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The expansion of the oil palm industry in Indonesia has improved livelihoods in rural communities, but comes at the cost of biodiversity and ecosystem degradation. Here, we investigated ways to balance ecological and economic outcomes of oil palm cultivation. We compared a wide range of production systems, including smallholder plantations, industrialized company estates, estates with improved agronomic management, and estates with native tree enrichment. Across all management types, we assessed multiple indicators of biodiversity, ecosystem functions, management, and landscape structure to identify factors that facilitate economic-ecological win-wins, using palm yields as measure of economic performance. Although, we found that yields in industrialized estates were, on average, twice as high as those in smallholder plantations, ecological indicators displayed substantial variability across systems, regardless of yield variations, highlighting potential for economic-ecological win-wins. Reducing management intensity (e.g., mechanical weeding instead of herbicide application) did not lower yields but improved ecological outcomes at moderate costs, making it a potential measure for balancing economic and ecological demands. Additionally, maintaining forest cover in the landscape generally enhanced local biodiversity and ecosystem functioning within plantations. Enriching plantations with native trees is also a promising strategy to increase ecological value without reducing productivity. Overall, we recommend closing yield gaps in smallholder cultivation through careful intensification, whereas conventional plantations could reduce management intensity without sacrificing yield. Our study highlights various pathways to reconcile the economics and ecology of palm oil production and identifies management practices for a more sustainable future of oil palm cultivation., Competing Interests: Competing interests statement:The authors declare no competing interest.

Terrestrial animal biodiversity is increasingly being lost because of land-use change 1,2 . However, functional and energetic consequences aboveground and belowground and across trophic levels in megadiverse tropical ecosystems remain largely unknown. To fill this gap, we assessed changes in energy fluxes across 'green' aboveground (canopy arthropods and birds) and 'brown' belowground (soil arthropods and earthworms) animal food webs in tropical rainforests and plantations in Sumatra, Indonesia. Our results showed that most of the energy in rainforests is channelled to the belowground animal food web. Oil palm and rubber plantations had similar or, in the case of rubber agroforest, higher total animal energy fluxes compared to rainforest but the key energetic nodes were distinctly different: in rainforest more than 90% of the total animal energy flux was channelled by arthropods in soil and canopy, whereas in plantations more than 50% of the energy was allocated to annelids (earthworms). Land-use change led to a consistent decline in multitrophic energy flux aboveground, whereas belowground food webs responded with reduced energy flux to higher trophic levels, down to -90%, and with shifts from slow (fungal) to fast (bacterial) energy channels and from faeces production towards consumption of soil organic matter. This coincides with previously reported soil carbon stock depletion 3 . Here we show that well-documented animal biodiversity declines with tropical land-use change 4-6 are associated with vast energetic and functional restructuring in food webs across aboveground and belowground ecosystem compartments., (© 2024. The Author(s).)

In agricultural landscapes, semi-natural habitats can support populations of beneficial species, such as natural enemies. Using artificial caterpillars made of plasticine, we compared arthropod and vertebrate predation rates in wheat fields adjacent to hedgerows or spontaneous grass margins in two contrasting agricultural landscapes. Overall, 25.3% of the caterpillars were attacked after 24 h, mainly by arthropods (67.1%). Predation rates in the heterogeneous and homogeneous landscapes were similar (mean ± SE; 24.9% ± 3.0% vs 25.7% ± 3.6%, n = 10 per landscape). However, we found a significant interaction between the margin and landscape type. Total (arthropods + vertebrates) predation rates were significantly higher in wheat fields next to hedgerows than in conventional wheat fields with grass margins in the homogeneous landscape (33.7% ± 4.5% vs 17.7% ± 2.5%, n = 5 per margin type and landscape), while no difference between predation rates in the two margin types was detected in the heterogeneous landscape. Total and vertebrate predation rates were positively affected by plant species richness, which suggest that one of the mechanisms through which hedgerows can support higher predator activity than spontaneous grass margins, particularly in homogeneous agricultural landscapes, is by providing higher plant diversity. Our results suggest that in simple landscapes, hedgerows may act as habitat islands of high conservation value for biodiversity, having a disproportionate value compared to hedgerows in landscapes that include forest fragments and other semi-natural habitats., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

Urbanization is a major driver of biodiversity change but how it interacts with spatial and temporal gradients to influence the dynamics of plant-pollinator networks is poorly understood, especially in tropical urbanization hotspots. Here, we analysed the drivers of environmental, spatial and temporal turnover of plant-pollinator interactions (interaction β-diversity) along an urbanization gradient in Bengaluru, a South Indian megacity. The compositional turnover of plant-pollinator interactions differed more between seasons and with local urbanization intensity than with spatial distance, suggesting that seasonality and environmental filtering were more important than dispersal limitation for explaining plant-pollinator interaction β-diversity. Furthermore, urbanization amplified the seasonal dynamics of plant-pollinator interactions, with stronger temporal turnover in urban compared to rural sites, driven by greater turnover of native non-crop plant species (not managed by people). Our study demonstrates that environmental, spatial and temporal gradients interact to shape the dynamics of plant-pollinator networks and urbanization can strongly amplify these dynamics., (© 2023 The Authors. Ecology Letters published by John Wiley & Sons Ltd.)

The impact of local biodiversity loss on ecosystem functioning is well established, but the role of larger-scale biodiversity dynamics in the delivery of ecosystem services remains poorly understood. Here we address this gap using a comprehensive dataset describing the supply of 16 cultural, regulating and provisioning ecosystem services in 150 European agricultural grassland plots, and detailed multi-scale data on land use and plant diversity. After controlling for land-use and abiotic factors, we show that both plot-level and surrounding plant diversity play an important role in the supply of cultural and aboveground regulating ecosystem services. In contrast, provisioning and belowground regulating ecosystem services are more strongly driven by field-level management and abiotic factors. Structural equation models revealed that surrounding plant diversity promotes ecosystem services both directly, probably by fostering the spill-over of ecosystem service providers from surrounding areas, and indirectly, by maintaining plot-level diversity. By influencing the ecosystem services that local stakeholders prioritized, biodiversity at different scales was also shown to positively influence a wide range of stakeholder groups. These results provide a comprehensive picture of which ecosystem services rely most strongly on biodiversity, and the respective scales of biodiversity that drive these services. This key information is required for the upscaling of biodiversity-ecosystem service relationships, and the informed management of biodiversity within agricultural landscapes., (© 2022. The Author(s), under exclusive licence to Springer Nature Limited.)

Urbanization poses a major threat to biodiversity and food security, as expanding cities, especially in the Global South, increasingly compete with natural and agricultural lands. However, the impact of urban expansion on agricultural biodiversity in tropical regions is overlooked. Here we assess how urbanization affects the functional response of farmland bees, the most important pollinators for crop production. We sampled bees across three seasons in 36 conventional vegetable-producing farms spread along an urbanization gradient in Bengaluru, an Indian megacity. We investigated how landscape and local environmental drivers affected different functional traits (sociality, nesting behavior, body size, and specialization) and functional diversity (functional dispersion) of bee communities. We found that the functional responses to urbanization were trait specific with more positive than negative effects of gray area (sealed surfaces and buildings) on species richness, functional diversity, and abundance of most functional groups. As expected, larger, solitary, cavity-nesting, and, surprisingly, specialist bees benefited from urbanization. In contrast to temperate cities, the abundance of ground nesters increased in urban areas, presumably because larger patches of bare soil were still available beside roads and buildings. However, overall bee abundance and the abundance of social bees (85% of all bees) decreased with urbanization, threatening crop pollination. Crop diversity promotes taxonomic and functional diversity of bee communities. Locally, flower resources promote the abundance of all functional groups, and natural vegetation can maintain diverse pollinator communities throughout the year, especially during the noncropping season. However, exotic plants decrease functional diversity and bee specialization. To safeguard bees and their pollination services in urban farms, we recommend (1) preserving seminatural vegetation (hedges) around cropping fields to provide nesting opportunities for aboveground nesters, (2) promoting farm-level crop diversification of beneficial crops (e.g., pulses, vegetables, and spices), (3) maintaining native natural vegetation along field margins, and (4) controlling and removing invasive exotic plants that disrupt native plant-pollinator interactions. Overall, our results suggest that urban agriculture can maintain functionally diverse bee communities and, if managed in a sustainable manner, be used to develop win-win solutions for biodiversity conservation of pollinators and food security in and around cities., (© 2022 The Authors. Ecological Applications published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Background: Intense conversion of tropical forests into agricultural systems contributes to habitat loss and the decline of ecosystem functions. Plant-pollinator interactions buffer the process of forest fragmentation, ensuring gene flow across isolated patches of forests by pollen transfer. In this study, we identified the composition of pollen grains stored in pot-pollen of stingless bees, Tetragonula laeviceps, via dual-locus DNA metabarcoding (ITS2 and rbcL) and light microscopy, and compared the taxonomic coverage of pollen sampled in distinct land-use systems categorized in four levels of management intensity (forest, shrub, rubber, and oil palm) for landscape characterization., Results: Plant composition differed significantly between DNA metabarcoding and light microscopy. The overlap in the plant families identified via light microscopy and DNA metabarcoding techniques was low and ranged from 22.6 to 27.8%. Taxonomic assignments showed a dominance of pollen from bee-pollinated plants, including oil-bearing crops such as the introduced species Elaeis guineensis (Arecaceae) as one of the predominant taxa in the pollen samples across all four land-use types. Native plant families Moraceae, Euphorbiaceae, and Cannabaceae appeared in high proportion in the analyzed pollen material. One-way ANOVA (p > 0.05), PERMANOVA (R² values range from 0.14003 to 0.17684, for all tests p-value > 0.5), and NMDS (stress values ranging from 0.1515 to 0.1859) indicated a lack of differentiation between the species composition and diversity of pollen type in the four distinct land-use types, supporting the influx of pollen from adjacent areas., Conclusions: Stingless bees collected pollen from a variety of agricultural crops, weeds, and wild plants. Plant composition detected at the family level from the pollen samples likely reflects the plant composition at the landscape level rather than the plot level. In our study, the plant diversity in pollen from colonies installed in land-use systems with distinct levels of forest transformation was highly homogeneous, reflecting a large influx of pollen transported by stingless bees through distinct land-use types. Dual-locus approach applied in metabarcoding studies and visual pollen identification showed great differences in the detection of the plant community, therefore a combination of both methods is recommended for performing biodiversity assessments via pollen identification., (© 2022. The Author(s).)

We challenge the widespread appraisal that organic farming is the fundamental alternative to conventional farming for harnessing biodiversity in agricultural landscapes. Certification of organic production is largely restricted to banning synthetic agrochemicals, resulting in limited benefits for biodiversity but high yield losses despite ongoing intensification and specialisation. In contrast, successful agricultural measures to enhance biodiversity include diversifying cropland and reducing field size, which can multiply biodiversity while sustaining high yields in both conventional and organic systems. Achieving a landscape-level mosaic of natural habitat patches and fine-grained cropland diversification in both conventional and organic agriculture is key for promoting large-scale biodiversity. This needs to be urgently acknowledged by policy makers for an agricultural paradigm shift., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Urbanization is a major driver of land use change and biodiversity decline. While most of the ongoing and future urbanization hotspots are located in the Global South, the impact of urban expansion on agricultural biodiversity and associated functions and services in these regions has widely been neglected. Additionally, most studies assess biodiversity responses at local scale (α-diversity), however, ecosystem functioning is strongly determined by compositional and functional turnover of communities (β-diversity) at regional scales. We investigated taxonomic and functional β-diversity of farmland birds across three seasons on 36 vegetable farms spread along a continuous urbanization gradient in Bangalore, a South Indian megacity. Increasing amount of grey area in the farm surroundings was the dominant driver affecting β-diversity and resulting in taxonomic and functional homogenization of farmland bird communities. Functional diversity losses were higher than expected from species declines (i.e., urbanization acts as an environmental filter), with particular losses of functionally important groups such as insectivores of crop pests. Moreover, urbanization reduced functional redundancy of bird communities, which may further weaken ecosystems resilience to future perturbations. Our study underscores urbanization as a major driver of taxonomic and functional homogenization of species communities in agricultural systems, potentially threatening crucial ecosystem services for food production., (© 2021 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Land-use intensification is a major driver of biodiversity loss. However, understanding how different components of land use drive biodiversity loss requires the investigation of multiple trophic levels across spatial scales. Using data from 150 agricultural grasslands in central Europe, we assess the influence of multiple components of local- and landscape-level land use on more than 4,000 above- and belowground taxa, spanning 20 trophic groups. Plot-level land-use intensity is strongly and negatively associated with aboveground trophic groups, but positively or not associated with belowground trophic groups. Meanwhile, both above- and belowground trophic groups respond to landscape-level land use, but to different drivers: aboveground diversity of grasslands is promoted by diverse surrounding land-cover, while belowground diversity is positively related to a high permanent forest cover in the surrounding landscape. These results highlight a role of landscape-level land use in shaping belowground communities, and suggest that revised agroecosystem management strategies are needed to conserve whole-ecosystem biodiversity.

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.

Worldwide pollinator declines lead to pollination deficits in crops and wild plants, and managed bees are frequently used to meet the increasing demand for pollination. However, their foraging can be affected by flower availability and colony size. We investigated how mass-flowering oilseed rape (OSR) can influence the pollen resource use of small and large honey bee (Apis mellifera L.) and bumble bee (Bombus terrestris L.) colonies. Colonies were placed adjacent to strawberry fields along a gradient of OSR availability in the landscapes. We used ITS2 metabarcoding to identify the pollen richness based on ITS2 amplicon sequencing and microscopy for quantification of target pollen. Bumble bees collected pollen from more different plant genera than honey bees. In both species, strawberry pollen collection decreased with high OSR availability but was facilitated by increasing strawberry flower cover. Colony size had no effect. The relationship between next-generation sequencing-generated ITS2 amplicon reads and microscopic pollen counts was positive but pollen type-specific. Bumble bees and, to a lesser degree, honey bees collected pollen from a wide variety of plants. Therefore, in order to support pollinators and associated pollination services, future conservation schemes should sustain and promote pollen plant richness in agricultural landscapes. Both bee species responded to the availability of flower resources in the landscape. Although honey bees collected slightly more strawberry pollen than bumble bees, both can be considered as crop pollinators. Metabarcoding could provide similar quantitative information to microscopy, taking into account the pollen types, but there remains high potential to improve the methodological weaknesses., (© 2020 The Authors. Molecular Ecology published by John Wiley & Sons Ltd.)

Bumble bees are important crop pollinators and provide important pollination services to their respective ecosystems. Their pollen diet and thus food preferences can be characterized through nucleic acid sequence analysis. We present ITS2 amplicon sequence data from pollen collected by bumble bees. The pollen was collected from six different bumble bee colonies that were placed in independent agricultural landscapes. We compared next-generation (Illumina), i.e., short-read, and third-generation (Nanopore), i.e., MinION, sequencing techniques. MinION data were preprocessed using traditional and Nanopore specific tools for comparative analysis and were evaluated in comparison to short-read sequence data with conventional processing. Based on the results, the dietary diary of bumble bee in the studied landscapes can be identified. It is known that short reads generated by next-generation sequencers have the advantage of higher quality scores while Nanopore yields longer read lengths. We show that assignments to taxonomic units yield comparable results when querying against an ITS2-specific sequence database. Thus, lower sequence quality is compensated by longer read lengths. However, the Nanopore technology is improving in terms of data quality, much cheaper, and suitable for portable applications. With respect to the studied agricultural landscapes we found that bumble bees require higher plant diversity than only crops to fulfill their foraging requirements., (Copyright © 2020 Leidenfrost, Bänsch, Prudnikow, Brenig, Westphal and Wünschiers.)

Human land use threatens global biodiversity and compromises multiple ecosystem functions critical to food production. Whether crop yield-related ecosystem services can be maintained by a few dominant species or rely on high richness remains unclear. Using a global database from 89 studies (with 1475 locations), we partition the relative importance of species richness, abundance, and dominance for pollination; biological pest control; and final yields in the context of ongoing land-use change. Pollinator and enemy richness directly supported ecosystem services in addition to and independent of abundance and dominance. Up to 50% of the negative effects of landscape simplification on ecosystem services was due to richness losses of service-providing organisms, with negative consequences for crop yields. Maintaining the biodiversity of ecosystem service providers is therefore vital to sustain the flow of key agroecosystem benefits to society., (Copyright © 2019 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works. Distributed under a Creative Commons Attribution NonCommercial License 4.0 (CC BY-NC).)

The PREDICTS project-Projecting Responses of Ecological Diversity In Changing Terrestrial Systems (www.predicts.org.uk)-has collated from published studies a large, reasonably representative database of comparable samples of biodiversity from multiple sites that differ in the nature or intensity of human impacts relating to land use. We have used this evidence base to develop global and regional statistical models of how local biodiversity responds to these measures. We describe and make freely available this 2016 release of the database, containing more than 3.2 million records sampled at over 26,000 locations and representing over 47,000 species. We outline how the database can help in answering a range of questions in ecology and conservation biology. To our knowledge, this is the largest and most geographically and taxonomically representative database of spatial comparisons of biodiversity that has been collated to date; it will be useful to researchers and international efforts wishing to model and understand the global status of biodiversity.

Land-use intensification is a major driver of biodiversity loss. Alongside reductions in local species diversity, biotic homogenization at larger spatial scales is of great concern for conservation. Biotic homogenization means a decrease in β-diversity (the compositional dissimilarity between sites). Most studies have investigated losses in local (α)-diversity and neglected biodiversity loss at larger spatial scales. Studies addressing β-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 α-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 β-diversity across groups, causing a homogenization of soil microbial, fungal pathogen, plant and arthropod communities. These effects were nonlinear and the strongest declines in β-diversity occurred in the transition from extensively managed to intermediate intensity grassland. LUI tended to reduce local α-diversity in aboveground groups, whereas the α-diversity increased in belowground groups. Correlations between the β-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.

Many experiments have shown that loss of biodiversity reduces the capacity of ecosystems to provide the multiple services on which humans depend. However, experiments necessarily simplify the complexity of natural ecosystems and will normally control for other important drivers of ecosystem functioning, such as the environment or land use. In addition, existing studies typically focus on the diversity of single trophic groups, neglecting the fact that biodiversity loss occurs across many taxa and that the functional effects of any trophic group may depend on the abundance and diversity of others. Here we report analysis of the relationships between the species richness and abundance of nine trophic groups, including 4,600 above- and below-ground taxa, and 14 ecosystem services and functions and with their simultaneous provision (or multifunctionality) in 150 grasslands. We show that high species richness in multiple trophic groups (multitrophic richness) had stronger positive effects on ecosystem services than richness in any individual trophic group; this includes plant species richness, the most widely used measure of biodiversity. On average, three trophic groups influenced each ecosystem service, with each trophic group influencing at least one service. Multitrophic richness was particularly beneficial for 'regulating' and 'cultural' services, and for multifunctionality, whereas a change in the total abundance of species or biomass in multiple trophic groups (the multitrophic abundance) positively affected supporting services. Multitrophic richness and abundance drove ecosystem functioning as strongly as abiotic conditions and land-use intensity, extending previous experimental results to real-world ecosystems. Primary producers, herbivorous insects and microbial decomposers seem to be particularly important drivers of ecosystem functioning, as shown by the strong and frequent positive associations of their richness or abundance with multiple ecosystem services. Our results show that multitrophic richness and abundance support ecosystem functioning, and demonstrate that a focus on single groups has led to researchers to greatly underestimate the functional importance of biodiversity.

Land-use change and intensification threaten bee populations worldwide, imperilling pollination services. Global models are needed to better characterise, project, and mitigate bees' responses to these human impacts. The available data are, however, geographically and taxonomically unrepresentative; most data are from North America and Western Europe, overrepresenting bumblebees and raising concerns that model results may not be generalizable to other regions and taxa. To assess whether the geographic and taxonomic biases of data could undermine effectiveness of models for conservation policy, we have collated from the published literature a global dataset of bee diversity at sites facing land-use change and intensification, and assess whether bee responses to these pressures vary across 11 regions (Western, Northern, Eastern and Southern Europe; North, Central and South America; Australia and New Zealand; South East Asia; Middle and Southern Africa) and between bumblebees and other bees. Our analyses highlight strong regionally-based responses of total abundance, species richness and Simpson's diversity to land use, caused by variation in the sensitivity of species and potentially in the nature of threats. These results suggest that global extrapolation of models based on geographically and taxonomically restricted data may underestimate the true uncertainty, increasing the risk of ecological surprises.

Species diversity promotes the delivery of multiple ecosystem functions (multifunctionality). However, the relative functional importance of rare and common species in driving the biodiversity-multifunctionality relationship remains unknown. We studied the relationship between the diversity of rare and common species (according to their local abundances and across nine different trophic groups), and multifunctionality indices derived from 14 ecosystem functions on 150 grasslands across a land-use intensity (LUI) gradient. The diversity of above- and below-ground rare species had opposite effects, with rare above-ground species being associated with high levels of multifunctionality, probably because their effects on different functions did not trade off against each other. Conversely, common species were only related to average, not high, levels of multifunctionality, and their functional effects declined with LUI. Apart from the community-level effects of diversity, we found significant positive associations between the abundance of individual species and multifunctionality in 6% of the species tested. Species-specific functional effects were best predicted by their response to LUI: species that declined in abundance with land use intensification were those associated with higher levels of multifunctionality. Our results highlight the importance of rare species for ecosystem multifunctionality and help guiding future conservation priorities., (© 2016 The Author(s).)

Biodiversity loss can affect the viability of ecosystems by decreasing the ability of communities to respond to environmental change and disturbances. Agricultural intensification is a major driver of biodiversity loss and has multiple components operating at different spatial scales: from in-field management intensity to landscape-scale simplification. Here we show that landscape-level effects dominate functional community composition and can even buffer the effects of in-field management intensification on functional homogenization, and that animal communities in real-world managed landscapes show a unified response (across orders and guilds) to both landscape-scale simplification and in-field intensification. Adults and larvae with specialized feeding habits, species with shorter activity periods and relatively small body sizes are selected against in simplified landscapes with intense in-field management. Our results demonstrate that the diversity of land cover types at the landscape scale is critical for maintaining communities, which are functionally diverse, even in landscapes where in-field management intensity is high.

Plant size is a major predictor of ecological functioning. We tested the hypothesis that feeding damage to plants increases with plant size, as the conspicuousness of large plants makes resource finding and colonisation easier. Further, large plants can be attractive to herbivores, as they offer greater amounts and ranges of resources and niches, but direct evidence from experiments testing size effects on feeding damage and consequently on plant fitness is so far missing. We established a common garden experiment with a plant size gradient (10-130 cm height) using 21 annual Brassicaceae species, and quantified plant size, biomass and number of all aboveground components (flowers, fruits, leaves, stems) and their proportional feeding damage. Plant reproductive fitness was measured using seed number, 1000 seed weight and total seed weight. Feeding damage to the different plant components increased with plant size or component biomass, with mean damage levels being approximately 30 % for flowers, 5 % for fruits and 1 % for leaves and stems. Feeding damage affected plant reproductive fitness depending on feeding damage type, with flower damage having the strongest effect, shown by greatly reduced seed number, 1000 seed weight and total seed weight. Finally, we found an overall negative effect of plant size on 1000 seed weight, but not on seed number and total seed weight. In conclusion, being conspicuous and attractive to herbivores causes greater flower damage leading to higher fitness costs for large plants, which might be partly counterbalanced by benefits such as enhanced competitive/compensatory abilities or more mutualistic pollinator visits.

Large plants are often more conspicuous and more attractive for associated animals than small plants, e.g. due to their wider range of resources. Therefore, plant size can positively affect species richness of associated animals, as shown for single groups of herbivores, but studies usually consider intraspecific size differences of plants in unstandardised environments. As comprehensive tests of interspecific plant size differences under standardised conditions are missing so far, we investigated effects of plant size on species richness of all associated arthropods using a common garden experiment with 21 Brassicaceae species covering a broad interspecific plant size gradient from 10 to 130 cm height. We recorded plant associated ecto- and endophagous herbivores, their natural enemies and pollinators on and in each aboveground plant organ, i.e. flowers, fruits, leaves and stems. Plant size (measured as height from the ground), the number of different plant organ entities and their biomass were assessed. Increasing plant size led to increased species richness of associated herbivores, natural enemies and pollinating insects. This pattern was found for ectophagous and endophagous herbivores, their natural enemies, as well as for herbivores associated with leaves and fruits and their natural enemies, independently of the additional positive effects of resource availability (i.e. organ biomass or number of entities and, regarding natural enemies, herbivore species richness). We found a lower R2 for pollinators compared to herbivores and natural enemies, probably caused by the high importance of flower characteristics for pollinator species richness besides plant size. Overall, the increase in plant height from 10 to 130 cm led to a 2.7-fold increase in predicted total arthropod species richness. In conclusion, plant size is a comprehensive driver of species richness of the plant associated arthropods, including pollinators, herbivores and their natural enemies, whether they are endophagous or ectophagous or associated with leaves or fruits.

Global change, especially land-use intensification, affects human well-being by impacting the delivery of multiple ecosystem services (multifunctionality). However, whether biodiversity loss is a major component of global change effects on multifunctionality in real-world ecosystems, as in experimental ones, remains unclear. Therefore, we assessed biodiversity, functional composition and 14 ecosystem services on 150 agricultural grasslands differing in land-use intensity. We also introduce five multifunctionality measures in which ecosystem services were weighted according to realistic land-use objectives. We found that indirect land-use effects, i.e. those mediated by biodiversity loss and by changes to functional composition, were as strong as direct effects on average. Their strength varied with land-use objectives and regional context. Biodiversity loss explained indirect effects in a region of intermediate productivity and was most damaging when land-use objectives favoured supporting and cultural services. In contrast, functional composition shifts, towards fast-growing plant species, strongly increased provisioning services in more inherently unproductive grasslands., (© 2015 The Authors Ecology Letters published by John Wiley & Sons Ltd and CNRS.)

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.

Background. Up to 75% of crop species benefit at least to some degree from animal pollination for fruit or seed set and yield. However, basic information on the level of pollinator dependence and pollinator contribution to yield is lacking for many crops. Even less is known about how insect pollination affects crop quality. Given that habitat loss and agricultural intensification are known to decrease pollinator richness and abundance, there is a need to assess the consequences for different components of crop production. Methods. We used pollination exclusion on flowers or inflorescences on a whole plant basis to assess the contribution of insect pollination to crop yield and quality in four flowering crops (spring oilseed rape, field bean, strawberry, and buckwheat) located in four regions of Europe. For each crop, we recorded abundance and species richness of flower visiting insects in ten fields located along a gradient from simple to heterogeneous landscapes. Results. Insect pollination enhanced average crop yield between 18 and 71% depending on the crop. Yield quality was also enhanced in most crops. For instance, oilseed rape had higher oil and lower chlorophyll contents when adequately pollinated, the proportion of empty seeds decreased in buckwheat, and strawberries' commercial grade improved; however, we did not find higher nitrogen content in open pollinated field beans. Complex landscapes had a higher overall species richness of wild pollinators across crops, but visitation rates were only higher in complex landscapes for some crops. On the contrary, the overall yield was consistently enhanced by higher visitation rates, but not by higher pollinator richness. Discussion. For the four crops in this study, there is clear benefit delivered by pollinators on yield quantity and/or quality, but it is not maximized under current agricultural intensification. Honeybees, the most abundant pollinator, might partially compensate the loss of wild pollinators in some areas, but our results suggest the need of landscape-scale actions to enhance wild pollinator populations.

Although temporal heterogeneity is a well-accepted driver of biodiversity, effects of interannual variation in land-use intensity (LUI) have not been addressed yet. Additionally, responses to land use can differ greatly among different organisms; therefore, overall effects of land-use on total local biodiversity are hardly known. To test for effects of LUI (quantified as the combined intensity of fertilization, grazing, and mowing) and interannual variation in LUI (SD in LUI across time), we introduce a unique measure of whole-ecosystem biodiversity, multidiversity. This synthesizes individual diversity measures across up to 49 taxonomic groups of plants, animals, fungi, and bacteria from 150 grasslands. Multidiversity declined with increasing LUI among grasslands, particularly for rarer species and aboveground organisms, whereas common species and belowground groups were less sensitive. However, a high level of interannual variation in LUI increased overall multidiversity at low LUI and was even more beneficial for rarer species because it slowed the rate at which the multidiversity of rare species declined with increasing LUI. In more intensively managed grasslands, the diversity of rarer species was, on average, 18% of the maximum diversity across all grasslands when LUI was static over time but increased to 31% of the maximum when LUI changed maximally over time. In addition to decreasing overall LUI, we suggest varying LUI across years as a complementary strategy to promote biodiversity conservation.

Pollination improves the yield of most crop species and contributes to one-third of global crop production, but comprehensive benefits including crop quality are still unknown. Hence, pollination is underestimated by international policies, which is particularly alarming in times of agricultural intensification and diminishing pollination services. In this study, exclusion experiments with strawberries showed bee pollination to improve fruit quality, quantity and market value compared with wind and self-pollination. Bee-pollinated fruits were heavier, had less malformations and reached higher commercial grades. They had increased redness and reduced sugar-acid-ratios and were firmer, thus improving the commercially important shelf life. Longer shelf life reduced fruit loss by at least 11%. This is accounting for 0.32 billion US$ of the 1.44 billion US$ provided by bee pollination to the total value of 2.90 billion US$ made with strawberry selling in the European Union 2009. The fruit quality and yield effects are driven by the pollination-mediated production of hormonal growth regulators, which occur in several pollination-dependent crops. Thus, our comprehensive findings should be transferable to a wide range of crops and demonstrate bee pollination to be a hitherto underestimated but vital and economically important determinant of fruit quality.

Pollination contributes to an estimated one third of global food production, through both the improvement of the yield and the quality of crops. Volatile compounds emitted by crop flowers mediate plant-pollinator interactions, but differences between crop varieties are still little explored. We investigated whether the visitation of crop flowers is determined by variety-specific flower volatiles using strawberry varieties (Fragaria x ananassa Duchesne) and how this affects the pollination services of the wild bee Osmia bicornis L. Flower volatile compounds of three strawberry varieties were measured via headspace collection. Gas chromatography showed that the three strawberry varieties produced the same volatile compounds but with quantitative differences of the total amount of volatiles and between distinct compounds. Electroantennographic recordings showed that inexperienced females of Osmia bicornis had higher antennal responses to all volatile compounds than to controls of air and paraffin oil, however responses differed between compounds. The variety Sonata was found to emit a total higher level of volatiles and also higher levels of most of the compounds that evoked antennal responses compared with the other varieties Honeoye and Darselect. Sonata also received more flower visits from Osmia bicornis females under field conditions, compared with Honeoye. Our results suggest that differences in the emission of flower volatile compounds among strawberry varieties mediate their attractiveness to females of Osmia bicornis. Since quality and quantity of marketable fruits depend on optimal pollination, a better understanding of the role of flower volatiles in crop production is required and should be considered more closely in crop-variety breeding.

The diversity and abundance of wild insect pollinators have declined in many agricultural landscapes. Whether such declines reduce crop yields, or are mitigated by managed pollinators such as honey bees, is unclear. We found universally positive associations of fruit set with flower visitation by wild insects in 41 crop systems worldwide. In contrast, fruit set increased significantly with flower visitation by honey bees in only 14% of the systems surveyed. Overall, wild insects pollinated crops more effectively; an increase in wild insect visitation enhanced fruit set by twice as much as an equivalent increase in honey bee visitation. Visitation by wild insects and honey bees promoted fruit set independently, so pollination by managed honey bees supplemented, rather than substituted for, pollination by wild insects. Our results suggest that new practices for integrated management of both honey bees and diverse wild insect assemblages will enhance global crop yields.

Understanding how landscape characteristics affect biodiversity patterns and ecological processes at local and landscape scales is critical for mitigating effects of global environmental change. In this review, we use knowledge gained from human-modified landscapes to suggest eight hypotheses, which we hope will encourage more systematic research on the role of landscape composition and configuration in determining the structure of ecological communities, ecosystem functioning and services. We organize the eight hypotheses under four overarching themes. Section A: 'landscape moderation of biodiversity patterns' includes (1) the landscape species pool hypothesis-the size of the landscape-wide species pool moderates local (alpha) biodiversity, and (2) the dominance of beta diversity hypothesis-landscape-moderated dissimilarity of local communities determines landscape-wide biodiversity and overrides negative local effects of habitat fragmentation on biodiversity. Section B: 'landscape moderation of population dynamics' includes (3) the cross-habitat spillover hypothesis-landscape-moderated spillover of energy, resources and organisms across habitats, including between managed and natural ecosystems, influences landscape-wide community structure and associated processes and (4) the landscape-moderated concentration and dilution hypothesis-spatial and temporal changes in landscape composition can cause transient concentration or dilution of populations with functional consequences. Section C: 'landscape moderation of functional trait selection' includes (5) the landscape-moderated functional trait selection hypothesis-landscape moderation of species trait selection shapes the functional role and trajectory of community assembly, and (6) the landscape-moderated insurance hypothesis-landscape complexity provides spatial and temporal insurance, i.e. high resilience and stability of ecological processes in changing environments. Section D: 'landscape constraints on conservation management' includes (7) the intermediate landscape-complexity hypothesis-landscape-moderated effectiveness of local conservation management is highest in structurally simple, rather than in cleared (i.e. extremely simplified) or in complex landscapes, and (8) the landscape-moderated biodiversity versus ecosystem service management hypothesis-landscape-moderated biodiversity conservation to optimize functional diversity and related ecosystem services will not protect endangered species. Shifting our research focus from local to landscape-moderated effects on biodiversity will be critical to developing solutions for future biodiversity and ecosystem service management., (© 2012 The Authors. Biological Reviews © 2012 Cambridge Philosophical Society.)

Sustainable agricultural landscapes by definition provide high magnitude and stability of ecosystem services, biodiversity and crop productivity. However, few studies have considered landscape effects on the stability of ecosystem services. We tested whether isolation from florally diverse natural and semi-natural areas reduces the spatial and temporal stability of flower-visitor richness and pollination services in crop fields. We synthesised data from 29 studies with contrasting biomes, crop species and pollinator communities. Stability of flower-visitor richness, visitation rate (all insects except honey bees) and fruit set all decreased with distance from natural areas. At 1 km from adjacent natural areas, spatial stability decreased by 25, 16 and 9% for richness, visitation and fruit set, respectively, while temporal stability decreased by 39% for richness and 13% for visitation. Mean richness, visitation and fruit set also decreased with isolation, by 34, 27 and 16% at 1 km respectively. In contrast, honey bee visitation did not change with isolation and represented > 25% of crop visits in 21 studies. Therefore, wild pollinators are relevant for crop productivity and stability even when honey bees are abundant. Policies to preserve and restore natural areas in agricultural landscapes should enhance levels and reliability of pollination services., (© 2011 Blackwell Publishing Ltd/CNRS.)

Global change may substantially affect biodiversity and ecosystem functioning but little is known about its effects on essential biotic interactions. Since different environmental drivers rarely act in isolation it is important to consider interactive effects. Here, we focus on how two key drivers of anthropogenic environmental change, climate change and the introduction of alien species, affect plant-pollinator interactions. Based on a literature survey we identify climatically sensitive aspects of species interactions, assess potential effects of climate change on these mechanisms, and derive hypotheses that may form the basis of future research. We find that both climate change and alien species will ultimately lead to the creation of novel communities. In these communities certain interactions may no longer occur while there will also be potential for the emergence of new relationships. Alien species can both partly compensate for the often negative effects of climate change but also amplify them in some cases. Since potential positive effects are often restricted to generalist interactions among species, climate change and alien species in combination can result in significant threats to more specialist interactions involving native species., (© 2010 The Authors. Biological Reviews © 2010 Cambridge Philosophical Society.)

Although habitat fragmentation and agricultural intensification are known as threads to pollinator diversity, little is known about consequences for population size and genetic diversity. Here, we combined detailed field observations, molecular approaches and GIS-based quantification of landscape structure (measured by proportions of seminatural habitats and proportions of mass flowering crops) to get new insights into driving forces of population dynamics of the bumblebee species Bombus pascuorum. Comparing 13 agriculturally dominated landscape sectors, we found the proportion of mass flowering crops to positively influence bumblebee abundance whereas the proportion of seminatural habitats was of minor importance. We used microsatellites to quantify landscape-related colony densities, inbreeding and population substructure. Detected colony densities did not correlate with landscape parameters or with local worker abundance, measured by field observations. These results indicate that increased worker abundances within landscapes are rather due to greater colony sizes than due to an increased number of nests. We found significant population substructure, measured by F(ST) and seven landscape sectors to bear significantly increased inbreeding values (F(IS)). F(IS) was strongly varying between sectors but did not correlate with landscape structure. Moreover, F(IS) had a significantly negative effect on colony size, demonstrating the importance of genetic diversity on population fitness at a landscape scale. We suggest that inbreeding levels might be related to the temporal variation of food resources and population sizes in agricultural landscapes.

Coexistence in bumblebee communities has largely been investigated at local spatial scales. However, local resource partitioning does not fully explain the species diversity of bumblebee communities. Theoretical studies provide new evidence that partitioning of space can promote species coexistence, when species interact with their environment at different spatial scales. If bumblebee species possess specific foraging ranges, different spatial resource utilisation patterns might operate as an additional mechanism of coexistence in bumblebee communities. We investigated the effects of the landscape-wide availability of different resources (mass flowering crops and semi-natural habitats) on the local densities of four bumblebee species at 12 spatial scales (landscape sectors with 250-3,000 m radius) to indirectly identify the spatial scales at which the bumblebees perceive their environment. The densities of all bumblebee species were enhanced in landscapes with high proportions of mass flowering crops (mainly oilseed rape). We found the strongest effects for Bombus terrestris agg. and Bombus lapidarius at large spatial scales, implying foraging distances of 3,000 and 2,750 m, respectively. The densities of Bombus pascuorum were most strongly influenced at a medium spatial scale (1,000 m), and of Bombus pratorum (with marginal significance) at a small spatial scale (250 m). The estimated foraging ranges tended to be related to body and colony sizes, indicating that larger species travel over larger distances than smaller species, presumably enabling them to build up larger colonies through a better exploitation of food resources. We conclude that coexistence in bumblebee communities could potentially be mediated by species-specific differences in the spatial resource utilisation patterns, which should be considered in conservation schemes.