In contrast, leaf damage associated with generalist leaf chewers was consistently higher on North American tree species. Interestingly, several species of European weevils were the most abundant leaf chewers on both continents and the dominant herbivores at the Canadian site. Thus, most observed leaf damage is likely to reflect the effect of generalist herbivores that feed heavily on plant species with which they have not evolved. At the German site, sap suckers were the dominant herbivores and showed a pattern consistent with enemy release. Taken together, the consistently lower herbivory on exotics on both continents is not purely a pattern of enemy release in the strictest sense, but to some degree additionally reflects the susceptibility of native plants to invasive herbivores. In conclusion, our cross-Atlantic study is consistent with the idea that nonnative trees have generally reduced herbivory, regardless of tree community diversity and species identity, but for different reasons depending on the dominant herbivore guild.

Plant–frugivore networks often display vertical structure in forest communities because of vertical distribution of fruit resources and animal feeding niches on and under trees. However, how vertical stratification of fruit sources and animal feeding niches affect the structure and functioning of frugivory networks has not previously been well studied. We used infrared cameras under trees and direct observations on trees to extensively sample the whole frugivory networks in a subtropical forest in Southwest China. We analyzed the structure of on‐the‐tree, under‐the‐tree, space‐free aggregated (where links lacked vertical spatial information), and space‐based aggregated frugivory networks (where links included vertical spatial information). We uncovered vertical differentiation in frugivore composition (dissimilarity > 0.7) and less than 5% overlap of links occurring both under and on trees. Network analyses revealed high complementarity between the under‐ and on‐the‐tree frugivory networks. Modularity and specialization increased when space‐based hidden links (i.e. considering spatial locations) were included. Overall, we conclude that consideration of vertical structure (on‐the‐tree versus under‐the‐tree) in frugivore networks can enhance our understanding of the seed dispersal process and frugivore co‐existence in forests.

Arable plants and wild bees are negatively affected by agricultural intensification, one of the major drivers of global biodiversity loss. However, it remains unclear how endangered and low competitive arable plants (rare arable plants) contribute to the persistence of flower-visiting wild bees by providing additional flower resources in agricultural landscapes. Thus, the effects of sowing 10 rare arable plant species on wild bees were investigated in an experimental field and on 10 different arable farms on nutrient-poor soils. Sowing of rare arable plants on cropped and uncropped plots was compared to annual and perennial wildflower strips. Results showed that rare arable plants on uncropped plots attracted as many wild bees as wildflower strips. Wild bee abundance and species richness increased in the autumn-sown crops in the second year, likely because winter annual rare arable plants were preferred. In particular, rare arable plants provided flowers preferred by long-tongued bumblebees, which are often lacking in intensively managed arable fields. Our study shows that sowing of rare arable plants can increase niche diversity and therefore resource availability for wild bees, and it can also conserve diversity of arable plants in degraded agricultural landscapes. Conservation of arable plants through sowing can also support wild bee communities and may become an important tool in pollinator-friendly management of arable land.

Ecological interactions link species in networks. Loss of species from or introduction of new species into an existing network may have substantial effects for interaction patterns. Predicting changes in interaction frequency while allowing for rewiring of existing interactions-and hence estimating the consequences of community compositional changes-is thus a central challenge for network ecology. Interactions between species groups, such as pollinators and flowers or parasitoids and hosts, are moderated by matching morphological traits or sensory clues, most of which are unknown to us. If these traits are phylogenetically conserved, however, we can use phylogenetic distances to construct latent, surrogate traits and try to match those across groups, in addition to observed traits. Understanding how important traits and trait matching are, relative to abundances and chance, is crucial to estimating the fundamental predictability of network interactions. Here, we present a statistically sound approach ("tapnet") to fitting abundances, traits, and phylogeny to observed network data to predict interaction frequencies. We thereby expand existing approaches to quantitative bipartite networks, which so far have failed to correctly represent the nonindependence of network interactions. Furthermore, we use simulations and cross validation on independent data to evaluate the predictive power of the fit. Our results show that tapnet is on a par with abundance-only, matching centrality, and machine learning approaches. This approach also allows us to evaluate how well current concepts of trait matching work. On the basis of our results, we expect that interactions in well-sampled networks can be well predicted if traits and abundances are the main driver of interaction frequency.

While an increasing number of studies indicate that the range, diversity and abundance of many wild pollinators has declined, the global area of pollinator-dependent crops has significantly increased over the last few decades. Crop pollination studies to date have mainly focused on either identifying different guilds pollinating various crops, or on factors driving spatial changes and turnover observed in these communities. The mechanisms driving temporal stability for ecosystem functioning and services, however, remain poorly understood. Our study quantifies temporal variability observed in crop pollinators in 21 different crops across multiple years at a global scale. Using data from 43 studies from six continents, we show that (i) higher pollinator diversity confers greater inter-annual stability in pollinator communities, (ii) temporal variation observed in pollinator abundance is primarily driven by the three-most dominant species, and (iii) crops in tropical regions demonstrate higher inter-annual variability in pollinator species richness than crops in temperate regions. We highlight the importance of recognizing wild pollinator diversity in agricultural landscapes to stabilize pollinator persistence across years to protect both biodiversity and crop pollination services. Short-term agricultural management practices aimed at dominant species for stabilizing pollination services need to be considered alongside longer term conservation goals focussed on maintaining and facilitating biodiversity to confer ecological stability.

The world is astoundingly variable, and individuals to whole communities must respond to variability to survive. One example of nature’s variability is the massive fluctuations in spruce budworm (Choristoneura fumiferana Clemens, Lepidoptera: Tortricidae) populations that occur over 35 years. We examined how the parasitoid community altered its parasitism of budworm and other caterpillar species in response to these fluctuations. Budworm and other caterpillar species were sampled from balsam fir in three plots for 14 years in Atlantic Canada, and then reared to identify any emerging parasitoids. We found that the parasitoid community showed a simple linear, indiscriminate response (i.e., no preference, where densities purely dictated parasitism rates) to changes in budworm densities relative to other caterpillar species on balsam fir. We also observed strong changes in topology and distributions of interaction strengths between the parasitoids, budworm and other caterpillar species as budworm densities fluctuated. Our study contributes to the suggestion that hardwood trees are a critical part of the budworm-parasitoid food web, where parasitoids attack other caterpillar species on hardwood trees when budworm populations are low. Taken together, our study shows that a parasitoid community collectively alters species interactions in response to variable budworm densities, fundamentally shifting food web pathways.

In recent decades, European temperate forests have repeatedly suffered from severe droughts. Drought‐weakened forests have often become more susceptible to pest outbreaks such as bark beetle infestations. Tree species diversity is expected to increase resistance to drought and pests, but evidence for a positive tree diversity effect on insect pest reduction is largely circumstantial. Here we tested the effects of tree diversity and biogeographic origin of tree species on bark beetle infestation in a large, young tree diversity experiment, with six broadleaved and six conifer species from Europe and North America. Lower infestation risk was expected for the exotic tree species in each congeneric pair (spruce, larch and pine) and for mixtures with higher species richness and higher broadleaf proportion. Following a severe drought in summer 2018, the conifer trees were attacked by the six‐toothed spruce bark beetle Pityogenes chalcographus . Bark beetle boreholes were recorded in winter 2018/2019 on all conifer species. Norway spruce Picea abies and European larch Larix decidua were the most infested species and thus considered main hosts of the bark beetle. For these two species, probability of infestation decreased with increasing tree diversity (although this was only significant for Larix ). In contrast, Pinus , which were less infested overall, were more likely to be infested in plots with high tree diversity. Exotic trees tended to be less infested, with clearest support for enemy release found at the level of infestation intensity when considering pure conifer stands. Overall, the effects of tree diversity and tree species origin were not as strong as the effect of position within the experimental site, where higher rates of infestation were observed at the edge than in the centre. Synthesis . Increasing tree diversity may reduce the risk of bark beetle infestation for genera prone to high infestation rates ( Picea and Larix ), but risk for less preferred genera ( Pinus , and to some extent the exotic tree species) may increase with tree diversity due to spillover from preferred hosts. In mixed forests, the risk of infestation, even by relatively specialized insect pests, may be redistributed among tree species rather than reduced for all.

Temporal variability of plant–pollinator interactions is important for fully understanding the structure, function, and stability of plant–pollinator networks, but most network studies so far have ignored within-day dynamics. Strong diel dynamics (e.g., a regular daily cycle) were found for networks with Cichorieae, which typically close their flowers around noon. Here, we experimentally prevented early flower closure to test whether these dynamics are driven by the temporally limited availability of Cichorieae, or by timing of pollinator activity. We further tested if the dynamics involving Cichorieae and their pollinators also affect the dynamics on other plants in the network. Finally, we explored the structure of such manipulated networks (with Cichorieae available in the morning and afternoon) compared to unmanipulated controls (Cichorieae available only in the morning). We found that flower closure of Cichorieae is indeed an important driver of diel network dynamics, while other drivers of pollinator timing appeared less important. If Cichorieae flowers were available in the afternoon, they were visited by generalist and specialist pollinators, which overall decreased link turnover between morning and afternoon. Effects of afternoon availability of Cichorieae on other plants in the network were inconclusive: pollinator switching to and from Cichorieae tended to increase. On the level of the aggregated (full-day) network, the treatment resulted in increased dominance of Cichorieae, reducing modularity and increasing plant generality. These results highlight that network dynamics can be predicted by knowledge of diel or seasonal phenology, and that fixed species timing assumptions will misrepresent the expected dynamics.

AbstractDescribing variation of species interaction networks across space and time promises a better understanding of how species communities respond to global change. To understand this variation, it has been suggested to partition network dissimilarity into one component driven by species turnover, that is, changes in community composition, and another component reflecting rewiring, that is, flexibility of interactions among shared species. The latter makes a strong case for investing the enormous effort in empirically recording interactions, instead of simply building networks based on community data. Here, I present a flexible R function (available in the R package bipartite) to calculate network dissimilarity and its components, with binary and quantitative networks. With this new tool, I compare two published methods for partitioning network dissimilarity, using conceptual examples, published plant–pollinator networks, and a set of simulations. This comparison highlights that the method that has received most attention overestimates the importance of rewiring for total network dissimilarity. In contrast, an earlier‐proposed method is derived from additive partitioning of the sets of interactions and thus accurately represents the two dissimilarity components. Furthermore, I argue that the term rewiring is not well defined in network ecology and that there are reasons why both methods overestimate the importance of rewiring, in particular with quantitative networks. The availability of a unified function to calculate multiple aspects of network dissimilarity will foster its critical application to characterize network dynamics and to identify underlying drivers. Studies on network dissimilarity and rewiring will have to be more careful in the choice of method and its interpretation.

Most studies of plant-animal mutualistic networks have come from a temporally static perspective. This approach has revealed general patterns in network structure, but limits our ability to understand the ecological and evolutionary processes that shape these networks and to predict the consequences of natural and human-driven disturbance on species interactions. We review the growing literature on temporal dynamics of plant-animal mutualistic networks including pollination, seed dispersal and ant defence mutualisms. We then discuss potential mechanisms underlying such variation in interactions, ranging from behavioural and physiological processes at the finest temporal scales to ecological and evolutionary processes at the broadest. We find that at the finest temporal scales (days, weeks, months) mutualistic interactions are highly dynamic, with considerable variation in network structure. At intermediate scales (years, decades), networks still exhibit high levels of temporal variation, but such variation appears to influence network properties only weakly. At the broadest temporal scales (many decades, centuries and beyond), continued shifts in interactions appear to reshape network structure, leading to dramatic community changes, including loss of species and function. Our review highlights the importance of considering the temporal dimension for understanding the ecology and evolution of complex webs of mutualistic interactions.

Understanding how biodiversity and interaction networks change across environmental gradients is a major challenge in ecology. We integrated metacommunity and metanetwork perspectives to test species' functional roles in bird-plant frugivory interactions in a fragmented forest landscape in Southwest China, with consequences for seed dispersal. Availability of fruit resources both on and under trees created vertical feeding stratification for frugivorous birds. Bird-plant interactions involving birds feeding only on-the-tree or both on and under-the-tree (shared) had a higher centrality and contributed more to metanetwork organisation than interactions involving birds feeding only under-the-tree. Moreover, bird-plant interactions associated with large-seeded plants disproportionately contributed to metanetwork organisation and centrality. Consequently, on-the-tree and shared birds contributed more to metanetwork organisation whereas under-the-tree birds were more involved in local processes. We would expect that species' roles in the metanetwork will translate into different conservation values for maintaining functioning of seed-dispersal networks.

The study of mutualistic interaction networks has led to valuable insights into ecological and evolutionary processes. However, our understanding of network structure may depend upon the temporal scale at which we sample and analyze network data. To date, we lack a comprehensive assessment of the temporal scale‐dependence of network structure across a wide range of temporal scales and geographic locations. If network structure is temporally scale‐dependent, networks constructed over different temporal scales may provide very different perspectives on the structure and composition of species interactions. Furthermore, it remains unclear how various factors – including species richness, species turnover, link rewiring and sampling effort – act in concert to shape network structure across different temporal scales. To address these issues, we used a large database of temporally‐resolved plant–pollinator networks to investigate how temporal aggregation from the scale of one day to multiple years influences network structure. In addition, we used structural equation modeling to explore the direct and indirect effects of temporal scale, species richness, species turnover, link rewiring and sampling effort on network structural properties. We find that plant–pollinator network structure is strongly temporally‐scale dependent. This general pattern arises because the temporal scale determines the degree to which temporal dynamics (i.e. phenological turnover of species and links) are included in the network, in addition to how much sampling effort is put into constructing the network. Ultimately, the temporal scale‐dependence of our plant–pollinator networks appears to be mostly driven by species richness, which increases with sampling effort, and species turnover, which increases with temporal extent. In other words, after accounting for variation in species richness, network structure is increasingly shaped by its underlying temporal dynamics. Our results suggest that considering multiple temporal scales may be necessary to fully appreciate the causes and consequences of interaction network structure.

Biodiversity-ecosystem functioning (BEF) research grew rapidly following concerns that biodiversity loss would negatively affect ecosystem functions and the ecosystem services they underpin. However, despite evidence that biodiversity strongly affects ecosystem functioning, the influence of BEF research upon policy and the management of ‘real-world’ ecosystems, i.e., semi-natural habitats and agroecosystems, has been limited. Here, we address this issue by classifying BEF research into three clusters based on the degree of human control over species composition and the spatial scale, in terms of grain, of the study, and discussing how the research of each cluster is best suited to inform particular fields of ecosystem management. Research in the first cluster, small-grain highly controlled studies, is best able to provide general insights into mechanisms and to inform the management of species-poor and highly managed systems such as croplands, plantations, and the restoration of heavily degraded ecosystems. Research from the second cluster, small-grain observational studies, and species removal and addition studies, may allow for direct predictions of the impacts of species loss in specific semi-natural ecosystems. Research in the third cluster, large-grain uncontrolled studies, may best inform landscape-scale management and national-scale policy. We discuss barriers to transfer within each cluster and suggest how new research and knowledge exchange mechanisms may overcome these challenges. To meet the potential for BEF research to address global challenges, we recommend transdisciplinary research that goes beyond these current clusters and considers the social-ecological context of the ecosystems in which BEF knowledge is generated. This requires recognizing the social and economic value of biodiversity for ecosystem services at scales, and in units, that matter to land managers and policy makers.

Agricultural intensification can impact agrobiodiversity in several ways such as in terms of population densities, community composition and food web interactions across all trophic levels. This effect can be investigated at two scales: field-scale and landscape scale. Here it was assessed how the impact of fertilisation (within field) and landscape complexity (within landscape) impact cereal aphid-primary parasitoid-secondary parasitoid systems in winter wheat in Germany. A newly developed molecular technique was used to quantify species-specific linkages between aphids, primary parasitoids and secondary parasitoids sampled in fertilised and unfertilised plots in either simple or complex structured landscapes. The results show a stronger effect of fertilisation than landscape complexity on the groups: fertilisation positively affected the crop plants while it negatively affected both the density of the cereal aphid Sitobion avenae and its primary parasitism rates whereas no effect on the level of secondary parasitism rates was observed. Landscape complexity had no effect on plants, aphids, as well as on primary parasitism rate. In case of secondary parasitism rate there was an effect in interaction with sampling date. Field identity accounted for the strongest effect on parasitoid community composition (10.6% of the variance) from all tested variables, while fertilisation and landscape complexity had almost no effect (1.1% and no effect). Nevertheless, a weak cascading effect of both environmental factors could be observed as the primary-secondary parasitoid network structure responded to both. However, these observed effects on food webs strongly depended on species identity, highlighting the need of species-level food web assessment.

Defaunation by humans causes a loss of large animals in many ecosystems globally. Recent work has emphasized the consequences of downsizing in animal communities for ecosystem functioning. However, no study so far has integrated network theory and life-history trade-offs to mechanistically evaluate the functional consequences of defaunation in plant–animal networks. Here, we simulated an avian seed-dispersal network and its derived ecosystem function seedling recruitment to assess the relative importance of different size-related mechanisms. Specifically, we considered size matching (between bird size and seed size) and size trade-offs, which are driven by differences in plant or animal species abundance (negative size–quantity relationship) as well as in recruitment probability and disperser quality (positive size–quality relationship). Defaunation led to impoverished seedling communities in terms of diversity and seed size, but only if models accounted for size matching. In addition, size trade-off in plants, in concert with size matching, provoked rapid decays in seedling abundance in response to defaunation. These results underscore a disproportional importance of large animals for ecosystem functions. Downsizing in ecological networks will have severe consequences for ecosystem functioning, especially in interaction networks that are structured by size matching between plants and animals.