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The idea that noncrop habitat enhances pest control and represents a win–win opportunity to conserve biodiversity and bolster yields has emerged as an agroecological paradigm. However, while noncrop habitat in landscapes surrounding farms sometimes benefits pest predators, natural enemy responses remain heterogeneous across studies and effects on pests are inconclusive. The observed heterogeneity in species responses to noncrop habitat may be biological in origin or could result from variation in how habitat and biocontrol are measured. Here, we use a pest-control database encompassing 132 studies and 6,759 sites worldwide to model natural enemy and pest abundances, predation rates, and crop damage as a function of landscape composition. Our results showed that although landscape composition explained significant variation within studies, pest and enemy abundances, predation rates, crop damage, and yields each exhibited different responses across studies, sometimes increasing and sometimes decreasing in landscapes with more noncrop habitat but overall showing no consistent trend. Thus, models that used landscape-composition variables to predict pest-control dynamics demonstrated little potential to explain variation across studies, though prediction did improve when comparing studies with similar crop and landscape features. Overall, our work shows that surrounding noncrop habitat does not consistently improve pest management, meaning habitat conservation may bolster production in some systems and depress yields in others. Future efforts to develop tools that inform farmers when habitat conservation truly represents a win–win would benefit from increased understanding of how landscape effects are modulated by local farm management and the biology of pests and their enemies., This work was supported through the National Socio-Environmental Synthesis Center (SESYNC) National Science Foundation Award DBI-1052875 for the project “Evidence and Decision-Support Tools for Controlling Agricultural Pests with Conservation Interventions” organized by D.S.K. and R.C.-K

In patchy environments, such as agricultural landscapes, both spatial and temporal scales of habitat heterogeneity can affect population dynamics and trophic interactions. As a result of crop rotation, landscapes and local resource availability may change dramatically within and between years. We used a tritrophic interaction constituted by pollen beetles, their host plant oilseed rape, and their parasitoids, as a model system to investigate how the effect of landscape composition on insect abundance changes with time and whether system dynamics showed carry-over effects of previous years. We employ path analysis models that allow us to study whole networks of hypotheses rather than univariate cause–effect relationships. We exposed pan traps in a 5 × 5 grid design within 10 landscapes in June 2011 (after oilseed rape flowering) and May 2012 (at peak oilseed rape flowering). Additionally, we assessed parasitism rates of pollen beetle larvae in May 2011 and measured changes in landscape composition. The effect of the oilseed rape proportion on beetle abundance changed with time from negative (during flowering) to positive (after flowering). Parasitism had a negative effect on the number of newly emerged pollen beetles, but only in landscapes with a low proportion of oilseed rape. Interestingly, our path analysis showed that landscape composition affected herbivore abundance 1 or 2 years later, mediated by changes in parasitism. Our results suggest that plant–herbivore–parasitoid interactions in dynamic agricultural landscapes can show interannual carry-over effects, as they are affected by landscape composition and top-down effects in previous years.

AimBiodiversity across the globe is heavily eroded by intensified management at local and landscape scales. Species communities of calcareous grasslands, which are among Europe's most diverse habitats, are severely threatened by the cessation of appropriate traditional management, loss of habitat connectivity and simplification of the surrounding landscape. However, our understanding of these often interrelated factors remains limited, in particular for trait-mediated responses across taxa. Here, we test the independent effects of local management (grazing, mowing and abandonment), habitat connectivity (measured by a connectivity index) and landscape complexity (indicated by the percentage of arable land) on nine taxa: plants, butterflies, bees, grasshoppers, hoverflies, spiders, true bugs, rove beetles and leafhoppers on small semi-natural calcareous grassland remnants (

Avian ecosystem services such as the suppression of pests are considered to be of high ecological and economic importance in a range of ecosystems, especially in tropical agroforestry. However, how bird predation success is related to the diversity and composition of the bird community, as well as local and landscape factors, is poorly understood. We quantified arthropod predation in relation to the identity and diversity of insectivorous birds using experimental exposure of artificial, caterpillar-like prey in 15 smallholder cacao agroforestry systems differing in local shade-tree management and distance to primary forest. The bird community was assessed using both mist-netting (targeting active understorey insectivores) and point counts (higher completeness of species inventories). Bird predation was not related to local shade-tree management or overall bird species diversity, but to the activity of insectivorous bird species and the proximity to primary forest. Insectivore activity was best predicted by mist-netting-based data, not by point counts. We identified the abundant Indonesian endemic lemon-bellied white-eye Zosterops chloris as the main driver of predation on artificial prey.Synthesis and applications. The suppression of arthropods is a major ecosystem service provided by insectivorous birds in agricultural systems world-wide, potentially reducing herbivore damage on plants and increasing yields. Our results show that avian predation success can be driven by single and abundant insectivorous species, rather than by overall bird species richness. Forest proximity was important for enhancing the density of this key species, but did also promote bird species richness. Hence, our findings are both of economical as well as ecological interest because the conservation of nearby forest remnants will likely benefit human needs and biodiversity conservation alike. The suppression of arthropods is a major ecosystem service provided by insectivorous birds in agricultural systems world-wide, potentially reducing herbivore damage on plants and increasing yields. Our results show that avian predation success can be driven by single and abundant insectivorous species, rather than by overall bird species richness. Forest proximity was important for enhancing the density of this key species, but did also promote bird species richness. Hence, our findings are both of economical as well as ecological interest because the conservation of nearby forest remnants will likely benefit human needs and biodiversity conservation alike. (Less)

In fragmented farmland landscapes struc- tural complexity and low agricultural intensification should decrease the abundance of crop aphids due to increased abundances and species diversity of aphid enemies, including hymenopteran parasitoids. Here we study the effects of landscape structure and agricultural intensification on parasitism rates, abun- dances, and species richness of aphids and their parasitoids in five different regions in Europe. While total aphid numbers did not differ significantly among regions, we observed marked differences between Scandinavian and central European sites with respect to the species composition of aphids and their parasitoids and parasitism rates. In the cross country comparison landscape complexity and agricultural intensification did not significantly affect total aphid densities, although we observed species-specific reac- tions to land use. We also observed a tendency towards increased parasitoid species richness at low agricul- tural intensification but not at high landscape structure.

Summary Landscape heterogeneity represents two aspects of landscape simplification: (i) compositional heterogeneity (diversity of habitat types); and (ii) configurational heterogeneity (number, size and arrangement of habitat patches), both with different ecological implications for community composition. We examined how independent gradients of compositional and configurational landscape heterogeneity, at eight spatial scales, shape taxonomic and functional composition of butterfly communities in 91 managed grasslands across Germany. We used landscape metrics that were calculated from functional maps based on habitat preferences of individual species during different life stages. The relative effects of compositional and configurational landscape heterogeneity were compared with those of local land-use intensity on butterfly taxonomic diversity, community composition and functional diversity of traits related to body size, feeding breadth and migratory tendency. As expected, compositional heterogeneity had strong positive effects on taxonomic diversity, while configurational heterogeneity had strong positive effects on trait dominance within the community. When landscapes had smaller mean patch size and greater boundary area, communities were dominated by species with more specialized larval feeding, decreased forewing length and limited migratory tendency. The positive effects of increased configurational landscape heterogeneity outweighed the negative effects of local land-use intensity on larval-feeding specialization, at all spatial scales, highlighting its importance for specialists of all dispersal capabilities. Synthesis and applications. We show that landscapes with high compositional heterogeneity support communities with greater taxonomic diversity, while landscapes with high configurational heterogeneity support communities that include vulnerable species (feeding specialists with larger body size, sedentary nature and more negatively affected by local management intensity). A decline in functional community composition can lead to functional homogenization, affecting the viability of the ecosystems by decreasing the variability in their responses to disturbance and altering their functioning. A landscape management for grasslands that promotes the maintenance of small patch sizes and a diversity of land uses in the surrounding landscape (within 250–1000 m) is recommended for the conservation of diverse butterfly communities. These strategies could also benefit government programmes such as the EU 2020 Biodiversity Strategy in their efforts to reduce the loss of biodiversity in agricultural landscapes.

Habitat fragmentation is a primary threat to biodiversity, but how it affects the structure and stability of ecological networks is poorly understood. Here, we studied plant–pollinator and host–parasitoid networks on 32 calcareous grassland fragments covering a size gradient of several orders of magnitude and with amounts of additional habitat availability in the surrounding landscape that varied independent of fragment size. We find that additive and interactive effects of habitat fragmentation at local (fragment size) and landscape scales (1,750 m radius) directly shape species communities by altering the number of interacting species and, indirectly, their body size composition. These, in turn, affect plant–pollinator, but not host–parasitoid, network structure: the nestedness and modularity of plant–pollinator networks increase with pollinator body size. Moreover, pollinator richness increases modularity. In contrast, the modularity of host–parasitoid networks decreases with host richness, whereas neither parasitoid richness nor body size affects network structure. Simulating species coextinctions also reveals that the structure–stability relationship depends on species’ sensitivity to coextinctions and their capacity for adaptive partner switches, which differ between mutualistic and antagonistic interaction partners. While plant–pollinator communities may cope with future habitat fragmentation by responding to species loss with opportunistic partner switches, past effects of fragmentation on the current structure of host–parasitoid networks may strongly affect their robustness to coextinctions under future habitat fragmentation. Analysing the structure of both plant–pollinator and host–parasitoid networks in calcareous grasslands, the authors reveal scale-dependent responses to habitat fragmentation in the structure and stability of different network types.

Tropical tree crops such as cacao and coffee are produced around the tropics in diverse, multistrata agroforests as well as monoculture plantations Box 16.1 and references therein). The smallholders cultivating these systems battle pests and diseases that differ regionally and change over time, but often take a significant part of their yield, and therefore their revenue. In these perennial systems, ants are tremendously diverse and abundant, and affect pests and diseases directly as well as indirectly. Management by farmers of particular ant species to control insect pests has a long history (Offenberg, 2015). It is not until recently that the effects of ants on yields have been quantified. The complex interactions through which ants affect the crop plants, and how their mediation by species- and community-level characteristics, are starting to be better understood. The extent of the impact ants have on yields and revenue justifies the anthropocentric framing of the outcome of these interactions in terms of ecosystem services and disservices. In this chapter we present the current state of knowledge on agroforest ant communities, economically relevant ecological interactions driven by these communities and the way landscape-scale land-use change and climate change can be expected to influence ants and ant effects on insect communities and yields. Finally, we discuss how farmers may adapt their management to support ant-mediated ecosystem services and minimize potential disservices. We refer to Del Toro et al. (2012) and Choate and Drummond (2011) for more broad reviews of the role of ants in agriculture, as providers of biological control and other ecosystem services and disservices. Taxonomically and Functionally Rich Ant Communities. Ant surveys from cacao and coffee systems from throughout the range of these crops show a very high species richness that in most cases is comparable to that found in undisturbed forests (Table 16.1). Agroforests harbor arboreal and ground-dwelling ants. This includes species that nest in the canopy and trunk of the trees (dead wood, hollow twigs, foliage, sometimes with carton/silk/dirt nesting structures), in the herb layer, in the litter layer, on open ground, in epiphytic and parasitic plants, dead wood debris and other plant residues, such as dry cacao pods on the ground or on the tree (Room, 1971; De la Mora et al., 2013; Castano-Meneses et al., 2015). (Less)

Urban cultural ecosystem services are understood differently by experts and laypersons. Yet, unaccounted differences can lead to management problems for urban green spaces, as experts may recommend practices that do not meet the laypersons’ wishes. Qualitative research on the perception of cultural ecosystem services can be one tool to analyze these differences. We use expert and problem-centered interviews to assess differences in cultural ecosystem service perceptions for experts and laypersons in Berlin. Using an innovative approach, we combine inductive qualitative content analysis with a frequency analysis and multidimensional scaling. This explorative study innovatively merges qualitative and quantitative approaches to show new ways of analysis. Our results show that the experts’ perceptions of nature appear to be more practical, management-centered, whereas laypersons appear to prioritize enjoyment of nature. Overall, multidimensional scaling indicates different perceptions and conceptualizations of cultural ecosystem service bundles, emphasizing the diverging understandings. If these different perceptions are not accounted for it could lead to social and political contrast. They should therefore be acknowledged in decision-making and goal formulation for the management of urban green.

Closing yield gaps within existing croplands, and thereby avoiding further habitat conversions, is a prominently and controversially discussed strategy to meet the rising demand for agricultural products, while minimizing biodiversity impacts. The agricultural intensification associated with such a strategy poses additional threats to biodiversity within agricultural landscapes. The uneven spatial distribution of both yield gaps and biodiversity provides opportunities for reconciling agricultural intensification and biodiversity conservation through spatially optimized intensification. Here, we integrate distribution and habitat information for almost 20,000 vertebrate species with land-cover and land-use datasets. We estimate that projected agricultural intensification between 2000 and 2040 would reduce the global biodiversity value of agricultural lands by 11%, relative to 2000. Contrasting these projections with spatial land-use optimization scenarios reveals that 88% of projected biodiversity loss could be avoided through globally coordinated land-use planning, implying huge efficiency gains through international cooperation. However, global-scale optimization also implies a highly uneven distribution of costs and benefits, resulting in distinct "winners and losers" in terms of national economic development, food security, food sovereignty or conservation. Given conflicting national interests and lacking effective governance mechanisms to guarantee equitable compensation of losers, multinational land-use optimization seems politically unlikely. In turn, 61% of projected biodiversity loss could be avoided through nationally focused optimization, and 33% through optimization within just 10 countries. Targeted efforts to improve the capacity for integrated land-use planning for sustainable intensification especially in these countries, including the strengthening of institutions that can arbitrate subnational land-use conflicts, may offer an effective, yet politically feasible, avenue to better reconcile future trade-offs between agriculture and conservation. The efficiency gains of optimization remained robust when assuming that yields could only be increased to 80% of their potential. Our results highlight the need to better integrate real-world governance, political and economic challenges into sustainable development and global change mitigation research.

Effects of habitat fragmentation on species diversity and herbivore-parasitoid interactions were analyzed using the insect community of seed feeders and their parasitoids in the pods of the bush vetch (Vicia sepium L.). Field studies were carried out on 18 old meadows differing in area and isolation. The area of these meadows was found to be the major determinant of species diversity and population abundance of endophagous insects. Effects of isolation were further analyzed experimentally using 16 small plots with potted vetch plants isolated by 100–500 m from vetch populations on large old meadows. The results showed that colonization success greatly decreased with increasing isolation. In both cases, insect species were not equally affected. Parasitoids suffered more from habitat loss and isolation than their phytophagous hosts. Minimum area requirements, calculated from logistic regressions, were higher for parasitoids than for herbivores. In addition, percent parasitism of the herbivores significantly decreased with area loss and increasing isolation of Vicia sepium plots, supporting the trophic-level hypothesis of island biogeography. Species with high rates of absence on meadows and isolated plant plots were not only characterized by their high trophic level, but also by low abundance and high spatial population variability. Thus conservation of large and less isolated habitat remnants enhances species diversity and parasitism of potential pest insects, i.e., the stability of ecosystem functions.

Effects of habitat fragmentation on the insect community of stinging nettle (Urtica dioica L.) were studied, using 32 natural nettle patches of different area and degree of isolation in an agricultural landscape. Habitat fragmentation reduced the species richness of Heteroptera, Auchenorrhyncha, and Coleoptera, and the abundance of populations. Habitat isolation and area reduction did not affect all insect species equally. Monophagous herbivores had a higher probability of absence from small patches than all (monophagous and polyphagous) herbivore species, and the percentage of monophagous herbivores increased with habitat area. Abundance and population variability of species were negatively correlated and could both be used as a predictor of the percentage of occupied habitats. Species richness of herbivores correlated (positively) with habitat area, while species richness of predators correlated (negatively) with habitat isolation. In logistic regressions, the probability of absence of monophagous herbivores from habitat patches could only be explained by habitat area (in 4 out of 10 species) and predator absence probability only by habitat isolation (in 3 out of 14 species). Presumably because of the instability of higher-trophic-level populations and dispersal limitation, predators were more affected by habitat isolation than herbivores, while they did not differ from herbivore populations with respect to abundance or variability. Thus increasing habitat connectivity in the agricultural landscape should primarily promote predator populations.

1)Autonomous sound recording techniques have gained considerable traction in the last decade, but the question still remains whether they can replace human observation surveys to sample some animal taxa. Especially bird survey methods have been tested using classical point counts and autonomous sound recording techniques.2)We review the latest information by comparing both survey methods' standardization, verifiability, sampling completeness, data types, compatibility, and practicality by means of a systematic review and a meta-analysis of alpha and gamma species richness levels sampled by both methods across 20 separate studies.3)Although sound recording surveys have hitherto not enjoyed the most effective setups, they yield very similar results in terms of alpha and gamma species richness. We also reveal the crucial importance of the microphone (high signal-to-noise ratio) as the sensor that replaces human senses.4)We discuss key differences between both methods, while richness estimates are closely related and 81% of all species were detected by both methods. Sound recording techniques provide a more powerful and promising tool to monitor birds in a standardized, verifiable, and exhaustive way against the golden standard of point counts. Advantages include the capability of sampling continuously through day or season and of difficult-to-reach regions in an autonomous way, avoidance of observer bias and human disturbance effects and higher detection probability of rare species due to extensive recordings.

Agricultural intensification is a leading cause of global biodiversity loss, which can reduce the provisioning of ecosystem services in managed ecosystems. Organic farming and plant diversification are farm management schemes that may mitigate potential ecological harm by increasing species richness and boosting related ecosystem services to agroecosystems. What remains unclear is the extent to which farm management schemes affect biodiversity components other than species richness, and whether impacts differ across spatial scales and landscape contexts. Using a global metadataset, we quantified the effects of organic farming and plant diversification on abundance, local diversity (communities within fields), and regional diversity (communities across fields) of arthropod pollinators, predators, herbivores, and detritivores. Both organic farming and higher in‐field plant diversity enhanced arthropod abundance, particularly for rare taxa. This resulted in increased richness but decreased evenness. While these responses were stronger at local relative to regional scales, richness and abundance increased at both scales, and richness on farms embedded in complex relative to simple landscapes. Overall, both organic farming and in‐field plant diversification exerted the strongest effects on pollinators and predators, suggesting these management schemes can facilitate ecosystem service providers without augmenting herbivore (pest) populations. Our results suggest that organic farming and plant diversification promote diverse arthropod metacommunities that may provide temporal and spatial stability of ecosystem service provisioning. Conserving diverse plant and arthropod communities in farming systems therefore requires sustainable practices that operate both within fields and across landscapes., EML and DC were supported by the National Institute of Food and Agriculture, U.S. Department of Agriculture, under award number 2014‐51106‐22096. BMF was supported by the National Council for Scientific and Technological Development‐CNPq, Brasília, Brazil #305062/2007‐7. SGP and MO were supported by the Felix Trust and STEP Project (EC FP7 244090).

In agriculture, both valuable ecosystem services and unwanted ecosystem disservices can be produced by the same organism group. For example, small rodents can provide biological control through weed seed consumption but may also act as pests, causing crop damage. We studied the hypothesized causal relationships between ecosystem services (removal of weed seeds) and disservices (removal of wheat grains and crop damage) derived by small rodents (voles and mice) at multiple spatial scales. At the landscape scale, we studied the effects of landscape compositional and configurational heterogeneity on the abundance of voles and mice and their related ecosystem services and disservices along the former inner German border in east and west Germany. At the local scale, we studied how abundance and ecosystem functions are affected by management intensity (organic vs. conventional winter wheat), associated differences in crop characteristics and edge effects. Linear mixed-effects models and path analysis show that voles drove ecosystem disservices, but not ecosystem services, in agricultural fields. Daily wheat seed removal by voles was influenced by increasing wheat height and was almost three times higher than weed seed removal, which was not related to local- or landscape-scale effects. Abundance of voles and associated crop damage decreased with lower crop density and higher wheat height, which were associated with organic farming. Abundance of voles and crop damage were highest in conventional fields in west Germany. Synthesis and applications. As the ecosystem disservice of wheat seed consumption by small rodents must be considered mainly during crop sowing, management before crop harvest should focus on decreasing the pest potential of voles but not mice. Our results suggest that densities of voles and their ecosystem disservices could be reduced by having fields with low crop density and high wheat height, practices associated with organic farming. Surrounding landscapes with low compositional and configurational heterogeneity could further reduce voles’ pest potential, but with probable negative effects on farmland biodiversity.

In the past two decades, a large number of studies have investigated the relationship between biodiversity and ecosystem functioning, most of which focussed on a limited set of ecosystem variables. The Jena Experiment was set up in 2002 to investigate the effects of plant diversity on element cycling and trophic interactions, using a multi-disciplinary approach. Here, we review the results of 15 years of research in the Jena Experiment, focussing on the effects of manipulating plant species richness and plant functional richness. With more than 85,000 measures taken from the plant diversity plots, the Jena Experiment has allowed answering fundamental questions important for functional biodiversity research.[br/] First, the question was how general the effect of plant species richness is, regarding the many different processes that take place in an ecosystem. About 45% of different types of ecosystem processes measured in the 'main experiment', where plant species richness ranged from 1 to 60 species, were significantly affected by plant species richness, providing strong support for the view that biodiversity is a significant driver of ecosystem functioning. Many measures were not saturating at the 60-species level, but increased linearly with the logarithm of species richness. There was, however, great variability in the strength of response among different processes. One striking pattern was that many processes, in particular belowground processes, took several years to respond to the manipulation of plant species richness, showing that biodiversity experiments have to be long-term, to distinguish trends from transitory patterns. In addition, the results from the Jena Experiment provide further evidence that diversity begets stability, for example stability against invasion of plant species, but unexpectedly some results also suggested the opposite, e.g. when plant communities experience severe perturbations or elevated resource availability. This highlights the need to revisit diversity-stability theory.[br/] Second, we explored whether individual plant species or individual plant functional groups, or biodiversity itself is more important for ecosystem functioning, in particular biomass production. We found strong effects of individual species and plant functional groups on biomass production, yet these effects mostly occurred in addition to, but not instead of, effects of plant species richness.[br/] Third, the Jena Experiment assessed the effect of diversity on multitrophic interactions. The diversity of most organisms responded positively to increases in plant species richness, and the effect was stronger for above-than for belowground organisms, and stronger for herbivores than for carnivores or detritivores. Thus, diversity begets diversity. In addition, the effect on organismic diversity was stronger than the effect on species abundances.[br/] Fourth, the Jena Experiment aimed to assess the effect of diversity on N, P and C cycling and the water balance of the plots, separating between element input into the ecosystem, element turnover, element stocks, and output from the ecosystem. While inputs were generally less affected by plant species richness, measures of element stocks, turnover and output were often positively affected by plant diversity, e.g. carbon storage strongly increased with increasing plant species richness. Variables of the N cycle responded less strongly to plant species richness than variables of the C cycle.[br/] Fifth, plant traits are often used to unravel mechanisms underlying the biodiversity-ecosystem functioning relationship. In the Jena Experiment, most investigated plant traits, both above-and belowground, were plastic and trait expression depended on plant diversity in a complex way, suggesting limitation to using database traits for linking plant traits to particular functions.[br/] Sixth, plant diversity effects on ecosystem processes are often caused by plant diversity effects on species interactions. Analyses in the Jena Experiment including structural equation modelling suggest complex interactions that changed with diversity, e.g. soil carbon storage and greenhouse gas emission were affected by changes in the composition and activity of the belowground microbial community. Manipulation experiments, in which particular organisms, e.g. belowground invertebrates, were excluded from plots in split-plot experiments, supported the important role of the biotic component for element and water fluxes.[br/] Seventh, the Jena Experiment aimed to put the results into the context of agricultural practices in managed grasslands. The effect of increasing plant species richness from 1 to 16 species on plant biomass was, in absolute terms, as strong as the effect of a more intensive grassland management, using fertiliser and increasing mowing frequency. Potential bioenergy production from high-diversity plots was similar to that of conventionally used energy crops. These results suggest that diverse 'High Nature Value Grasslands' are multifunctional and can deliver a range of ecosystem services including production-related services.[br/] A final task was to assess the importance of potential artefacts in biodiversity-ecosystem functioning relationships, caused by the weeding of the plant community to maintain plant species composition. While the effort ( in hours) needed to weed a plot was often negatively related to plant species richness, species richness still affected the majority of ecosystem variables. Weeding also did not negatively affect monoculture performance; rather, monocultures deteriorated over time for a number of biological reasons, as shown in plant-soil feedback experiments.[br/] To summarize, the Jena Experiment has allowed for a comprehensive analysis of the functional role of biodiversity in an ecosystem. A main challenge for future biodiversity research is to increase our mechanistic understanding of why the magnitude of biodiversity effects differs among processes and contexts. It is likely that there will be no simple answer. For example, among the multitude of mechanisms suggested to underlie the positive plant species richness effect on biomass, some have received limited support in the Jena Experiment, such as vertical root niche partitioning. However, others could not be rejected in targeted analyses. Thus, from the current results in the Jena Experiment, it seems likely that the positive biodiversity effect results from several mechanisms acting simultaneously in more diverse communities, such as reduced pathogen attack, the presence of more plant growth promoting organisms, less seed limitation, and increased trait differences leading to complementarity in resource uptake. Distinguishing between different mechanisms requires careful testing of competing hypotheses. Biodiversity research has matured such that predictive approaches testing particular mechanisms are now possible.

Farmland biodiversity dramatically declined in Europe during the 20th century. Agri-environment schemes (AES) were introduced in the late 1980s in European Union countries as a solution to combat biodiversity decline. We examined the effectiveness of AES in enhancing biodiversity in a new EU member country (Estonia) over the period of 2010–2012. We compared species numbers and abundance of bumblebees and birds, plus cover of flowers, between three farming systems in two regions of Estonia. Farm types included conventional and two under AES (organic and a less strict environmentally friendly management agreement). Environmentally friendly management practices in Estonia include diversified crop rotations, at least 15% of arable land (including rotational grasslands) under legumes, permanent grassland strips, protection of landscape elements, reduced applications of agrochemicals, etc. The two selected regions (North and South) differed in landscape structure, soil types and crop yields. Flower cover and bird species richness and abundance without the dominant species (skylark) were higher on organic farms. Bumblebee species richness was significantly higher under both types of AES than under conventional farming. Flower cover, abundance and species richness of bumblebees and birds were significantly higher in the more heterogeneous landscapes of Southern Estonia. Environmentally friendly management may be a viable alternative to organic farming for a widely accepted, simple but large-scale greening of agricultural landscapes.

The modification of pristine wildlife habitats by land use is a worldwide cause of species extinction, as most native species cannot persist at high levels of human land use intensity. In this study, we gathered 90 lists of local interactions between herbivorous insects and their host plants to investigate the effects of land use intensity on the species richness and taxonomic diversity of insect herbivore assemblages. The effect of land use intensity on these insect herbivore assemblages was assessed using path analysis that controlled for differences in host plant richness, sampling effort, and the taxonomic range of plants and herbivores. We included the proportion of exotic host plant species in the path models to evaluate the extent to which the effect of land use intensity is mediated by the replacement of native with exotic host plants. Overall, we found negative effects of land use intensity on the species richness and taxonomic diversity of the insect herbivore assemblages, and these effects were mediated by an increase in the proportion of exotic host plant species. Since the effect of host plant richness was controlled for, our findings imply that the impoverishment of insect herbivore assemblages due to human land use is even greater than the loss of their host plant. A likely implication of such reduction in the species richness and taxonomic diversity of the insect herbivore assemblages is the accelerated loss of specialized plant-herbivore interactions, thus favoring interactions among generalist species and the biotic homogenization of species interactions across human-disturbed habitats.

Voluntary sustainability standards and certification offer a promising mechanism to mitigate the severe negative impacts of agricultural expansion and intensification on tropical biodiversity. From a conservation standpoint, certification of tropical agroforestry crops, especially coffee and cocoa, is of particular interest given the potentially high biodiversity value of agroforestry systems and the substantial market penetration of coffee and cocoa certification in recent years. Here, we review experience with coffee and cocoa certification, summarize evidence on conservation impacts, and explore future needs. While there is much evidence that environmental criteria behind certification support biodiversity conservation, it is less clear to what extent certification is the cause of improved conservation outcomes. Additionally, the farm-scale focus of current certification models may limit delivery of biodiversity conservation benefits, as maintenance of biodiversity depends on processes at larger landscape scales. To address this scale mismatch, we suggest that investment and innovation in certification over the next decade prioritize landscape conservation outcomes. This may be achieved by (1) linking existing certification mechanisms with broader landscape and ecosystem service management approaches and/or (2) expanding current certification models to consider the landscape itself as the certified unit.

Habitat fragmentation is a major driver of species loss. Here we test the hypotheses that high tree diversity in a large deciduous forest enhances bird diversity and nest survival. We further expect that forest edges support higher bird diversity when different habitat types adjoin, whereas nest predation is not higher, because the large forest area mitigates potential edge effects. We studied how edge-centre differences and tree diversity (beech-dominated vs. tree-species rich) affect the bird community and survival rates of ground breeding birds’ nests based on an artificial nest predation experiment in the Hainich National Park, Germany. We surveyed birds three times during the breeding season. We selected six forest stands with low tree diversity (i.e. dominated by beech) and six with high tree diversity (i.e. tree-species rich). Each forest stand contained four bird survey plots (plot 1: 0–30 m, plot 2: 60–90 m, plot 3: 120–150 m and plot 4: 180–210 m distant from edge; altogether 48 bird survey plots). Additionally each plot corner contained one artificial ground nest baited with one Blue-breasted Quail egg and one plasticine egg for eight days of exposure in the middle of the breeding season. Bird abundance and diversity were higher in the first 30 m of the forest. Bird diversity, including ground breeding birds, was also enhanced by higher percentages of bushes, which can provide enhanced food supply, perches as well as sheltering. Nest predation showed no edge effect, supporting the idea that small area of forest fragments causes more important negative effects than the edge in large forest remnants. Predation rates were higher in tree-species rich stands compared to beech-dominated stands, probably due to greater diversity and density of mammalian predators. Edge effects shaped the bird community composition and positively affected abundances of tree and shrub breeding birds, but did not affect ground breeders and the nest predation of ground nests. Shrub breeders accumulating in forest edges might, however, suffer more from nest predation in forest fragments. In conclusion, bird diversity and avian egg predation were affected by both forest edges and tree diversity in surprisingly different ways.

Intensification of agriculture reduces heterogeneity at local and landscape levels and thereby impact biodiversity and ecosystem processes. We studied a host-antagonist system of cavity-nesting bees, wasps and their antagonists and hypothesised that hosts and antagonists show different responses to local land-use intensity, the diversity of landscape in terms of composition and the spatial structure of landscape in terms of configuration. In a highly replicated study, we established nesting resources on 95 grasslands in three geographic regions across Germany and measured species richness and abundance of hosts (bees and wasps) and their antagonists, and rates of parasitism. For each site, we quantified local land-use intensity as well as landscape heterogeneity in terms of composition and configuration at spatial scales ranging from 250 m to 2000 m. Increasing landscape heterogeneity enhanced species richness, abundance and parasitism rate, whereas local land-use intensity only marginally negatively affected total abundance. Bee and wasp abundance as well as wasp species richness were enhanced by landscape composition at 250 m, whereas their antagonists were enhanced by landscape configuration at 1500 m. In conclusion, landscape composition and configuration affect trophic levels differently and are more relevant than local land-use intensity. Solitary bees and wasps, which offer important pollination and pest control services, could be supported by enhancing landscape diversity, while their antagonists could benefit from measures that promote landscape connectivity. Hence, scale-dependent and trophic group specific conservation management schemes are required, that address different components of landscape heterogeneity to enhance functional diversity and trophic interactions in agricultural landscapes.

Sustainable rice production is critical to food security especially in Asia. Effective biocontrol of major rice pests such as the White-Backed Planthopper (Sogatella furcifera, Horváth; WBP) is, hence, of eminent importance. We use newly compiled data from Thailand on the Wrinkle-Lipped Bat (Tadarida plicata, Buchanan), WBP distributions and an iterative modelling approach to quantify the importance of biological pest control by a common bat species on WBP. In Thailand, this single species interaction may prevent rice loss of almost 2,900 tons per year, which translates into a national economic value of more than 1.2. million USD or rice meals for almost 26,200 people annually. For the first time, our results show not only the critical importance of bat pest control services in economic terms, but also for sustaining food security. Thus, bat population decline as currently observed in Southeast Asia, will directly affect people by food and money. Functionally important populations, not just rare and endangered species, should be included in conservation management of human-dominated landscapes.

Summary 1. Human-dominated landscapes are characterized by a mosaic of natural and managed eco- systems, affecting arthropod communities on different spatial scales. Effective landscape man- agement for functionally important organisms suffers from little understanding of organism spillover between semi-natural habitats and adjacent crops, and of how it is affected by the surrounding landscape. 2. We examined syrphid abundance (Diptera: Syrphidae) in three types of linear semi-natural habitats, differing in connectedness to annual crops and forest [forest edges ( n = 12), forest- connected hedges ( n = 11) and isolated hedges ( n = 12)], as well as in the adjacent oilseed rape or winter wheat fields (i.e. altogether n = 70 sites in 35 landscapes). The landscape cir- cles with 1 km radius around the study sites differed in the proportion of oilseed rape (rang- ing from 0% to 35% oilseed rape) enabling us to test landscape-scale effects of oilseed rape. 3. Aphidophagous syrphids were more abundant in forest-connected hedgerows than in for- est edges (with isolated hedges being intermediate), and more abundant in crop fields adjacent to hedgerows than adjacent to forest edges, indicating spillover from semi-natural habitats to the adjacent crop fields. Aphidophagous syrphid abundance was higher in semi-natural habi- tats adjacent to oilseed rape fields than adjacent to wheat fields if the proportion of oilseed rape in the landscape was low (indicating local concentration). 4. Synthesis and applications. This study highlights the potential of hedgerows to enhance the abundances of beneficial syrphids and their spillover to adjacent crop fields, especially when they are connected with forests. We provide evidence that this local exchange is moderated by the extent of mass-flowering crops in the surrounding landscapes due to local concentration. There- fore, measurements for the improvement in local biological functioni ng should be evaluated by simultaneously investigating local and regio nal aspects of crop configurations to allow for region-specific management recommendations. Increasing the total amount of hedgerows in the agricultural matrix under moderate landscape- scale proportions of mass-flowering crops may serve best for the conservation of biodiversity and augmentation of i mportant ecosystem services such as biological control and pollination in lands capes dominated by agricultural cultivations.

Current biodiversity conservation policies have so far had limited success because they are mainly targeted to the scale of individual fields with little concern on different responses of organism groups at larger spatial scales. We investigated the relative impacts of multi-scale factors, including local land use intensity, landscape context and region, on functional groups of beetles (Coleoptera). In 2008, beetles were suction-sampled from 95 managed grasslands in three regions, ranging from Southern to Northern Germany. The results showed that region was the most important factor affecting the abundance of herbivores and the abundance and species composition of predators and decomposers. Herbivores were not affected by landscape context and land use intensity. The species composition of the predator communities changed with land use intensity, but only in interaction with landscape context. Interestingly, decomposer abundance was negatively related to land use intensity in low-diversity landscapes, whereas in high-diversity landscapes the relation was positive, possibly due to enhanced spillover effects in complex landscapes. We conclude that (i) management at multiple scales, from local sites to landscapes and regions, is essential for managing biodiversity, (ii) beetle predators and decomposers are more affected than herbivores, supporting the hypothesis that higher trophic levels are more sensitive to environmental change, and (iii) sustaining biological control and decomposition services in managed grassland needs a diverse landscape, while effects of local land use intensity may depend on landscape context.

Biodiversity experiments have shown that plant diversity has largely positive effects on insect diversity and abundance. However, such relationships have rarely been studied in undisturbed and more complex ecosystems such as forests. Flies (Diptera) are among the most dominant taxa in temperate ecosystems, influencing many ecosystem processes. As it is unknown how Diptera respond to changes in forest biodiversity, we examined how community characteristics of Diptera respond to varying levels of tree and herb diversity and vegetation structure. The study was conducted in the Hainich National Park (Central Germany) on 84 plots along a gradient of tree (from two to nine species) and herb (from two to 28 species) diversity. We found that herb and canopy cover as well as spatial effects were the best predictors of Diptera community composition, consisting of 62 families, including 99 Empidoidea and 78 Phoridae species. Abundance of Empidoidea was positively influenced by herb diversity, indicating bottom-up control. A complex causal pathway influenced Dipteran species richness: species-rich forest stands, with low beech cover, had lower canopy cover, resulting in higher Dipteran species richness. In addition, Diptera benefited from a more dense and diverse herb community. Individual species responded differentially to herb layer diversity, indicating that effects of plant diversity on higher trophic levels depend on species identity. We conclude that tree and herb canopy cover as well as herb diversity predominately shape Dipteran communities in temperate deciduous forests, which is in contrast to expectations from grassland studies exhibiting much closer relationships between plant and insect diversity.

Coffee is one of the most important commodities globally and the Coffee Berry Borer (CBB) is its main pest, causing losses of more than half a billion dollars annually. In this systematic review, we quantify the available evidence of successful biological control in coffee agroforestry. There is a recent research trend of switching from the traditional focus on classical biological control to conservation biocontrol, considering a broad range of enemy groups inhabiting the cropping system. We found 19 papers proving CBB biocontrol success in the field and 25 studies suggesting potential biocontrol. Most papers showed effects of fungal infections, followed by papers on ant communities, parasitic Hymenoptera, birds, and nematodes. With respect to local coffee management, arboreal canopy cover providing shade as well as organic practices enhances biocontrol success. Landscape-scale studies are almost missing, although CBB predation by birds can be significant and benefits from the presence of forest patches in the surrounding landscape. Insectivorous birds successfully reduce yield losses by CBB, whereas in many other taxa there is a need for identifying economic impacts of biocontrol. In conclusion, understanding successful coffee management to reduce CBB pest pressure requires more studies on conservation biocontrol, and analyzing the relative importance of local and landscape management for fungal infections as well as for invertebrate and vertebrate predators.

Human welfare is significantly linked to ecosystem services such as the suppression of pest insects by birds and bats. However, effects of biocontrol services on tropical cash crop yield are still largely unknown. For the first time, we manipulated the access of birds and bats in an exclosure experiment (day, night and full exclosures compared to open controls in Indonesian cacao agroforestry) and quantified the arthropod communities, the fruit development and the final yield over a long time period (15 months). We found that bat and bird exclusion increased insect herbivore abundance, despite the concurrent release of mesopredators such as ants and spiders, and negatively affected fruit development, with final crop yield decreasing by 31% across local (shade cover) and landscape (distance to primary forest) gradients. Our results highlight the tremendous economic impact of common insectivorous birds and bats, which need to become an essential part of sustainable landscape management.

Agricultural intensification in terms of decreasing landscape complexity and connectivity has negatively affected biodiversity. Linear landscape elements composed of woody vegetation like hedges may counteract this negative trend by providing habitats and enhancing habitat connectivity for different organisms. Here, we tested the impacts of habitat type (forest edges vs. hedges) and hedges’ isolation (connected vs. isolated hedges) from forests as well as microhabitat conditions (percentage of bare ground and width) on trait-specific occurrence of ground-dwelling arthropods, namely spiders and carabids. Arthropods were grouped by habitat specialisation (forest vs. open-habitat species vs. generalists), hunting strategy (web-building or hunting spiders) and dispersal ability (wing morphology of carabids). Spider and carabid assemblage composition was strongly influenced by habitat type and isolation, but not by microhabitat conditions. Activity density of forest species and brachypterous carabids was higher in forest edges compared to hedges, whereas open-habitat species and macropterous carabids showed reverse patterns, with no effects of isolation. Occurrence of generalist carabids, but not spiders, was higher in hedges compared to forest edges. Habitat type and isolation did not affect spiders with different hunting strategy. Microhabitat conditions were less important for spider and carabid occurrence. Our study concludes that on a landscape scale, type of linear woody habitat is more important for arthropod occurrence than isolation effects and microhabitat conditions, depending on traits. Hedges provide refuges for species specialised to open habitats and species with high dispersal ability, such as macropterous carabids. Forest edges enhance persistence of species specialised to forests and species with low dispersal ability, such as brachypterous carabids.

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.

Biodiversity can provide insurance against environmental change, but only if species differ in their response to environmental conditions (response diversity). Wild bees provide pollination services to wild and crop plants, and response diversity might insure this function against changing climate. To experimentally test the hypothesis that bee species differ in their response to increasing winter temperature, we stored cocoons of nine bee species at different temperatures during the winter (1.5-9.5 °C). Bee species differed significantly in their responses (weight loss, weight at emergence and emergence date). The developmental stage during the winter explained some of these differences. Bee species overwintering as adults generally showed decreased weight and earlier emergence with increasing temperature, whereas bee species overwintering in pre-imaginal stages showed weaker or even opposite responses. This means that winter warming will likely affect some bee species negatively by increasing energy expenditure, while others are less sensitive presumably due to different physiology. Likewise, species phenologies will respond differently to winter warming, potentially affecting plant-pollinator interactions. Responses are not independent of current flight periods: bees active in spring will likely show the strongest phenological advances. Taken together, wild bee diversity provides response diversity to climate change, which may be the basis for an insurance effect.

Increasing yield has emerged as the most prominent element in strategies to deal with growing global demand for food and fibre. It is usually acknowledged that this needs to be done while minimising harm to the environment, but historically land-use intensification has been a major driver of biodiversity loss. The risk is now great that a singular focus on increasing yields will divert attention from the linked problem of biodiversity decline, and the historical pattern will continue. There are options that increase yields while reducing harm to biodiversity, which should be the focus of future strategies. The solutions are not universal, but are locally specific. This is because landscapes vary greatly in inherent biodiversity, the production systems they can support, and the potential for them to be adopted by landholders. While new production techniques might apply at local scale, biodiversity conservation inevitably requires strategies at landscape and larger scales. peerReviewed

Predatory arthropods are attracted to infochemicals emitted by their herbivore prey or by the prey’s host plants. We studied such a tritrophic system measuring the olfactory responses of three potter wasp species (Symmorphusmurarius, Symmorphusgracilis, Discoeliuszonalis, Hymenoptera: Eumeninae) to salicylaldehyde, sequestered as a defence compound by Chrysomela leaf beetle larvae when feeding on Salicaceae, and volatile organic compounds (VOCs) emitted by aspen (Populus tremula, Salicaceae). In electroantennographic recordings (EAG), the highly specialized S. murarius that almost exclusively feeds on larvae of Salicaceae-feeding Chrysomela species was more sensitive to salicylaldehyde than the less specialized S.gracilis, feeding on such Chrysomela species but also weevil larvae. In contrast the related D. zonalis, foraging for microlepidoptera caterpillars on various host plants, did not respond at all. Furthermore, the three wasp species responded differently to aspen VOCs in GC–MS/EAD measurements. These results indicate that the sense of smell of predatory potter wasps differs for prey and plant volatiles among related wasp species according to their degree of host specialization. The considerable differences in salicylaldehyde perception suggest that its originally defensive function has backfired as it is used by specialist potter wasps for prey location. This is an important clue on adaptive mechanisms of the highest trophic level of the well-studied evolutionary arms race among Chrysomela leaf beetles, their host plants and their enemies.

Agricultural intensification (AI) is a great threat to biodiversity and its negative effects on species richness of different communities have been repeatedly shown. The effects of AI on community composition and variability, however, are important for assessing the predictability of community responses, but have rarely been studied simultaneously and across different taxonomic groups. In this study, we focused on parasitoids (primary and secondary) and predators (hoverflies and carabid beetles) of aphids in winter wheat fields with contrasting AI regimes (low AI, i.e. organic fields in structurally complex landscapes vs. high AI, i.e. conventional fields in structurally simple landscapes). We found divergence in species composition of more specialised, low-dispersing primary and secondary parasitoids within high AI fields, probably due to the disruption of the exchanges of species between local populations in structurally simple landscapes. In contrast, species composition of less specialised, highly dispersing carabid beetles and hoverflies converged in fields with high AI, where they were characterised by the dominance of a single, vagile species adapted to high land-use conditions. Furthermore, temporal community shifts were only pronounced in primary parasitoids and hoverflies, with higher temporal changes in fields with high AI in primary parasitoids. Collectively, our results illustrate that environmental homogenisation due to AI does not necessarily induce spatio-temporal homogenisation of communities, but rather can have contrasting effects on more specialised, low-dispersive parasitoids versus more generalised, high-dispersive predators, thereby demonstrating great differences in the predictability of responses to AI among aphid natural enemies.

Current studies on trophic interactions in biodiversity experiments have largely relied on artificially sown gradients in plant diversity, but removal experiments with their more natural plant community composition are more realistic. Slugs are a major part of the invertebrate herbivore community, with some species being common pests in agriculture. We therefore investigated how strongly slugs are influenced by grassland management, plant biodiversity and composition. Here we analysed the effects of cutting frequency, fertilizer application and plant functional group composition on slug densities and their contribution to herbivory on Rumex acetosa in a removal experiment within a >100-year old grassland in Northern Germany. The experiment was laid out as a Latin rectangle with full factorial combinations of (i) plant functional group removal (3 levels) using herbicides, (ii) fertilizer application (2 levels) and (iii) cutting frequency (2 levels). The resulting 12 treatment combinations were replicated 6 times, resulting in 72 plots. We collected a total of 1020 individuals belonging to three species Arion distinctus (60.4% of individuals), Deroceras reticulatum (34.7%) and Arion lusitanicus (4.9%) using a cover board technique and additionally measured herbivore damage to R. acetosa . We found the highest slug abundance on plots with a low cutting frequency and high food resource availability (increased cover of forbs and taller vegetation). Fertilizer application had no significant effect on slug abundance, but caused higher herbivore damage to on R. acetosa , possibly as a result of increased tissue quality. The negative effect of higher cutting frequency on slug abundance was lowest in control plots with their naturally developed graminoid-forb communities (cutting reduced slug density by 6% in the control vs. 29% in herbicide plots). Our experiments therefore support the idea that more natural plant species compositions reduce the impact of disturbances (e.g. through cutting or grazing) on invertebrates.

Tropical small-holder farmers rely on sustainable food production. Crop seed predation by ants can cause substantial yield loss and result in high pesticide use. We conducted field experiments and questionnaire-based surveys aiming to assess the effect of sown-seed predation on four crop species (Cucumis sativus, Daucus carota, Capsicum frutescens and Solanum melongena) in 15 vegetable gardens and the resulting impact on the net income of Indonesian farmers. Furthermore, we tested a commonly applied insecticide and herbicide for seed, seedling and plant protection aiming to understand their effect on ant seed predation. We found that the mean percentage of seeds removed per garden was 42.0%, 49.4%, 48.0% and 50.6% for C. sativus, D. carota, C. frutescens and S. melongena, respectively, halving the farmers' income after considering initial and operational costs. Insecticide and herbicide treatments did not affect seed predation success or overall ant abundance, although they had positive and negative effects on ant species-specific abundance. High overall ant abundance caused high seed predation rate in all gardens as a result of a functional redundancy of ant species, which compensated for pesticide-related species loss. Environmentally-friendly and more sustainable practices such as overseeding or seedling production in nurseries could substitute for these inefficient approaches of chemical pest control, although this requires further research.

Grasses are a dominant component of meadows and pastures, harboring an often overlooked diversity of non-pest stem-boring insects that feed and develop exclusively enclosed by plant tissue inside grass shoots. Surprisingly, the effects of land-use management on these highly specialized communities have rarely been studied. Here, the applicability of short-term management reduction as a conservational tool, increasing stem-borer colonization success, was examined. On 41 grasslands in Germany a temporal gradient of set-aside treatments was established by experimentally excluding subplots from management, or by a priori selection of already abandoned grassland patches. Stem-borer abundances and attack heights on resulting managed, 1-season unmanaged and ≥2-seasons abandoned treatments were compared. Grassland management had a negative effect on stem-borer colonization success and spatial niche differentiation. Reducing management over a 1-season period did not enhance abundances. Two out of three species responded positively only to an abandoning treatment lasting ≥2-seasons, which was also reflected by the decreasing spatial overlap in this treatment. Even though grass shoots in unmanaged areas were on average 14.9 cm taller than in managed areas, stem-borer abundances did not differ between the latter treatments. Hence, preserving essential larval development and hibernation habitats in future rotational set-aside schemes requires a management exclusion period of at least two growing seasons. From these grassy strips, e.g. located at meadow edges, highly specialized stem-borers can re-colonize sward islets on surrounding intensively managed grasslands, despite of ever changing land-use practices and environmental conditions.

Summary Calcareous grasslands are among the most species-rich habitats in Europe, but are increasingly threatened due to abandonment and fragmentation. Little is known about how the surrounding landscape influences fragmentation effects. Here, we focus on the interaction between habitat fragmentation and landscape composition on leafhoppers, a highly diverse group of insects, including many species that are likely to be vulnerable to changes in their environment. We selected 14 small and 14 large fragments of calcareous grassland in Central Germany, differing in isolation from other calcareous grasslands and in composition of the surrounding landscape. Leafhoppers, sampled by sweep netting, were either specialists that depended on calcareous grasslands or generalists that could use the landscape matrix, but still required low-productivity habitats. Increasing habitat isolation reduced leafhopper species richness in simple (dominated by arable crops), but not in complex landscapes. This effect was driven by the generalist species. In simple landscapes, leafhoppers may find it more difficult to reach the next suitable fragment due to a lack of alternative resources during dispersal. Moreover, we found that generalist species richness increased with increasing connectivity on small fragments, whereas it remained stable with increasing connectivity on large fragments. In small, isolated fragments, a higher extinction rate combined with a lower probability of recolonization is thought to cause the reduced species richness. Synthesis and applications. Our results show for the first time that insect species richness can be negatively affected by increasing habitat isolation in simplified but not in complex landscapes and in small but not in large fragments. We provide evidence that mitigating the negative effects of habitat fragmentation needs to take the surrounding landscape into account. Management efforts should prioritize (i) an increase in connectivity of small, isolated fragments, (ii) an increase in connectivity of fragments in simple landscapes and (iii) enhanced dispersal by increasing heterogeneity of both landscape composition and configuration. Moreover, extensive management of fragments by grazing or mowing to increase local habitat quality for leafhoppers would benefit other insect groups as well.

Background: Plant diversity can influence numerous ecosystem processes, including plant–animal interactions, which, in turn, will affect plant growth and fitness. At present, little is known on how plant–animal interactions in forests respond to gradients in tree and herb-layer diversity. Aims: To quantify how invertebrate herbivory, pollination-dependent seed production and post-dispersal seed predation vary along a gradient of tree and herb diversity in a semi-natural temperate deciduous forest. Methods: Potted individuals of the understorey herbs Lathyrus vernus and Primula elatior were exposed in 40 forest plots along a natural gradient of tree and herb diversity for 3 months to record seed production and leaf damage caused by invertebrate herbivores. In half the plants, pollinators were experimentally excluded to test if seed production depended on insect pollination. In Lathyrus vernus linkages between below- and above-ground herbivory were tested by inoculating plants with root-feeding nematodes. To study seed predation, we measured seed removal from seed depots that were selectively accessible to different seed predator groups. Results: Herbivore damage decreased with increasing tree diversity in P. elatior. In L. vernus, above-ground herbivory was higher in nematode-treated plants than in control plants. Seed production in L. vernus, which strongly depended on insect pollination, showed a positive relationship with tree diversity. Seed predation was positively related to herb diversity in L. vernus, but only weakly so in P. elatior. While both vertebrates and invertebrates acted as seed predators of L. vernus, seeds of P. elatior were mainly predated by invertebrates. Conclusions: The fitness of understorey plants is linked to tree and herb diversity via changes in plant-invertebrate interactions. However, species-specific responses of study plants underline the importance of species identity effects in addition to effects of biodiversity per se.

Hedges and forest edges play a major role in providing nesting sites, food resources and shelter for birds in agricultural landscapes of western and central Europe. We investigated the response of farmland vs. woodland birds at two degrees of isolation of hedges from forest and to vegetation structure. We surveyed 200 m long sections of six forest edges, six hedges connected to forests and six isolated hedges. Species richness and abundance of farmland birds were higher in hedges than in forest edges, species richness and abundance of woodland birds were lower in hedges than in the forest edges. Species richness and abundance of both groups did not differ between connected and isolated hedges. Width and height of hedges and edges did not affect the species richness and abundance of either farmland or woodland birds. Furthermore, bird community composition differed between habitat types (hedge vs. forest edge) and also between hedge isolation levels (hedges connected to forest vs. isolated hedges). Based o...

Insect-feeding birds may interfere with trophic interactions in plant-insect food webs, which may be particularly important in agroecosystems. Here, we studied how Eurasian Tree Sparrows (Passer montanus) affect aphids and their predators in cereal fields using bird exclusion experiments. The Tree Sparrows fed their nestlings with aphid antagonists. Hoverflies and ladybird beetles accounted for 77% of the food for the nestlings during peak aphid density. When birds were excluded, densities of hoverfly larvae, which were the most abundant aphid predator group in the cereals, were 4% higher in wheat and 45% higher in oat, while aphid densities were 24% lower in wheat and 26% lower in oat. The demonstrated disruption of biological control by mesopredators through bird predation may be a common phenomenon in cropping systems characterized by small-sized and abundant pest species. Management of biotic interactions such as biological control needs a broad food-web perspective, even in simplified agroecosystems.

Agricultural intensification drives biodiversity loss and shapes farmers’ profit, but the role of legacy effects and detailed quantification of ecological–economic trade-offs are largely unknown. In Europe during the 1950s, the Eastern communist bloc switched to large-scale farming by forced collectivization of small farms, while the West kept small-scale private farming. Here we show that large-scale agriculture in East Germany reduced biodiversity, which has been maintained in West Germany due to >70% longer field edges than those in the East. In contrast, profit per farmland area in the East was 50% higher than that in the West, despite similar yield levels. In both regions, switching from conventional to organic farming increased biodiversity and halved yield levels, but doubled farmers’ profits. In conclusion, European Union policy should acknowledge the surprisingly high biodiversity benefits of small-scale agriculture, which are on a par with conversion to organic agriculture. East German collective farms reduced biodiversity but raised profits, in contrast to smaller private farms in West Germany, with legacy effects that continue today.

Local biodiversity can be expected to be similar worldwide if environmental conditions are similar. Here, we hypothesize that tropical ant communities with different types of regional species pools but at similar habitat types in Brazil and Indonesia show similar diversity patterns at multiple spatial scales, when comparing (1) the relative contribution of alpha and beta diversity to gamma diversity; (2) the number of distinct communities (community differentiation); and (3) the drivers of β-diversity (species replacement or species loss/gain) at each spatial scale. In both countries, rainforests and savannas (biome scale) were represented by three landscapes (landscape scale), each with four transects (site scale) and each transect with 10 pitfall traps (local scale). At the local scale, α-diversity was higher and β-diversity lower than expected from null models. Hence, we observed a high coexistence of species across biomes. The replacement of species seemed the most important factor for β-diversity among sites and among landscapes across biomes. Species sorting, landscape-moderated species distribution and neutral drift are potential mechanisms for the high β-diversity among sites within landscapes. At the biome scale, different evolutionary histories produced great differences in ant community composition, so the replacement of species is, at this scale, the most important driver of beta diversity. According to these key findings, we conclude that distinct regional ant species pools from similar tropical habitat types are similarly constrained across several spatial scales, regardless of the continent considered.

Land-use intensification is a major driver of biodiversity loss(1,2). Alongside reductions in local species diversity, biotic homogenization at larger spatial scales is of great concern for conservation. Biotic homogenization means a decrease in beta-diversity (the compositional dissimilarity between sites). Most studies have investigated losses in local (alpha)-diversity(1,3) and neglected biodiversity loss at larger spatial scales. Studies addressing beta-diversity have focused on single or a few organism groups (for example, ref. 4), and it is thus unknown whether land-use intensification homogenizes communities at different trophic levels, above-and belowground. Here we show that even moderate increases in local land-use intensity (LUI) cause biotic homogenization across microbial, plant and animal groups, both above- and belowground, and that this is largely independent of changes in alpha-diversity. We analysed a unique grassland biodiversity dataset, with abundances of more than 4,000 species belonging to 12 trophic groups. LUI, and, in particular, high mowing intensity, had consistent effects on beta-diversity across groups, causing a homogenization of soil microbial, fungal pathogen, plant and arthropod communities. These effects were nonlinear and the strongest declines in beta-diversity occurred in the transition from extensively managed to intermediate intensity grassland. LUI tended to reduce local alpha-diversity in aboveground groups, whereas the alpha-diversity increased in belowground groups. Correlations between the alpha-diversity of different groups, particularly between plants and their consumers, became weaker at high LUI. This suggests a loss of specialist species and is further evidence for biotic homogenization. The consistently negative effects of LUI on landscape-scale biodiversity underscore the high value of extensively managed grasslands for conserving multitrophic biodiversity and ecosystem service provision. Indeed, biotic homogenization rather than local diversity loss could prove to be the most substantial consequence of land-use intensification.

Ongoing biodiversity decline impairs ecosystem processes, including pollination. Flower visitation, an important indicator of pollination services, is influenced by plant species richness. However, the spatio-temporal responses of different pollinator groups to plant species richness have not yet been analyzed experimentally. Here, we used an experimental plant species richness gradient to analyze plant-pollinator interactions with an unprecedented spatio-temporal resolution. We observed four pollinator functional groups (honeybees, bumblebees, solitary bees, and hoverflies) in experimental plots at three different vegetation strata between sunrise and sunset. Visits were modified by plant species richness interacting with time and space. Furthermore, the complementarity of pollinator functional groups in space and time was stronger in species-rich mixtures. We conclude that high plant diversity should ensure stable pollination services, mediated via spatio-temporal niche complementarity in flower visitation. Declining plant diversity has been shown to affect ecosystem processes such as plant-pollinator interactions, but it is currently not known if or how spatio-temporal niche partitioning of pollinators is affected by plant biodiversity. In a grassland biodiversity experiment, where we manipulated plant species richness and studied spatio-temporal resource use of pollinators, we found that complementarity of pollinator groups in space and time was stronger in species-rich mixtures. Our study shows that high plant diversity may ensure stable pollination services due to increased pollinator complementarity.

There is compelling evidence that more diverse ecosystems deliver greater benefits to people, and these ecosystem services have become a key argument for biodiversity conservation. However, it is unclear how much biodiversity is needed to deliver ecosystem services in a cost-effective way. Here we show that, while the contribution of wild bees to crop production is significant, service delivery is restricted to a limited subset of all known bee species. Across crops, years and biogeographical regions, crop-visiting wild bee communities are dominated by a small number of common species, and threatened species are rarely observed on crops. Dominant crop pollinators persist under agricultural expansion and many are easily enhanced by simple conservation measures, suggesting that cost-effective management strategies to promote crop pollination should target a different set of species than management strategies to promote threatened bees. Conserving the biological diversity of bees therefore requires more than just ecosystem-service-based arguments.

Cultural ecosystem services are growing in importance and their substantial contribution to well-being is well recognized. Yet, significant conceptual and methodological gaps exist, especially for urban ecosystems. We analyzed perceptions of cultural ecosystem services in the urban context of Berlin, based on qualitative research methods. Using expert and problem-centered interviews, we show how cultural ecosystem services are understood and which focus areas were emphasized. We compared our inductive codes with the Millennium Ecosystem Assessment. While our findings show supporting evidence for the Millennium Ecosystem Assessment classification, some categories had to be substituted and others adjusted to local citizen understandings. The results reveal a variety of intricate cultural ecosystem service perceptions. Hence, selecting and emphasizing only a few services without prior studies could misinform decision-makers and lead to biased policy outcome. Regionally specific perceptions of cultural benefits from urban green are important information for planning processes. Cultural ecosystem services could be one way to achieve awareness of socio-ecological aspects, as our results show linkages between cultural ecosystem services and urban social sustainability. Using qualitative cultural ecosystem service research could foster public participation and increase the input of regionally specific perceptions.

Tropical lowland rainforests are increasingly threatened by the expansion of agriculture and the extraction of natural resources. In Jambi Province, Indonesia, the interdisciplinary EFForTS project focuses on the ecological and socio-economic dimensions of rainforest conversion to jungle rubber agroforests and monoculture plantations of rubber and oil palm. Our data confirm that rainforest transformation and land use intensification lead to substantial losses in biodiversity and related ecosystem functions, such as decreased above- and below-ground carbon stocks. Owing to rapid step-wise transformation from forests to agroforests to monoculture plantations and renewal of each plantation type every few decades, the converted land use systems are continuously dynamic, thus hampering the adaptation of animal and plant communities. On the other hand, agricultural rainforest transformation systems provide increased income and access to education, especially for migrant smallholders. Jungle rubber and rubber monocultures are associated with higher financial land productivity but lower financial labour productivity compared to oil palm, which influences crop choice: smallholders that are labour-scarce would prefer oil palm while land-scarce smallholders would prefer rubber. Collecting long-term data in an interdisciplinary context enables us to provide decision-makers and stakeholders with scientific insights to facilitate the reconciliation between economic interests and ecological sustainability in tropical agricultural landscapes.

Ecologists and farmers often have contrasting perceptions about the value of natural habitat in agricultural production landscapes, which so far has been little acknowledged in ecology and conservation. Ecologists and conservationists often appreciate the contribution of natural habitat to biodiversity and potential ecosystem services such as biological pest control, whereas many farmers see habitat remnants as a waste of cropland or source of pests. While natural habitat has been shown to increase pest control in many systems, we here identify five hypotheses for when and why natural habitat can fail to support biological pest control, and illustrate each with case studies from the literature: (1) pest populations have no effective natural enemies in the region, (2) natural habitat is a greater source of pests than natural enemies, (3) crops provide more resources for natural enemies than does natural habitat, (4) natural habitat is insufficient in amount, proximity, composition, or configuration to provide large enough enemy populations needed for pest control, and (5) agricultural practices counteract enemy establishment and biocontrol provided by natural habitat. In conclusion, we show that the relative importance of natural habitat for biocontrol can vary dramatically depending on type of crop, pest, predator, land management, and landscape structure. This variation needs to be considered when designing measures aimed at enhancing biocontrol services through restoring or maintaining natural habitat.