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Recent evidence highlights the value of wild-insect species richness and abundance for crop pollination worldwide. Yet, deliberate physical importation of single species (eg European honey bees) into crop fields for pollination remains the mainstream management approach, and implementation of practices to enhance crop yield (production per area) through wild insects is only just beginning. With few exceptions, studies measuring the impacts of pollinator-supporting practices on wild-insect richness and pollination service success - particularly in relation to long-term crop yield and economic profit - Are rare. Here, we provide a general framework and examples of approaches for enhancing pollinator richness and abundance, quantity and quality of pollen on stigmas, crop yield, and farmers' profit, including some benefits detected only through long-term monitoring. We argue for integrating the promotion of wild-insect species richness with single-species management to benefit farmers and society.

Substantial evidence suggests that biodiversity can stabilize ecosystem function, but how it does this is less clear. In very general terms, the hypothesis is that biodiversity stabilizes function because having more species increases the role of compensatory dynamics, which occur when species in a community show different responses to the environment. Here, we focus on two forms of compensatory dynamics, cross-scale redundancy (CSR) and response diversity (RD). CSR occurs when species respond to a disturbance at different scales such that scale-specific disturbances do not negatively affect all species. RD occurs when species contributing to the same function show different responses to an environmental change. We developed a new analytical approach that can compare the strength of CSR and RD in the same dataset and used it to study native bee pollination of blueberry at 16 farms that varied in surrounding agricultural land use. We then asked whether CSR and RD among bee species are associated with the stability of blueberry pollination. Although CSR and RD were both present, only RD was associated with higher stability of pollination. Furthermore, the effects of RD on stability were due to a single widespread species, Andrena bradleyi, that is a specialist on blueberry and, unlike other bee species, was highly abundant at farms surrounded by intensive blueberry agriculture. Thus, the stabilizing effect we observed was attributable to an "identity effect" more than to species richness per se. Our results demonstrate how CSR and RD can be empirically measured and compared and highlight how the theoretical expectations of the biodiversity-ecosystem functioning field are not always upheld when confronted with real-world data., (© 2024 The Ecological Society of America.)

Declining pollinator populations could threaten global food production, especially if current crop yields are limited by insufficient pollinator visitation to flowers, in a phenomenon referred to as 'pollinator limitation'. Here, we assess the global prevalence of pollinator limitation, explore the risk factors, such as crop type or geographic region, that predict where pollinator limitation is more likely and ask by how much increases in pollinator visitation could improve crop yields. We address these questions using 198,360 plant-pollinator interactions and 2,083 yield measurements from 32 crop species grown in 120 study systems. We find that 28-61% of global crop systems are pollinator limited and that this limitation most frequently occurs in blueberry, coffee and apple crops. For a few datasets, we note that the probability of pollinator limitation decreases with greater forest land cover surrounding a crop field at 1 km, although average effect sizes are small. Finally, we estimate that for those crops we identify as pollinator limited, increasing pollinator visitation at all farms to existing levels observed in the 90th percentile of each study system would close 63% of yield gaps between high- and low-yielding fields. Our findings show variations in sensitivity to pollinator limitation across diverse crop systems and indicate that realistic increases in pollinator visitation could mitigate crop yield shortfalls attributable to pollinator limitation., (© 2024. The Author(s), under exclusive licence to Springer Nature Limited.)

Plant-pollinator interactions are ecologically and economically important, and, as a result, their prediction is a crucial theoretical and applied goal for ecologists. Although various analytical methods are available, we still have a limited ability to predict plant-pollinator interactions. The predictive ability of different plant-pollinator interaction models depends on the specific definitions used to conceptualize and quantify species attributes (e.g., morphological traits), sampling effects (e.g., detection probabilities), and data resolution and availability. Progress in the study of plant-pollinator interactions requires conceptual and methodological advances concerning the mechanisms and species attributes governing interactions as well as improved modeling approaches to predict interactions. Current methods to predict plant-pollinator interactions present ample opportunities for improvement and spark new horizons for basic and applied research., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

Premise: Bees provision most of the pollen removed from anthers to their larvae and transport only a small proportion to stigmas, which can negatively affect plant fitness. Though most bee species collect pollen from multiple plant species, we know little about how the efficiency of bees' pollen transport varies among host plant species or how it relates to other aspects of generalist bee foraging behavior that benefit plant fitness, such as specialization on individual foraging bouts., Methods: We compared the pollen collected and transported by three bee species for 46 co-occurring plant species. Specifically, we compared the relative abundance of pollen taxa in the individual bees' scopae, structures where bees store pollen to provision larvae, with the relative abundance of pollen taxa on the rest of bees' bodies, which is more likely to be transferred to stigmas., Results: Bees carried five times more pollen grains in their scopae than elsewhere on their bodies. Within foraging bouts, bees were relatively specialized in their pollen collection, but transported proportionally less pollen for the host plants on which they specialized. Across foraging bouts, two bee species transported proportionally less pollen for some of their host plants than for others, though differences didn't consistently follow the same trend as at the foraging bout scale., Conclusions: Our results suggest that foraging-bout specialization, which is known to reduce heterospecific pollen transfer, also results in less-efficient pollen transport. Thus, bee foragers that visit predominantly one plant species may have contrasting effects on that plant's fitness., (© 2023 The Authors. American Journal of Botany published by Wiley Periodicals LLC on behalf of Botanical Society of America.)

Biodiversity promotes ecosystem function (EF) in experiments, but it remains uncertain how biodiversity loss affects function in larger-scale natural ecosystems. In these natural ecosystems, rare and declining species are more likely to be lost, and function needs to be maintained across space and time. Here, we explore the importance of rare and declining bee species to the pollination of three wildflowers and three crops using large-scale (72 sites across 5000 km 2 ), multi-year datasets. Half of the sampled bee species (82/164) were rare or declining, but these species provided only ~15% of overall pollination. To determine the number of species important to EF, we used two methods of "scaling up," both of which have previously been used for biodiversity-function analysis. First, we summed bee species' contributions to pollination across space and time and then found the minimum set of species needed to provide a threshold level of function across all sites; according to this method, effectively no rare and declining bee species were important to pollination. Second, we account for the "insurance value" of biodiversity by finding the minimum set of bee species needed to simultaneously provide a threshold level of function at each site in each year. The second method leads to the conclusion that 25 rare and eight declining bee species (36% and 53% of all rare and declining bee species, respectively) are included in the minimum set. Our findings provide some of the strongest evidence yet that rare and declining species are key to meeting threshold levels of EF, thereby providing a more direct link between real-world biodiversity loss and EF., (© 2022 The Ecological Society of America.)

The current biodiversity crisis underscores the need to understand how biodiversity loss affects ecosystem function in real-world ecosystems. At any one place and time, a few highly abundant species often provide the majority of function, suggesting that function could be maintained with relatively little biodiversity. However, biodiversity may be critical to ecosystem function at longer timescales if different species are needed to provide function at different times. Here we show that the number of wild bee species needed to maintain a threshold level of crop pollination increased steeply with the timescale examined: two to three times as many bee species were needed over a growing season compared to on a single day and twice as many species were needed over six years compared to during a single year. Our results demonstrate the importance of pollinator biodiversity to maintaining pollination services across time and thus to stable agricultural output., (© 2022. The Author(s), under exclusive licence to Springer Nature Limited.)

AbstractThe relationship between biodiversity and ecosystem function (BEF) remains unclear in many natural ecosystems, partially for lack of theoretical and analytical tools that match common characteristics of observational community data. The ecological Price equation promises to meet this need by organizing many different species-level changes into a few ecologically meaningful categories that sum to total ecosystem function change. Current versions of the ecological Price equation focus on species richness and presence-absence. However, abundance and relative abundance are better estimated in samples and are likely showing a stronger response to global change. Here, we present a novel, abundance-based version of the ecological Price equation in both discrete and continuous forms and explain the similarities and differences between this method and a related, previously developed richness-based method. We also present new empirical techniques for applying the Price equation to ecological data. Our two demonstration analyses reveal how additive effects of increasing abundance on total function are modified by concurrent selection effects due to shifts in species' composition as well as intraspecific change in species' per capita function. The ecological Price equations derived here complement existing approaches and together offer BEF researchers analytical tools and a unifying framework for studying BEF in observational community data.

It is important to understand how biodiversity, including that of rare species, affects ecosystem function. Here, we consider this question with regard to pollination. Studies of pollination function have typically focused on pollination of single plant species, or average pollination across plants, and typically find that pollination depends on a few common species. Here, we used data from 11 plant-bee visitation networks in New Jersey, USA, to ask whether the number of functionally important bee species changes as we consider function separately for each plant species in increasingly diverse plant communities. Using rarefaction analysis, we found the number of important bee species increased with the number of plant species. Overall, 2.5 to 7.6 times more bee species were important at the community scale, relative to the average plant species in the same community. This effect did not asymptote in any of our datasets, suggesting that even greater bee biodiversity is needed in real-world systems. Lastly, on average across plant communities, 25% of bee species that were important at the community scale were also numerically rare within their network, making this study one of the strongest empirical demonstrations to date of the functional importance of rare species.

Seventy five percent of the world's food crops benefit from insect pollination. Hence, there has been increased interest in how global change drivers impact this critical ecosystem service. Because standardized data on crop pollination are rarely available, we are limited in our capacity to understand the variation in pollination benefits to crop yield, as well as to anticipate changes in this service, develop predictions, and inform management actions. Here, we present CropPol, a dynamic, open, and global database on crop pollination. It contains measurements recorded from 202 crop studies, covering 3,394 field observations, 2,552 yield measurements (i.e., berry mass, number of fruits, and fruit density [kg/ha], among others), and 47,752 insect records from 48 commercial crops distributed around the globe. CropPol comprises 32 of the 87 leading global crops and commodities that are pollinator dependent. Malus domestica is the most represented crop (32 studies), followed by Brassica napus (22 studies), Vaccinium corymbosum (13 studies), and Citrullus lanatus (12 studies). The most abundant pollinator guilds recorded are honey bees (34.22% counts), bumblebees (19.19%), flies other than Syrphidae and Bombyliidae (13.18%), other wild bees (13.13%), beetles (10.97%), Syrphidae (4.87%), and Bombyliidae (0.05%). Locations comprise 34 countries distributed among Europe (76 studies), North America (60), Latin America and the Caribbean (29), Asia (20), Oceania (10), and Africa (7). Sampling spans three decades and is concentrated on 2001-2005 (21 studies), 2006-2010 (40), 2011-2015 (88), and 2016-2020 (50). This is the most comprehensive open global data set on measurements of crop flower visitors, crop pollinators and pollination to date, and we encourage researchers to add more datasets to this database in the future. This data set is released for non-commercial use only. Credits should be given to this paper (i.e., proper citation), and the products generated with this database should be shared under the same license terms (CC BY-NC-SA)., (© 2021 The Ecological Society of America.)

Floral constancy of foraging bees influences plant reproduction. Constancy as observed in nature arises from at least four distinct mechanisms frequently confounded in the literature: context-independent preferences for particular plant species, preferential visitation to the same species as the previous plant visited (simple constancy), the spatial arrangement of plants, and the relative abundances of co-flowering species. To disentangle these mechanisms, we followed individual bee flight paths within patches where all flowering plants were mapped, and we used step selection models to estimate how each mechanism influences the probability of selecting any particular plant given the available plants in a multi-species community. We found that simple constancy was positive: bees preferred to visit the same species sequentially. In addition, bees preferred to travel short distances and maintain their direction of travel between plants. After accounting for distance, we found no significant effect of site-level plant relative abundances on bee foraging choices. To explore the importance of the spatial arrangement of plants for bee foraging choices, we compared our full model containing all parameters to one with spatial arrangement removed. Due to bees' tendency to select nearby plants, combined with strong intraspecific plant clumping, spatial arrangement was responsible for about 50% of the total observed constancy. Our results suggest that floral constancy may be overestimated in studies that do not account for the spatial arrangement of plants, especially in systems with intraspecific plant clumping. Plant spatial patterns at within-site scales are important for pollinator foraging behavior and pollination success., (© 2022. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

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.

Most studies of plant-animal mutualistic networks have come from a temporally static perspective. This approach has revealed general patterns in network structure, but limits our ability to understand the ecological and evolutionary processes that shape these networks and to predict the consequences of natural and human-driven disturbance on species interactions. We review the growing literature on temporal dynamics of plant-animal mutualistic networks including pollination, seed dispersal and ant defence mutualisms. We then discuss potential mechanisms underlying such variation in interactions, ranging from behavioural and physiological processes at the finest temporal scales to ecological and evolutionary processes at the broadest. We find that at the finest temporal scales (days, weeks, months) mutualistic interactions are highly dynamic, with considerable variation in network structure. At intermediate scales (years, decades), networks still exhibit high levels of temporal variation, but such variation appears to influence network properties only weakly. At the broadest temporal scales (many decades, centuries and beyond), continued shifts in interactions appear to reshape network structure, leading to dramatic community changes, including loss of species and function. Our review highlights the importance of considering the temporal dimension for understanding the ecology and evolution of complex webs of mutualistic interactions., (© 2020 The Authors. Ecology Letters published by John Wiley & Sons Ltd.)

Individual pollinators that specialize on one plant species within a foraging bout transfer more conspecific and less heterospecific pollen, positively affecting plant reproduction. However, we know much less about pollinator specialization at the scale of a foraging bout compared to specialization by pollinator species. In this study, we measured the diversity of pollen carried by individual bees foraging in forest plant communities in the mid-Atlantic United States. We found that individuals frequently carried low-diversity pollen loads, suggesting that specialization at the scale of the foraging bout is common. Individuals of solitary bee species carried higher diversity pollen loads than did individuals of social bee species; the latter have been better studied with respect to foraging bout specialization, but account for a small minority of the world's bee species. Bee body size was positively correlated with pollen load diversity, and individuals of polylectic (but not oligolectic) species carried increasingly diverse pollen loads as the season progressed, likely reflecting an increase in the diversity of flowers in bloom. Furthermore, the seasonal increase in pollen load diversity was stronger for bees visiting trees and shrubs than for bees visiting herbaceous plants. Overall, our results showed that both plant and pollinator species' traits as well as community-level patterns of flowering phenology are likely to be important determinants of individual-level interactions in plant-pollinator communities., (© 2019 The Authors. Journal of Animal Ecology © 2019 British Ecological Society.)

Background: Intraspecific variation in foraging niche can drive food web dynamics and ecosystem processes. In particular, male and female animals can exhibit different, often cascading, impacts on their interaction partners. Despite this, studies of plant-pollinator interaction networks have focused on the partitioning of the floral community between pollinator species, with little attention paid to intraspecific variation in plant preference between male and female bees. We designed a field study to evaluate the strength and prevalence of sexually dimorphic foraging, and particularly resource preferences, in bees., Study Design: We observed bees visiting flowers in semi-natural meadows in New Jersey, USA. To detect differences in flower use against a shared background of resource (flower) availability, we maximized the number of interactions observed within narrow spatio-temporal windows. To distinguish observed differences in bee use of flower species, which can reflect abundance patterns and sampling effects, from underlying differences in bee preferences, we analyzed our data with both a permutation-based null model and random effects models., Findings: We found that the diets of male and female bees of the same species were often dissimilar as the diets of different species of bees. Furthermore, we demonstrate differences in preference between male and female bees. We show that intraspecific differences in preference can be robustly identified among hundreds of unique species-species interactions, without precisely quantifying resource availability, and despite high phenological turnover of both bees and plant bloom. Given the large differences in both flower use and preferences between male and female bees, ecological sex differences should be integrated into studies of bee demography, plant pollination, and coevolutionary relationships between flowers and insects., Competing Interests: The authors have declared that no competing interests exist.

[This corrects the article DOI: 10.1371/journal.pone.0151482.].

Ecologists have shown through hundreds of experiments that ecological communities with more species produce higher levels of essential ecosystem functions such as biomass production, nutrient cycling, and pollination, but whether this finding holds in nature (that is, in large-scale and unmanipulated systems) is controversial. This knowledge gap is troubling because ecosystem services have been widely adopted as a justification for global biodiversity conservation. Here we show that, to provide crop pollination in natural systems, the number of bee species must increase by at least one order of magnitude compared with that in field experiments. This increase is driven by species turnover and its interaction with functional dominance, mechanisms that emerge only at large scales. Our results show that maintaining ecosystem services in nature requires many species, including relatively rare ones., (Copyright © 2018 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works.)

Anthropogenic landscapes are associated with biodiversity loss and large shifts in species composition and traits. These changes predict the identities of winners and losers of future global change, and also reveal which environmental variables drive a taxon's response to land use change. We explored how the biodiversity of native bee species changes across forested, agricultural, and urban landscapes. We collected bee community data from 36 sites across a 75,000 km 2 region, and analyzed bee abundance, species richness, composition, and life-history traits. Season-long bee abundance and richness were not detectably different between natural and anthropogenic landscapes, but community phenologies differed strongly, with an early spring peak followed by decline in forests, and a more extended summer season in agricultural and urban habitats. Bee community composition differed significantly between all three land use types, as did phylogenetic composition. Anthropogenic land use had negative effects on the persistence of several life-history strategies, including early spring flight season and brood parasitism, which may indicate adaptation to conditions in forest habitat. Overall, anthropogenic communities are not diminished subsets of contemporary natural communities. Rather, forest species do not persist in anthropogenic habitats, but are replaced by different native species and phylogenetic lineages preadapted to open habitats. Characterizing compositional and functional differences is crucial for understanding land use as a global change driver across large regional scales., (© 2017 John Wiley & Sons Ltd.)

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., (© 2017 John Wiley & Sons Ltd.)

The relationship between biodiversity and the stability of ecosystem function is a fundamental question in community ecology, and hundreds of experiments have shown a positive relationship between species richness and the stability of ecosystem function. However, these experiments have rarely accounted for common ecological patterns, most notably skewed species abundance distributions and non-random extinction risks, making it difficult to know whether experimental results can be scaled up to larger, less manipulated systems. In contrast with the prolific body of experimental research, few studies have examined how species richness affects the stability of ecosystem services at more realistic, landscape scales. The paucity of these studies is due in part to a lack of analytical methods that are suitable for the correlative structure of ecological data. A recently developed method, based on the Price equation from evolutionary biology, helps resolve this knowledge gap by partitioning the effect of biodiversity into three components: richness, composition, and abundance. Here, we build on previous work and present the first derivation of the Price equation suitable for analyzing temporal variance of ecosystem services. We applied our new derivation to understand the temporal variance of crop pollination services in two study systems (watermelon and blueberry) in the mid-Atlantic United States. In both systems, but especially in the watermelon system, the stronger driver of temporal variance of ecosystem services was fluctuations in the abundance of common bee species, which were present at nearly all sites regardless of species richness. In contrast, temporal variance of ecosystem services was less affected by differences in species richness, because lost and gained species were rare. Thus, the findings from our more realistic landscapes differ qualitatively from the findings of biodiversity-stability experiments., (© 2017 by the Ecological Society of America.)

Recent studies of mutualistic networks show that interactions between partners change across years. Both biological mechanisms and chance could drive these patterns, but the relative importance of these factors has not been separated. We established a field experiment consisting of 102 monospecific plots of 17 native plant species, from which we collected 6713 specimens of 52 bee species over four years. We used these data and a null model to determine whether bee species' foraging choices varied more or less over time beyond the variation expected by chance. Thus we provide the first quantitative definition of rewiring and fidelity as these terms are used in the literature on interaction networks. All 52 bee species varied in plant partner choice across years, but for 27 species this variation was indistinguishable from random partner choice. Another 11 species showed rewiring, varying more across years than expected by chance, while 14 species showed fidelity, indicating that they both prefer certain plant species and are consistent in those preferences across years. Our study shows that rewiring and fidelity both exist in mutualist networks, but that once sampling effects have been accounted for, they are less common than has been reported in the ecological literature., (© 2016 by the Ecological Society of America.)

Allometric relationships among morphological traits underlie important patterns in ecology. These relationships are often phylogenetically shared; thus quantifying allometric relationships may allow for estimating difficult-to-measure traits across species. One such trait, proboscis length in bees, is assumed to be important in structuring bee communities and plant-pollinator networks. However, it is difficult to measure and thus rarely included in ecological analyses. We measured intertegular distance (as a measure of body size) and proboscis length (glossa and prementum, both individually and combined) of 786 individual bees of 100 species across 5 of the 7 extant bee families (Hymenoptera: Apoidea: Anthophila). Using linear models and model selection, we determined which parameters provided the best estimate of proboscis length. We then used coefficients to estimate the relationship between intertegular distance and proboscis length, while also considering family. Using allometric equations with an estimation for a scaling coefficient between intertegular distance and proboscis length and coefficients for each family, we explain 91% of the variance in species-level means for bee proboscis length among bee species. However, within species, individual-level intertegular distance was a poor predictor of individual proboscis length. To make our findings easy to use, we created an R package that allows estimation of proboscis length for individual bee species by inputting only family and intertegular distance. The R package also calculates foraging distance and body mass based on previously published equations. Thus by considering both taxonomy and intertegular distance we enable accurate estimation of an ecologically and evolutionarily important trait.

Wild and managed bees are well documented as effective pollinators of global crops of economic importance. However, the contributions by pollinators other than bees have been little explored despite their potential to contribute to crop production and stability in the face of environmental change. Non-bee pollinators include flies, beetles, moths, butterflies, wasps, ants, birds, and bats, among others. Here we focus on non-bee insects and synthesize 39 field studies from five continents that directly measured the crop pollination services provided by non-bees, honey bees, and other bees to compare the relative contributions of these taxa. Non-bees performed 25-50% of the total number of flower visits. Although non-bees were less effective pollinators than bees per flower visit, they made more visits; thus these two factors compensated for each other, resulting in pollination services rendered by non-bees that were similar to those provided by bees. In the subset of studies that measured fruit set, fruit set increased with non-bee insect visits independently of bee visitation rates, indicating that non-bee insects provide a unique benefit that is not provided by bees. We also show that non-bee insects are not as reliant as bees on the presence of remnant natural or seminatural habitat in the surrounding landscape. These results strongly suggest that non-bee insect pollinators play a significant role in global crop production and respond differently than bees to landscape structure, probably making their crop pollination services more robust to changes in land use. Non-bee insects provide a valuable service and provide potential insurance against bee population declines.

Anthropogenic change can have large impacts on wild bees and the pollination services they provide. However, the overall pattern of wild bee response to drivers such as land-use change, pesticides, pathogens, and climate change has been one of variability in both the magnitude and directionality of responses. We argue that two causes contribute to this variation. First, different species exhibit differential responses to the same anthropogenic drivers. Second, these anthropogenic drivers vary in type and magnitude that will drive variation in bee responses. For this second issue, we focus on land-use change, the most well-studied driver. We conclude by discussing how understanding species-level responses and the magnitude of land-use change can make bee conservation more effective., (Copyright © 2015 Elsevier Inc. All rights reserved.)

Biodiversity-ecosystem functioning experiments have established that species richness and composition are both important determinants of ecosystem function in an experimental context. Determining whether this result holds for real-world ecosystem services has remained elusive, however, largely due to the lack of analytical methods appropriate for large-scale, associational data. Here, we use a novel analytical approach, the Price equation, to partition the contribution to ecosystem services made by species richness, composition and abundance in four large-scale data sets on crop pollination by native bees. We found that abundance fluctuations of dominant species drove ecosystem service delivery, whereas richness changes were relatively unimportant because they primarily involved rare species that contributed little to function. Thus, the mechanism behind our results was the skewed species-abundance distribution. Our finding that a few common species, not species richness, drive ecosystem service delivery could have broad generality given the ubiquity of skewed species-abundance distributions in nature., (© 2015 John Wiley & Sons Ltd/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.

Modern agriculture relies on domesticated pollinators such as the honey bee (Apis mellifera L.), and to a lesser extent on native pollinators, for the production of animal-pollinated crops. There is growing concern that pollinator availability may not keep pace with increasing agricultural production. However, whether crop production is in fact pollen-limited at the field scale has rarely been studied. Here, we ask whether commercial highbush blueberry (Vaccinium corymbosum L.) production in New Jersey is limited by a lack of pollination even when growers provide honey bees at recommended densities. We studied two varieties of blueberry over 3 yr to determine whether blueberry crop production is pollen-limited and to measure the relative contributions of honey bees and native bees to blueberry pollination. We found two lines of evidence for pollen limitation. First, berries receiving supplemental hand-pollination were generally heavier than berries receiving ambient pollination. Second, mean berry mass increased significantly and nonasymptotically with honey bee flower visitation rate. While honey bees provided 86% of pollination and thus drove the findings reported above, native bees still contributed 14% of total pollination even in our conventionally managed, high-input agricultural system. Honey bees and native bees were also similarly efficient as pollinators on a per-visit basis. Overall, our study shows that pollination can be a limiting factor in commercial fruit production. Yields might increase with increased honey bee stocking rates and improved dispersal of hives within crop fields, and with habitat restoration to increase pollination provided by native bees.

Bacterial gut symbiont communities are critical for the health of many insect species. However, little is known about how microbial communities vary among host species or how they respond to anthropogenic disturbances. Bacterial communities that differ in richness or composition may vary in their ability to provide nutrients or defenses. We used deep sequencing to investigate gut microbiota of three species in the genus Bombus (bumble bees). Bombus are among the most economically and ecologically important non-managed pollinators. Some species have experienced dramatic declines, probably due to pathogens and land-use change. We examined variation within and across bee species and between semi-natural and conventional agricultural habitats. We categorized as 'core bacteria' any operational taxonomic units (OTUs) with closest hits to sequences previously found exclusively or primarily in the guts of honey bees and bumble bees (genera Apis and Bombus). Microbial community composition differed among bee species. Richness, defined as number of bacterial OTUs, was highest for B. bimaculatus and B. impatiens. For B. bimaculatus, this was due to high richness of non-core bacteria. We found little effect of habitat on microbial communities. Richness of non-core bacteria was negatively associated with bacterial abundance in individual bees, possibly due to deeper sampling of non-core bacteria in bees with low populations of core bacteria. Infection by the gut parasite Crithidia was negatively associated with abundance of the core bacterium Gilliamella and positively associated with richness of non-core bacteria. Our results indicate that Bombus species have distinctive gut communities, and community-level variation is associated with pathogen infection.

Theoretical and simulation studies predict that the order of species loss from mutualist networks with respect to how linked species are to other species within the network will determine the rate at which networks collapse. However, the empirical order of species loss with respect to linkage has rarely been investigated. Furthermore, a species' linkage is a composite of its diet breadth and its abundance, yet the relative importance of these two factors in determining species loss order is poorly known. Here we explore the order of pollinator species loss in two contrasting study systems undergoing land-use intensification, using >20,000 pollinator specimens. We found that a pollinator species' linkage, as measured independently within plant-pollinator networks, positively predicted its persistence at human-disturbed sites in three of four analyses. The strongest predictor of persistence in all analyses was pollinator species abundance. In contrast, diet breadth poorly predicted persistence. Overall, our results suggest that community disassembly order buffers plant-pollinator networks against environmental change by retaining the highly linked species that make a disproportionate contribution to network robustness. Furthermore, these highly linked species likely persist because they are also the most common species, not because they are dietary generalists.