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Biodiversity conservation faces a methodological conundrum: Biodiversity measurement often relies on species, most of which are rare at various scales, especially prone to extinction under global change, but also the most challenging to sample and model. Predicting the distribution change of rare species using conventional species distribution models is challenging because rare species are hardly captured by most survey systems. When enough data is available, predictions are usually spatially biased toward locations where the species is most likely to occur, violating the assumptions of many modelling frameworks. Workflows to predict and eventually map rare species distributions imply important trade-offs between data quantity, quality, representativeness, and model complexity that need to be considered prior to survey and analysis. Our opinion is that study designs need to carefully integrate the different steps, from species sampling to modelling, in accordance to the different types of rarity and available data in order to improve our capacity for sound assessment and prediction of rare species distribution. In this article, we summarize and comment on how different categories of species rarity lead to different types of occurrence and distribution data depending on choices made during the survey process, namely the spatial distribution of samples (where to sample) and the sampling protocol in each selected location (how to sample). We then clarify which species distribution models are suitable depending on the different types of distribution data (how to model). Among others, for most rarity forms, we highlight the insights from systematic species-targeted sampling coupled with hierarchical models that allow correcting for overdispersion and for spatial and sampling sources of bias. Our article provides scientists and practitioners with a much-needed guide through the ever-increasing diversity of methodological developments to improve prediction of rare species

Pollinators experience large spatiotemporal fluctuations in resource availability when mass-flowering crops are rotated with resource-poor cereal crops. Yet, few studies have considered the effect this has on pollinator population stability, nor how this might be mitigated to maintain consistent crop pollination services. We assess the potential of boundary features (standard narrow 1 m grassy margins, hedgerows and wide 4 m agri-environment margins) to support and stabilise pollinator populations and pollination service in agricultural landscapes under crop rotation. Assuming a 6-year rotation, we use a process-based pollinator model to predict yearly pollinator population size and in-crop visitation rates to oilseed rape and field bean across 117 study landscapes in England with varying amounts of boundary features. We model both ground-nesting bumblebees and solitary bees and compare the predictions including and excluding boundary features from the landscapes. Ground-nesting bumblebee populations, whose longer-lifetime colonies benefit from continuity of resources, were larger and more stable (relative to the no-features scenario) in landscapes with more boundary features. Ground-nesting solitary bee populations were also larger but not significantly more stable, except with the introduction of wide permanent agri-environment margins, due to their shorter lifetimes and shorter foraging/dispersal ranges. Crop visitation by ground-nesting bumblebees was greater and more stable in landscapes with more boundary features, partly due to increased colony growth prior to crop flowering. Time averaged crop visitation by ground-nesting solitary bees was slightly lower, due to females dividing their foraging time between boundary features and the crop. However, despite this, the minimum pollination service delivered was higher, due to the more stable delivery. Synthesis and applications. Field boundary features have an important role in stabilising pollinator populations a

Atmospheric nitrogen deposition and other sources of environmental eutrophication have increased substantially over the past century worldwide, notwithstanding the recent declining trends in Europe. Despite the recognized susceptibility of plants to eutrophication, few studies evaluated how impacts propagate to consumers, such as pollinators. Here we aim to test if soil eutrophication contributes to the temporal dynamics of pollinators and their larval resources.We used a temporally and spatially explicit historical dataset with information on species occurrences to test if soil eutrophication, and more specifically nitrogen deposition, contributes to the patterns of change of plant and pollinator richness in the Netherlands over an 80 yr period. We focus on bees and butterflies, two groups for which we have good knowledge of larval resources that allowed us to define groups of species with different nitrogen related diet preferences. For each group we estimated richness changes between different 20‐yr periods at local, regional and national scale, using analytical methods developed for analyzing richness changes based on collection data.Our findings suggest that the impacts of soil eutrophication on plant communities propagate to higher trophic levels, but with a time‐lag. Pollinators with nitrogen‐related diet preferences were particularly affected, in turn potentially impairing the performance of pollinator‐dependent plants. Pollinator declines continued even after their focal plants started to recover. In addition, our results suggest that current levels of nitrogen deposition still have a negative impact on most groups here analyzed, constraining richness recoveries and accentuating declines.Our results indicate that the global increase in nitrogen availability plays an important role in the ongoing pollinator decline. Consequently, species tolerances to soil nitrogen levels should be considered across all trophic levels in management plans that aim to halt bio

The study of ecosystem processes over multiple scales of space and time is often best achieved using comparable data from multiple sites. Yet, long-term ecological observatories have often developed their own data collection protocols. Here, we address this problem by proposing a set of ecological protocols suitable for widespread adoption by the ecological community. Scientists from the European ecological research community prioritized terrestrial ecosystem parameters that could benefit from a more consistent approach to data collection within the resources available at most long-term ecological observatories. Parameters for which standard methods are in widespread use, or for which methods are evolving rapidly, were not selected. Protocols were developed by domain experts, building on existing methods where possible, and refined through a process of field testing and training. They address above-ground plant biomass; decomposition; land use and management; leaf area index; soil mesofaunal diversity; soil C and N stocks, and greenhouse gas emissions from soils. These complement existing methods to provide a complete assessment of ecological integrity. These protocols offer integrated approaches to ecological data collection that are low cost and are starting to be used across the European Long Term Ecological Research community.

Nature conservation policies need to deliver on multiple criteria, including genetic diversity, population viability and species richness as well as ecosystem services. The challenge of integrating these may be addressed by simulation modelling.We used four models (MetaConnect, SPOMSIM, a community model and InVEST) to assess a variety of spatial habitat patterns with two levels of total habitat cover and realised at two spatial scales, exploring which landscape structures performed best according to five different conservation criteria assessed for four functional types of organisms (approximately representing trees, butterflies, small mammals and birds).The results display both synergies and trade-offs: population size and pollination services generally benefitted more from fragmentation than did genetic heterozygosity, and species richness more than allelic richness, although the latter two varied considerably among the functional types.No single landscape performed best across all conservation criteria, but averaging over criteria and functional types, overall performance improved with greater levels of habitat cover and intermediate fragmentation (or less fragmentation in cases with lower habitat cover).Policy implications. Using four simulation models, we show that different conservation objectives must be traded off in spatial conservation planning, and that considering only a single taxon or criterion may result in suboptimal choices when planning reserve networks. Nevertheless, heterogeneous spatial patterns of habitat can provide reasonable compromises for multiple criteria.

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Fundamental ecological research is both intrinsically interesting and provides the basic knowledge required to answer applied questions of importance to the management of the natural world. The 100th anniversary of the British Ecological Society in 2013 is an opportune moment to reflect on the current status of ecology as a science and look forward to high-light priorities for future work.To do this, we identified 100 important questions of fundamental importance in pure ecology. We elicited questions from ecologists working across a wide range of systems and disciplines. The 754 questions submitted (listed in the online appendix) from 388 participants were narrowed down to the final 100 through a process of discussion, rewording and repeated rounds of voting. This was done during a two-day workshop and thereafter.The questions reflect many of the important current conceptual and technical pre-occupations of ecology. For example, many questions concerned the dynamics of environmental change and complex ecosystem interactions, as well as the interaction between ecology and evolution.The questions reveal a dynamic science with novel subfields emerging. For example, a group of questions was dedicated to disease and micro-organisms and another on human impacts and global change reflecting the emergence of new subdisciplines that would not have been foreseen a few decades ago.The list also contained a number of questions that have perplexed ecologists for decades and are still seen as crucial to answer, such as the link between population dynamics and life-history evolution.Synthesis. These 100 questions identified reflect the state of ecology today. Using them as an agenda for further research would lead to a substantial enhancement in understanding of the discipline, with practical relevance for the conservation of biodiversity and ecosystem function.

Land-use changes can alter the spatial population structure of plant species, which may in turn affect the attractiveness of flower aggregations to different groups of pollinators at different spatial scales. To assess how pollinators respond to spatial heterogeneity of plant distributions and whether honeybees affect visitation by other pollinators we used an extensive data set comprising ten plant species and their flower visitors from five European countries. In particular we tested the hypothesis that the composition of the flower visitor community in terms of visitation frequencies by different pollinator groups were affected by the spatial plant population structure, viz. area and density measures, at a within-population (‘patch’) and among-population (‘population’) scale. We found that patch area and population density were the spatial variables that best explained the variation in visitation frequencies within the pollinator community. Honeybees had higher visitation frequencies in larger patches, while bumblebees and hoverflies had higher visitation frequencies in sparser populations. Solitary bees had higher visitation frequencies in sparser populations and smaller patches. We also tested the hypothesis that honeybees affect the composition of the pollinator community by altering the visitation frequencies of other groups of pollinators. There was a positive relationship between visitation frequencies of honeybees and bumblebees, while the relationship with hoverflies and solitary bees varied (positive, negative and no relationship) depending on the plant species under study. The overall conclusion is that the spatial structure of plant populations affects different groups of pollinators in contrasting ways at both the local (‘patch’) and the larger (‘population’) scales and, that honeybees affect the flower visitation by other pollinator groups in various ways, depending on th

The decline of bees has raised concerns regarding their conservation and the maintenance of ecosystem services they provide to bee-pollinated wild flowers and crops. Although the Mediterranean region is a hotspot for bee species richness, their status remains poorly studied. There is an urgent need for cost-effective, reliable, and unbiased sampling methods that give good bee species richness estimates. This study aims: (a) to assess bee species richness in two common Mediterranean habitat types: semi-natural scrub (phrygana) and managed olive groves; (b) to compare species richness in those systems to that of other biogeographic regions, and (c) to assess whether six different sampling methods (pan traps, variable and standardized transect walks, observation plots and trap nests), previously tested in other European biogeographic regions, are suitable in Mediterranean communities. Eight study sites, four per habitat type, were selected on the island of Lesvos, Greece. The species richness observed was high compared to other habitat types worldwide for which comparable data exist. Pan traps collected the highest proportion of the total bee species richness across all methods at the scale of a study site. Variable and standardized transect walks detected the highest total richness over all eight study sites. Trap nests and observation plots detected only a limited fraction of the bee species richness. To assess the total bee species richness in bee diversity hotspots, such as the studied habitats, we suggest a combination of transect walks conducted by trained bee collectors and pan trap sampling.

Aim The biogeography of arbuscular mycorrhizal (AM) fungi is poorly understood, and consequently the potential of AM fungi to determine plant distribution has been largely overlooked. We aimed to describe AM fungal communities associating with a single host-plant species across a wide geographical area, including the plant’s native, invasive and experimentally introduced ranges. We hypothesized that an alien AM plant associates primarily with the geographically widespread generalist AM fungal taxa present in a novel range. Location Europe, China. Methods We transplanted the palm Trachycarpus fortunei into nine European sites where it does not occur as a native species, into one site where it is naturalized (Switzerland), and into one glasshouse site. We harvested plant roots after two seasons. In addition, we sampled palms at three sites in the plant’s native range (China). Roots were subjected to DNA extraction, polymerase chain reaction (PCR) and 454 sequencing of AM fungal sequences. We analysed fungal communities with non-metric multidimensional scaling (NMDS) ordination and cluster analysis and studied the frequency of geographically widespread fungal taxa with log-linear analysis. We compared fungal communities in the roots of the palm with those in resident plants at one site in the introduced range (Estonia) where natural AM fungal communities had previously been studied. Results We recorded a total of 73 AM fungal taxa. AM fungal communities in the native and introduced ranges differed from one another, while those in the invasive range contained taxa present in both other ranges. Geographically widespread AM fungal taxa were over-represented in palm roots in all regions, but especially in the introduced range. At the Estonian site, the palm was colonized by the same community of widespread AM fungal taxa as associate with resident habitat-generalist plants; by contrast, resident forest-specialist plants were colonized by a diverse community of widespre

Pollinating insects form a key component of European biodiversity, and provide a vital ecosystem service to crops and wild plants. There is growing evidence of declines in both wild and domesticated pollinators, and parallel declines in plants relying upon them. The STEP project (Status and Trends of European Pollinators, 2010-2015, www.stepproject.net) is documenting critical elements in the nature and extent of these declines, examining key functional traits associated with pollination deficits, and developing a Red List for some European pollinator groups. Together these activities are laying the groundwork for future pollinator monitoring programmes. STEP is also assessing the relative importance of potential drivers of pollinator declines, including climate change, habitat loss and fragmentation, agrochemicals, pathogens, alien species, light pollution, and their interactions. We are measuring the ecological and economic impacts of declining pollinator services and floral resources, including effects on wild plant populations, crop production and human nutrition. STEP is reviewing existing and potential mitigation options, and providing novel tests of their effectiveness across Europe. Our work is building upon existing and newly developed datasets and models, complemented by spatially-replicated campaigns of field research to fill gaps in current knowledge. Findings are being integrated into a policy-relevant framework to create evidence-based decision support tools. STEP is establishing communication links to a wide range of stakeholders across Europe and beyond, including policy makers, beekeepers, farmers, academics and the general public. Taken together, the STEP research programme aims to improve our understanding of the nature, causes, consequences and potential mitigation of declines in pollination services at local, national, continental and global scales.

Pollination mode is an important reproductive characteristic, often assumed to play a considerable role in plant species invasiveness. We asked (1) whether alien and native species differed in the frequency of pollination modes (insect pollination, self-pollination, wind pollination, water pollination), (2) whether the pollination modes affected the invasion success of two groups of aliens, differing in their residence time in Central Europe: archaeophytes (introduced before 1500) and neophytes (introduced more recently), and (3) whether there were differences in the diversity of insect pollinators of native species, and of alien species at different stages of invasion and with different residence time. The analysis was carried out using 2817 species occurring in the Czech Republic (1596 native and 1221 alien, the latter comprising 331 archaeophytes and 890 neophytes). Data were analyzed using generalized linear models. The alien flora introduced to Central Europe contained a higher proportion of insect-pollinated species than did the Central European native flora and linked to a higher diversity of pollinators per species. However, the frequency of pollination modes in the introduced alien flora gradually changed during the process of naturalization, becoming more similar to that of native species, and eventually, the naturalized species that became invasive did not differ in their frequency of pollination modes from native species. The frequency of self-pollination increased from casual through naturalized to invasive alien species. This suggests a remarkable role for pollination mode in successful invasions; indeed, self-pollination tends to support spread of neophytes more than any other mode of pollination. The range of habitats occupied by plants of different invasion status affected the diversity of insect pollinator species. In contrast, regional commonness of plant species only affected the number of pollinator functional groups. In native species and archa

Pollinators are a key component of global biodiversity, providing vital ecosystem services to crops and wild plants. There is clear evidence of recent declines in both wild and domesticated pollinators, and parallel declines in the plants that rely upon them. Here we describe the nature and extent of reported declines, and review the potential drivers of pollinator loss, including habitat loss and fragmentation, agrochemicals, pathogens, alien species, climate change and the interactions between them. Pollinator declines can result in loss of pollination services which have important negative ecological and economic impacts that could significantly affect the maintenance of wild plant diversity, wider ecosystem stability, crop production, food security and human welfare.

1. Habitat fragmentation can affect pollinator and plant population structure in terms of species composition, abundance, area covered and density of flowering plants. This, in turn, may affect pollinator visitation frequency, pollen deposition, seed set and plant fitness.2. A reduction in the quantity of flower visits can be coupled with a reduction in the quality of pollination service and hence the plants' overall reproductive success and long-term survival. Understanding the relationship between plant population size and/or isolation and pollination limitation is of fundamental importance for plant conservation.3. We examined flower visitation and seed set of 10 different plant species from five European countries to investigate the general effects of plant populations size and density, both within (patch level) and between populations (population level), on seed set and pollination limitation.4. We found evidence that the effects of area and density of flowering plant assemblages were generally more pronounced at the patch level than at the population level. We also found that patch and population level together influenced flower visitation and seed set, and the latter increased with increasing patch area and density, but this effect was only apparent in small populations.5. Synthesis. By using an extensive pan-European data set on flower visitation and seed set we have identified a general pattern in the interplay between the attractiveness of flowering plant patches for pollinators and density dependence of flower visitation, and also a strong plant species-specific response to habitat fragmentation effects. This can guide efforts to conserve plant-pollinator interactions, ecosystem functioning and plant fitness in fragmented habitats.

The field site network (FSN) plays a central role in conducting joint research within all Assessing Large-scale Risks for biodiversity with tested Methods (ALARM) modules and provides a mechanism for integrating research on different topics in ALARM on the same site for measuring multiple impacts on biodiversity. The network covers most European climates and biogeographic regions, from Mediterranean through central European and boreal to subarctic. The project links databases with the European-wide field site network FSN, including geographic information system (GIS)-based information to characterise the test location for ALARM researchers for joint on-site research. Maps are provided in a standardised way and merged with other site-specific information. The application of GIS for these field sites and the information management promotes the use of the FSN for research and to disseminate the results. We conclude that ALARM FSN sites together with other research sites in Europe jointly could be used as a future backbone for research proposals.

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Bee pollinators are currently recorded with many different sampling methods. However, the relative performances of these methods have not been systematically evaluated and compared. In response to the strong need to record ongoing shifts in pollinator diversity and abundance, global and regional pollinator initiatives must adopt standardized sampling protocols when developing large-scale and long-term monitoring schemes.We systematically evaluated the performance of six sampling methods (observation plots, pan traps, standardized and variable transect walks, trap nests with reed internodes or paper tubes) that are commonly used across a wide range of geographical regions in Europe and in two habitat types (agricultural and seminatural). We focused on bees since they represent the most important pollinator group worldwide. Several characteristics of the methods were considered in order to evaluate their performance in assessing bee diversity: sample coverage, observed species richness, species richness estimators, collector biases (identified by subunit-based rarefaction curves), species composition of the samples, and the indication of overall bee species richness (estimated from combined total samples).The most efficient method in all geographical regions, in both the agricultural and seminatural habitats, was the pan trap method. It had the highest sample coverage, collected the highest number of species, showed negligible collector bias, detected similar species as the transect methods, and was the best indicator of overall bee species richness. The transect methods were also relatively efficient, but they had a significant collector bias. The observation plots showed poor performance. As trap nests are restricted to cavity-nesting bee species, they had a naturally low sample coverage. However, both trap nest types detected additional species that were not recorded by any of the other methods.For large-scale and long-term monitoring schemes with surveyors with di

Climate change, environmental chemicals, biological invasions, and loss of pollinator are the main research topics in the ALARM project. All of theses environmental pressures are more or less of anthropogenic origin. These have generally been studied independently of one another. But it is clear that they interact, potentially producing effects on biodiversity and ecosystems that exceed all current assessments of potential risks. ALARM develops relevant assessment methods in two phases: a) concept and data base integration and b) ecosystem risk evaluation. These methods will be tested and standardised protocols will be developed.The project links databases with a European-wide field site network and then generates common methods. At these field sites GIS is used as a basis to characterise the test location for ALARM researchers for joint on-site research. Maps are provided in a standardized way and merged with other site specific information. The application of GIS for these field sites and the information management is necessary to communicate the research and the results. Problems of data homogenisation and of unique quality standards of databases arise, but ways to solve this will be shown. GIS as a basis of risk maps on a European level and support the scientists to provide their results.

Experimental studies have shown that many species show preferences for different climatic conditions, or may die in unsuitable conditions. Climate envelope models have been used frequently in recent years to predict the presence and absence of species at large spatial scales. However, many authors have postulated that the distributions of species at smaller spatial scales are determined by factors such as habitat availability and biotic interactions. Climatic effects are often assumed by modellers to be unimportant at fine resolutions, but few studies have actually tested this. We sampled the distributions of 20 beetle species of the family Carabidae across three study sites by pitfall trapping, and at the national scale from monitoring data. Statistical models were constructed to determine which of two sets of environmental variables (temperature or broad habitat type) best accounted for the observed data at the three sites and at the national (Great Britain) scale. High-resolution temperature variables frequently produced better models (as determined by AIC) than habitat features when modelling the distributions of species at a local scale, within the three study sites. Conversely, habitat was always a better predictor than temperature when describing species’ distributions at a coarse scale within Great Britain. Northerly species were most likely to occur in cool micro-sites within the study sites, whereas southerly species were most likely to occur in warm micro-sites. Effects of microclimate were not limited to species at the edges of their distribution, and fine-resolution temperature surfaces should therefore ideally be utilised when undertaking climate-envelope modelling.