Evidence of a European seed dispersal crisis.

Seed dispersal is crucial for ecosystem persistence, especially in fragmented landscapes, such as those common in Europe. Ongoing defaunation might compromise effective seed dispersal, but the conservation status of pairwise interactions remains unknown. With a literature review, we reconstructed the first European-wide seed dispersal network and evaluated the conservation status of interactions by assessing each interacting partner's IUCN (International Union for Conservation of Nature) conservation status and population trends. We found that a third of the disperser species and interactions face potential extinction and that 30% of the plant species have most of their dispersers threatened or declining. Our study reveals a developing seed dispersal crisis in Europe and highlights large knowledge gaps regarding the dispersers and conservation status of zoochorous plants, urging further scrutiny and action to conserve the seed dispersal service.

Isolation limits spring pollination in a UK fragmented landscape.

Animal-mediated pollination determines the reproductive success of most flowering plants; this process however can be disrupted by environmental degradation, with habitat loss and fragmentation highlighted as a top driver of pollination deficits. Despite being a pervasive stressor worldwide, we still have rather limited empirical evidence on its effects on pollination services, especially for early spring pollination syndromes. We investigate this using a potted plant phytometry experiment in which we placed English Bluebell (Hyacinthoides non-scripta)-a species largely pollinated in spring-into a fragmented woodland habitat. We selected 51 woodland patches which varied in both size and distance from each other and placed 153 pots of bluebell plants in the patches for c.4 weeks to measure pollination. The woodlands were located in a matrix of grassland, the latter being of low plant species richness and overall in the patches, woodland plants showed a positive species-area relationship. We collected traits on bluebell reproduction, these included the number, size, quality of seeds, the number of seed capsules and the number of flowers that failed to set any seeds. We found that seed traits responded differently to patch area and isolation. Patch isolation negatively affected the number of seeds and capsules, whilst it did not affect the size and quality of seeds. Patch area had no effect on any traits, suggesting that patch area might not necessarily be a factor that affects pollination in this species. The number of flowers that failed to set seed was unaffected by either patch area or isolation. Our study suggests that woodland fragmentation impacts the pollination of understory spring flowering plants. Our results highlight the use of multiple traits of phytometer plants to evaluate pollination and the importance of connectivity in maintaining pollination services in small-fragmented landscapes.

Multi-habitat landscapes are more diverse and stable with improved function.

Conservation, restoration and land management are increasingly implemented at landscape scales1,2. However, because species interaction data are typically habitat- and/or guild-specific, exactly how those interactions connect habitats and affect the stability and function of communities at landscape scales remains poorly understood. We combine multi-guild species interaction data (plant-pollinator and three plant-herbivore-parasitoid communities, collected from landscapes with one, two or three habitats), a field experiment and a modelling approach to show that multi-habitat landscapes support higher species and interaction evenness, more complementary species interactions and more consistent robustness to species loss. These emergent network properties drive improved pollination success in landscapes with more habitats and are not explained by simply summing component habitat webs. Linking landscape composition, through community structure, to ecosystem function, highlights mechanisms by which several contiguous habitats can support landscape-scale ecosystem services.

Spillover effects from invasive Acacia alter the plant-pollinator networks and seed production of native plants.

Invasive flowering plants can disrupt plant-pollinator networks. This is well documented where invasives occur amongst native plants; however, the potential for 'spillover' effects of invasives that form stands in adjacent habitats are less well understood. Here we quantify the impact of two invasive Australian species, Acacia saligna and Acacia longifolia, on the plant-pollinator networks in fynbos habitats in South Africa. We compared networks from replicate 1 ha plots of native vegetation (n = 21) that were subjected to three treatments: (1) at least 400 m from flowering Acacia; (2) adjacent to flowering Acacia, or (3) adjacent to flowering Acacia where all Acacia flowers were manually removed. We found that native flowers adjacent to stands of flowering Acacia received significantly more insect visits, especially from beetles and Apis mellifera capensis, and that visitation was more generalized. We also recorded visitation to, and the seed set of, three native flowering species and found that two received more insect visits, but produced fewer seeds, when adjacent to flowering Acacia. Our research shows that 'spillover' effects of invasive Acacia can lead to significant changes in visitation and seed production of native co-flowering species in neighbouring habitats-a factor to be considered when managing invaded landscapes.

Decline in Honeybees and Its Consequences for Beekeepers and Crop Pollination in Western Nepal.

In understudied regions of the world, beekeeper records can provide valuable insights into changes in pollinator population trends. We conducted a questionnaire survey of 116 beekeepers in a mountainous area of Western Nepal, where the native honeybee Apis cerana cerana is kept as a managed bee. We complemented the survey with field data on insect-crop visitation, a household income survey, and an interview with a local lead beekeeper. In total, 76% of beekeepers reported declines in honeybees, while 86% and 78% reported declines in honey yield and number of beehives, respectively. Honey yield per hive fell by 50% between 2012 and 2022, whilst the number of occupied hives decreased by 44%. Beekeepers ranked climate change and declining flower abundance as the most important drivers of the decline. This raises concern for the future food and economic security of this region, where honey sales contribute to 16% of total household income, and where Apis cerana cerana plays a major role in crop pollination, contributing more than 50% of all flower visits to apple, cucumber, and pumpkin. To mitigate further declines, we promote native habitat and wildflower preservation, and using well-insulated log hives to buffer bees against the increasingly extreme temperature fluctuations.

Quantifying the production of plant pollen at the farm scale.

Plant pollen is rich in protein, sterols and lipids, providing crucial nutrition for many pollinators. However, we know very little about the quantity, quality and timing of pollen availability in real landscapes, limiting our ability to improve food supply for pollinators. We quantify the floral longevity and pollen production of a whole plant community for the first time, enabling us to calculate daily pollen availability. We combine these data with floral abundance and nectar measures from UK farmland to quantify pollen and nectar production at the landscape scale throughout the year. Pollen and nectar production were significantly correlated at the floral unit, and landscape level. The species providing the highest quantity of pollen on farmland were Salix spp. (38%), Filipendula ulmaria (14%), Rubus fruticosus (10%) and Taraxacum officinale (9%). Hedgerows were the most pollen-rich habitats, but permanent pasture provided the majority of pollen at the landscape scale, because of its large area. Pollen and nectar were closely associated in their phenology, with both peaking in late April, before declining steeply in June and remaining low throughout the year. Our data provide a starting point for including pollen in floral resource assessments and ensuring the nutritional requirements of pollinators are met in farmland landscapes.

Fertilization using manure minimizes the trade-offs between biodiversity and forage production in agri-environment scheme grasslands.

A common practice used to restore and maintain biodiversity in grasslands is to stop or decrease the use of fertilizers as they are a major cause of biodiversity loss. This practice is problematic for farmers who need fertilizers to increase forage and meet the nutritional needs of livestock. Evidence is needed that helps identify optimal fertilizer regimes that could benefit biodiversity and livestock production simultaneously over the long-term. Here, we evaluated the impact of different fertilizer regimes on indicators related to both biodiversity (plant, pollinator, leaf miners and parasitoid Shannon-Weiner diversity, bumblebee abundance, nectar productivity and forb species richness), and forage production (ash, crude protein, ruminant metabolizable energy and dry matter). To this end, we used data from a grassland restoration experiment managed under four nutrient inputs schemes for 27 years: farmyard manure (FYM; 72 kg N ha-1 yr-1), artificial nitrogen-phosphorus and potassium (NPK; 25 kg N ha-1 yr-1), FYM + NPK (97 kg N ha-1 yr-1) and no-fertilizer. Results showed strong trade-offs between biodiversity and forage production under all treatments even in applications lower than the critical load in the EU. Overall, farmyard manure was the fertilizer that optimized production and biodiversity while 97 kg N ha-1 yr-1 of fertilizer addition (FYM+NPK) had the most negative impact on biodiversity. Finally, forage from places where no fertilizer has been added for 27 years did not meet the nutritional requirements of cattle, but it did for sheep. Rethinking typical approaches of nutrient addition could lead to land management solutions suitable for biological conservation and agriculture.

Corridor quality buffers extinction under extreme droughts in experimental metapopulations.

Corridors with good-quality habitats maintain the spatial dynamics of metapopulations by promoting dispersal between habitat patches, potentially buffering populations, and communities against continued global change. However, this function is threatened by habitats becoming increasingly fragmented, and habitat matrices becoming increasingly inhospitable, potentially reducing the resilience and persistence of populations. Yet, we lack a clear understanding of how reduced corridor quality interacts with rates of environmental change to destabilize populations. Using laboratory microcosms containing metapopulations of the Collembola Folsomia candida, we investigate the impact of corridor quality on metapopulation persistence under a range of simulated droughts, a key stressor for this species. We manipulated both drought severity and the number of patches affected by drought across landscapes connected by either good- or poor-quality corridors. We measured the time of metapopulation extinction, the maximum rate of metapopulation decline, and the variability of abundance among patches as criteria to evaluate the persistence ability of metapopulations. We show that while drought severity negatively influenced the time of metapopulation extinction and the increase in drought patches caused metapopulation decline, these results were mitigated by good-quality corridors, which increased metapopulation persistence time and decreased both how fast metapopulations declined and the interpatch variability in abundances. Our results suggest that enhancing corridor quality can increase the persistence of metapopulations, increasing the time available for conservation actions to take effect, and/or for species to adapt or move in the face of continued stress. Given that fragmentation increases the isolation of habitats, improving the quality of habitat corridors may provide a useful strategy to enhance the resistance of spatially structured populations.

Landscape configuration affects probability of apex predator presence and community structure in experimental metacommunities.

Biodiversity is declining at an unprecedented rate, highlighting the urgent requirement for well-designed protected areas. Design tactics previously proposed to promote biodiversity include enhancing the number, connectivity, and heterogeneity of reserve patches. However, how the importance of these features changes depending on what the conservation objective is remains poorly understood. Here we use experimental landscapes containing ciliate protozoa to investigate how the number and heterogeneity in size of habitat patches, rates of dispersal between neighbouring patches, and mortality risk of dispersal across the non-habitat 'matrix' interact to affect a number of diversity measures. We show that increasing the number of patches significantly increases γ diversity and reduces the overall number of extinctions, whilst landscapes with heterogeneous patch sizes have significantly higher γ diversity than those with homogeneous patch sizes. Furthermore, the responses of predators depended on their feeding specialism, with generalist predator presence being highest in a single large patch, whilst specialist predator presence was highest in several-small patches with matrix dispersal. Our evidence emphasises the importance of considering multiple diversity measures to disentangle community responses to patch configuration.

Landscape-scale drivers of pollinator communities may depend on land-use configuration.

Research into pollinators in managed landscapes has recently combined approaches of pollination ecology and landscape ecology, because key stressors are likely to interact across wide areas. While laboratory and field experiments are valuable for furthering understanding, studies are required to investigate the interacting drivers of pollinator health and diversity across a broader range of landscapes and a wider array of taxa. Here, we use a network of 96 study landscapes in six topographically diverse regions of Britain, to test the combined importance of honeybee density, insecticide loadings, floral resource availability and habitat diversity to pollinator communities. We also explore the interactions between these drivers and the cover and proximity of semi-natural habitat. We found that among our four drivers, only honeybee density was positively related to wild pollinator abundance and diversity, and the positive association between abundance and floral resources depended on insecticide loadings and habitat diversity. By contrast, our exploratory models including habitat composition metrics revealed a complex suite of interactive effects. These results demonstrate that improving pollinator community composition and health is unlikely to be achieved with general resource enhancements only. Rather, local land-use context should be considered in fine-tuning pollinator management and conservation. This article is part of the theme issue 'Natural processes influencing pollinator health: from chemistry to landscapes'.

Interaction generalisation and demographic feedbacks drive the resilience of plant-insect networks to extinctions.

Understanding the processes driving ecological resilience, that is the extent to which systems retain their structure while absorbing perturbations, is a central challenge for theoretical and applied ecologists. Plant-insect assemblages are well-suited for the study of ecological resilience as they are species-rich and encompass a variety of ecological interactions that correspond to essential ecosystem functions. Mechanisms affecting community response to perturbations depend on both the natural history and structure of ecological interactions. Natural history attributes of the interspecific interactions, for example whether they are mutualistic or antagonistic, may affect the ecological resilience by controlling the demographic feedbacks driving ecological dynamics at the community level. Interaction generalisation may also affect resilience, by defining opportunities for interaction rewiring, the extent to which species are able to switch interactions in fluctuating environments. These natural history attributes may also interact with network structure to affect ecological resilience. Using adaptive network models, we investigated the resilience of plant-pollinator and plant-herbivore networks to species loss. We specifically investigated how fundamental natural history differences between these systems, namely the demographic consequences of the interaction and their level of generalisation-mediating rewiring opportunities-affect the resilience of dynamic ecological networks to extinctions. We also create a general benchmark for the effect of network structure on resilience simulating extinctions on theoretical networks with controlled structures. When network structure was static, pollination networks were less resilient than herbivory networks; this is related to their high levels of nestedness and the reciprocally positive feedbacks that define mutualisms, which made co-extinction cascades more likely and longer in plant-pollinator assemblages. When considering interaction rewiring, the high generalisation and the structure of pollination networks boosted their resilience to extinctions, which approached those of herbivory networks. Simulation results using theoretical networks suggested that the empirical structure of herbivory networks may protect them from collapse. Elucidating the ecological and evolutionary processes driving interaction rewiring is key to understanding the resilience of plant-insect assemblages. Accounting for rewiring requires ecologists to combine natural history with network models that incorporate feedbacks between species abundances, traits and interactions. This combination will elucidate how perturbations propagate at community level, reshaping biodiversity structure and ecosystem functions.

Compreender os processos que governam a resiliência dos sistemas ecológicos, i.e. o quanto sistemas ecológicos conservam sua estrutura enquanto absorvem perturbações, é um desafio central para ecólogos teóricos e aplicados. Comunidades de insetos e plantas são bons modelos para o estudo da resiliência ecológica pois são ricos em espécies, representando uma grande diversidade de interações ecológicas que correspondem a serviços ecossistêmicos essenciais. Os mecanismos que afetam a resposta de comunidades ecológicas a perturbações dependem tanto da história natural quanto da estrutura das interações ecológicas. A história natural de interações interespecíficas, e.g. se a interação é mutualística ou antagonística, pode afetar a resiliência do sistema ao controlar as retroalimentações demográficas que governam a dinâmica ecológica no nível da comunidade. Generalismo nas interações também pode afetar resiliência ao definir as oportunidades de rewiring de interações, i.e. o quanto espécies são capazes de mudar interações em ambientes instáveis. Atributos da história natural das interações podem também interagir com a estrutura de redes ecológicas de forma a influenciar a resiliência de sistemas ecológicos. Usando modelos de redes adaptativas, investigamos a resiliência de redes polinizador-planta e herbívoro-planta à perda de espécies. Especificamente, investigamos como diferenças fundamentais na história natural dos dois sistemas, isto é, as consequências demográficas da interação e seu grau de generalização - que mediam as oportunidades de rewiring - afetam a resiliência de redes ecológicas dinâmicas a extinções. Também criamos um referencial teórico e abrangente para o efeito da estrutura das redes em sua resiliência, simulando extinções em redes teóricas com estruturas controladas. Quando a estrutura das redes foi considerada estática, redes de polinização foram menos resilientes do que redes de herbivoria; o que está associado aos maiores níveis de aninhamento e aos efeitos demográficos positivos e recíprocos que definem mutualismos, aumentando a probabilidade e o comprimento das cascatas de extinção em redes polinizador-planta. Ao incorporar rewiring de interações, a alta generalização e a estrutura das redes de polinização impulsionou sua resiliência a extinções, que se aproximou da resiliência de redes de herbivoria. Os resultados das simulações com redes teóricas sugerem que estrutura de redes de herbivoria protegem esses sistemas do colapso. Compreender quais processos ecológicos e evolutivos governam o rewiring de interações é chave se queremos prever a resiliência de sistemas inseto-planta. Para incorporar rewiring de interações, será necessário combinar conhecimento sobre história natural com modelos de rede que incorporem a retroalimentação entre abundâncias, atributos e interações das espécies envolvidas. Essa combinação elucidará como perturbações se propagam no nível de comunidades ecológicas, reconfigurando a estrutura da biodiversidade e suas funções ecossistêmicas.

Corridor quality affects net movement, size of dispersers, and population growth in experimental microcosms.

Corridors are expected to increase species dispersal in fragmented habitats. However, it remains unclear how the quality of corridors influences the dispersal process, and how it interacts with corridor length and width. Here we investigate these factors using a small-scale laboratory system where we track the dispersal of the model organism Collembola Folsomia candida. Using this system, we study the effects of corridor length, width, and quality on the probability of dispersal, net movement, body size of dispersers, and the rate of change in population size after colonization. We show that corridor quality positively affected dispersal probability, net movement, and the rate of change in population size in colonised patches. Moreover, corridor quality significantly affected the size of dispersers, with only larger individuals dispersing through poor quality corridors. The length and width of corridors affected both the rate at which populations increased in colonised patches and the net number of individuals which dispersed, suggesting that these physical properties may be important in maintaining the flow of individuals in space. Our results thus suggest that corridor quality can have an important role in determining not only the probability of dispersal occurs but also the phenotypes of the individuals which disperse, with concomitant effects on the net movement of individuals and the rate of change in population size in the colonised patches.

Urban gulls adapt foraging schedule to human-activity patterns.

Numerous animals are able to adapt to temporal patterns in natural food availability, but whether species living in relatively novel environments such as cities can adapt to anthropogenic activity cycles is less well understood. We aimed to assess the extent to which urban gulls have adapted their foraging schedule to anthropogenic food source fluctuations related to human activity by combining field observations at three distinct urban feeding grounds (park, school and waste centre) with global positioning system (GPS) tracking data of gulls visiting similar types of feeding grounds throughout the same city. We found that the birds' foraging patterns closely matched the timing of school breaks and the opening and closing times of the waste centre, but gull activity in the park appeared to correspond to the availability of natural food sources. Overall, this suggests that gulls may have the behavioural flexibility to adapt their foraging behaviour to human time schedules when beneficial and that this trait could potentially enable them to thrive in cities.

Differential effects of fertilisers on pollination and parasitoid interaction networks.

Grassland fertilisation drives non-random plant loss resulting in areas dominated by perennial grass species. How these changes cascade through linked trophic levels, however, is not well understood. We studied how grassland fertilisation propagates change through the plant assemblage into the plant-flower-visitor, plant-leaf miner and leaf miner-parasitoid networks using a year's data collection from a long-term grassland fertiliser application experiment. Our experiment had three fertiliser treatments each applied to replicate plots 15 m2 in size: mineral fertiliser, farmyard manure, and mineral fertiliser and farmyard manure combined, along with a control of no fertiliser. The combined treatment had the most significant impact, and both plant species richness and floral abundance decreased with the addition of fertiliser. While insect species richness was unaffected by fertiliser treatment, fertilised plots had a significantly higher abundance of leaf miners and parasitoids and a significantly lower abundance of bumblebees. The plant-flower-visitor and plant-herbivore networks showed higher values of vulnerability and lower modularity with fertiliser addition, while leaf miner-parasitoid networks showed a rise in generality. The different groups of insects were impacted by fertilisers to varying degrees: while the effect on abundance was the highest for leaf miners, the vulnerability and modularity of flower-visitor networks was the most affected. The impact on the abundance of leaf miners was positive and three times higher than the impact on parasitoids, and the impact on bumblebee abundance was negative and double the magnitude of impact on flower abundance. Overall, our results show that while insect species richness was unaffected by fertilisers, network structure changed significantly as the replacement of forbs by grasses resulted in changes in relative abundance across trophic levels, with the direction of change depending on the type of network. Synthesis. By studying multiple networks simultaneously, we were able to rank the relative impact of habitat change on the different groups of species within the community. This provided a more holistic picture of the impact of agricultural intensification and provides useful information when deciding on priorities for mitigation.

Reshaping our understanding of species' roles in landscape-scale networks.

In network ecology, landscape-scale processes are often overlooked, yet there is increasing evidence that species and interactions spill over between habitats, calling for further study of interhabitat dependencies. Here, we investigate how species connect a mosaic of habitats based on the spatial variation of their mutualistic and antagonistic interactions using two multilayer networks, combining pollination, herbivory and parasitism in the UK and New Zealand. Developing novel methods of network analysis for landscape-scale ecological networks, we discovered that few plant and pollinator species acted as connectors or hubs, both within and among habitats, whereas herbivores and parasitoids typically have more peripheral network roles. Insect species' roles depend on factors other than just the abundance of taxa in the lower trophic level, exemplified by larger Hymenoptera connecting networks of different habitats and insects relying on different resources across different habitats. Our findings provide a broader perspective for landscape-scale management and ecological community conservation.

A systems approach reveals urban pollinator hotspots and conservation opportunities.

Urban areas are often perceived to have lower biodiversity than the wider countryside, but a few small-scale studies suggest that some urban land uses can support substantial pollinator populations. We present a large-scale, well-replicated study of floral resources and pollinators in 360 sites incorporating all major land uses in four British cities. Using a systems approach, we developed Bayesian network models integrating pollinator dispersal and resource switching to estimate city-scale effects of management interventions on plant-pollinator community robustness to species loss. We show that residential gardens and allotments (community gardens) are pollinator 'hotspots': gardens due to their extensive area, and allotments due to their high pollinator diversity and leverage on city-scale plant-pollinator community robustness. Household income was positively associated with pollinator abundance in gardens, highlighting the influence of socioeconomic factors. Our results underpin urban planning recommendations to enhance pollinator conservation, using increasing city-scale community robustness as our measure of success.

Impact of cyber-invasive species on a large ecological network.

As impacts of introduced species cascade through trophic levels, they can cause indirect and counter-intuitive effects. To investigate the impact of invasive species at the network scale, we use a generalized food web model, capable of propagating changes through networks with a series of ecologically realistic criteria. Using data from a small British offshore island, we quantify the impacts of four virtual invasive species (an insectivore, a herbivore, a carnivore and an omnivore whose diet is based on a rat) and explore which clusters of species react in similar ways. We find that the predictions for the impacts of invasive species are ecologically plausible, even in large networks. Species in the same taxonomic group are similarly impacted by a virtual invasive species. However, interesting differences within a given taxonomic group can occur. The results suggest that some native species may be at risk from a wider range of invasives than previously believed. The implications of these results for ecologists and land managers are discussed.

A global synthesis of the effects of diversified farming systems on arthropod diversity within fields and across agricultural landscapes.

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.

Modest enhancements to conventional grassland diversity improve the provision of pollination services.

Grassland for livestock production is a major form of land use throughout Europe and its intensive management threatens biodiversity and ecosystem functioning in agricultural landscapes. Modest increases to conventional grassland biodiversity could have considerable positive impacts on the provision of ecosystem services, such as pollination, to surrounding habitats.Using a field-scale experiment in which grassland seed mixes and sward management were manipulated, complemented by surveys on working farms and phytometer experiments, the impact of conventional grassland diversity and management on the functional diversity and ecosystem service provision of pollinator communities were investigated.Increasing plant richness, by the addition of both legumes and forbs, was associated with significant enhancements in the functional diversity of grassland pollinator communities. This was associated with increased temporal stability of flower-visitor interactions at the community level. Visitation networks revealed pasture species Taraxacum sp. (Wigg.) (dandelion) and Cirsium arvense (Scop.) (creeping thistle) to have the highest pollinator visitation frequency and richness. Cichorium intybus (L.) (chichory) was highlighted as an important species having both high pollinator visitation and desirable agronomic properties.Increased sward richness was associated with an increase in the pollination of two phytometer species; Fragaria × ananassa (strawberry) and Silene dioica (red campion), but not Vicia faba (broad bean). Enhanced functional diversity, richness and abundance of the pollinator communities associated with more diverse neighbouring pastures were found to be potential mechanisms for improved pollination. Synthesis and applications. A modest increase in conventional grassland plant diversity with legumes and forbs, achievable with the expertise and resources available to most grassland farmers, could enhance pollinator functional diversity, richness and abundance. Moreover, our results suggest that this could improve pollination services and consequently surrounding crop yields (e.g. strawberry) and wildflower reproduction in agro-ecosystems.

Food for Pollinators: Quantifying the Nectar and Pollen Resources of Urban Flower Meadows.

Planted meadows are increasingly used to improve the biodiversity and aesthetic amenity value of urban areas. Although many 'pollinator-friendly' seed mixes are available, the floral resources these provide to flower-visiting insects, and how these change through time, are largely unknown. Such data are necessary to compare the resources provided by alternative meadow seed mixes to each other and to other flowering habitats. We used quantitative surveys of over 2 million flowers to estimate the nectar and pollen resources offered by two exemplar commercial seed mixes (one annual, one perennial) and associated weeds grown as 300m2 meadows across four UK cities, sampled at six time points between May and September 2013. Nectar sugar and pollen rewards per flower varied widely across 65 species surveyed, with native British weed species (including dandelion, Taraxacum agg.) contributing the top five nectar producers and two of the top ten pollen producers. Seed mix species yielding the highest rewards per flower included Leontodon hispidus, Centaurea cyanus and C. nigra for nectar, and Papaver rhoeas, Eschscholzia californica and Malva moschata for pollen. Perennial meadows produced up to 20x more nectar and up to 6x more pollen than annual meadows, which in turn produced far more than amenity grassland controls. Perennial meadows produced resources earlier in the year than annual meadows, but both seed mixes delivered very low resource levels early in the year and these were provided almost entirely by native weeds. Pollen volume per flower is well predicted statistically by floral morphology, and nectar sugar mass and pollen volume per unit area are correlated with flower counts, raising the possibility that resource levels can be estimated for species or habitats where they cannot be measured directly. Our approach does not incorporate resource quality information (for example, pollen protein or essential amino acid content), but can easily do so when suitable data exist. Our approach should inform the design of new seed mixes to ensure continuity in floral resource availability throughout the year, and to identify suitable species to fill resource gaps in established mixes.