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Sunflower ( Helianthus annuus ) pollination in California's Central Valley is limited by native bee nest site location
Ecological Applications
Abstract
The delivery of ecosystem services by mobile organisms depends on the distribution of those organisms, which is, in turn, affected by resources at local and landscape scales. Pollinator-dependent crops rely on mobile animals like bees for crop production, and the spatial relationship between floral resources and nest location for these central-place foragers influences the delivery of pollination services. Current models that map pollination coverage in agricultural regions utilize landscape-level estimates of floral availability and nesting incidence inferred from expert opinion, rather than direct assessments. Foraging distance is often derived from proxies of bee body size, rather than direct measurements of foraging that account for behavioral responses to floral resource type and distribution. The lack of direct measurements of nesting incidence and foraging distances may lead to inaccurate mapping of pollination services. We examined the role of local-scale floral resource presence from hedgerow plantings on nest incidence of ground-nesting bees in field margins and within monoculture, conventionally managed sunflower fields in California's Central Valley. We tracked bee movement into fields using fluorescent powder. We then used these data to simulate the distribution of pollination services within a crop field. Contrary to expert opinion, we found that ground-nesting native bees nested both in fields and edges, though nesting rates declined with distance into field. Further, we detected no effect of field-margin floral enhancements on nesting. We found evidence of an exponential decay rate of bee movement into fields, indicating that foraging predominantly occurred in less than 1% of medium-sized bees' predicted typical foraging range. Although we found native bees nesting within agricultural fields, their restricted foraging movements likely centralize pollination near nest sites. Our data thus predict a heterogeneous distribution of pollination services within sunflower fields, with edges receiving higher coverage than field centers. To generate more accurate maps of services, we advocate directly measuring the autecology of ecosystem service providers, which vary by crop system, pollinator species, and region. Improving estimates of the factors affecting pollinator populations can increase the accuracy of pollination service maps and help clarify the influence of farming practices on wild bees occurring in agricultural landscapes.
As the global agricultural footprint expands, it is increasingly important to address the link between the resource pulses characteristic of monoculture farming and wildlife epidemiology. To understand how mass-flowering crops impact host communities and subsequently amplify or dilute parasitism, we surveyed wild and managed bees in a monoculture landscape with varying degrees of floral diversification. We screened 1509 bees from 16 genera in sunflower fields and in non-crop flowering habitat across 200 km2 of the California Central Valley. We found that mass-flowering crops increase bee abundance. Wild bee abundance was subsequently associated with higher parasite presence, but only in sites with a low abundance of non-crop flowers. Bee traits related to higher dispersal ability (body size) and diet breadth (pollen lecty) were also positively related to parasite presence. Our results highlight the importance of non-crop flowering habitat for supporting bee communities. We suggest monoculture alone cannot support healthy bees.
Wild bees, especially ground-nesting species, are important for crop pollination. However, the extent to which regular disturbance, such as tillage, impacts ground-nesting bees is poorly known. We conducted a two-year tillage experiment on the squash bee, Peponapis pruinosa (Say), to quantify how tillage impacted offspring survival, offspring sex ratio, and timing of emergence. We established P. pruinosa nests in twenty 3 m by 3 m cages and then randomly assigned a tillage treatment to half of those cages. The following summer we trapped emerging offspring. We used a Bayesian framework to analyse the data. Offspring emergence varied greatly, but there was modest evidence that tillage reduced offspring emergence in treatment cages. There was limited evidence that tillage reduced the proportion of male bees emerging into cages, suggesting that tillage did not impact sex ratio. There was strong evidence that tillage delayed emergence of surviving offspring. Tillage practices should be considered when managing fields for this ground nesting bee. However, even disturbed squash fields may contribute to this species’ persistence in agricultural landscapes.
A major challenge in habitat restoration is targeting the key aspects of a species' niche for enhancement, particularly for species that use a diverse set of habitat features. However, restoration that focuses on limited aspects of a species' niche may neglect other resources that are critical to population persistence. We evaluated the ability of native plant hedgerows, planted to increase pollen and nectar resources for wild bees in agricultural landscapes, to provide suitable nesting habitat and enhance nesting rates of ground-nesting bees. We found that, when compared to unmanaged field edges (controls), hedgerows did not augment most indicators of nest habitat quality (bare ground, soil surface irregularity, and soil hardness), although coarser soils were associated with higher incidence and richness of nesting bees. Hedgerows did not augment nesting rates when compared to control edges. Although all the bee species we detected nesting were also found foraging on floral resources, the foraging versus nesting assemblages found within a site were highly dissimilar. These results may reflect sampling error; or, species found foraging but not nesting in hedgerows could be utilizing hedgerows as “partial habitats,” nesting outside hedgerow plantings but foraging on the floral resources they provide. We conclude that although hedgerows are known to provide critical floral resources to wild bees especially in resource-poor intensive agricultural landscapes, simply increasing vegetative diversity and structure may not be simultaneously enhancing nesting habitat for ground-nesting bees.
Research relating to ecosystem services has increased, partly because of drastic declines in biodiversity in agricultural landscapes. However, the mechanistic linkages between land use, biodiversity and service provision are poorly understood and synthesized. This is particularly true for many ecosystem services provided by mobile organisms such as natural enemies to crop pests. These species are not only influenced by local land use but also by landscape composition at larger spatial scales.
We present a conceptual ecological production function framework for predicting land‐use impact on biological control of pests by natural enemies. We develop a novel, mechanistic landscape model for biological control of cereal aphids, explicitly accounting for the influence of landscape composition on natural enemies varying in mobility, feeding rates and other life history traits. Finally, we use the model to map biological control services across cereal fields in a Swedish agricultural region with varying landscape complexity.
The model predicted that biological control would reduce crop damage by 45–70% and that the biological control effect would be higher in complex landscapes. In a validation with independent data, the model performed well and predicted a significant proportion of biological control variation in cereal fields. However, much variability remains to be explained, and we propose that the model could be improved by refining the mechanistic understanding of predator dynamics and accounting for variation in aphid colonization.
We encourage scientists working with biological control to adopt the conceptual framework presented here and to develop production functions for other crop‐pest systems. If this kind of ecological production function is combined with production functions for other services, the joint model will be a powerful tool for managing ecosystem services and planning for sustainable agriculture at the landscape scale.
Natural ecosystems benefit human communities by providing ecosystem services such as water purification and crop pollination. Mapping ecosystem service values has become popular, but most are static snapshots of average value. Estimating instead the economic impacts of specific ecosystem changes can better inform typical resource decisions. Here we develop an approach to mapping marginal values, those resulting from the next unit of ecosystem change, across landscapes. We demonstrate the approach with a recent model of crop pollination services in Costa Rica, simulating deforestation events to predict resulting marginal changes in pollination services to coffee farms. We find that marginal losses from deforestation vary from zero to US$700/ha across the landscape. Financial risks for farmers from these losses and marginal benefits of forest restoration show similar spatial variation. Marginal values are concentrated in relatively few forest parcels not identified using average value. These parcels lack substitutes: nearby forest parcels that can supply services in the event of loss. Indeed, the marginal value of forest parcels declines exponentially with the density of surrounding forest cover. The approach we develop is applicable to any ecosystem service. Combined with information on costs, it can help target conservation or restoration efforts to optimize benefits to people and biodiversity.
Pollinators provide essential ecosystem services, and declines in some pollinator communities around the world have been reported. Understanding the fundamental components defining these communities is essential if conservation and restoration are to be successful. We examined the structure of plant-pollinator communities in a dynamic Mediterranean landscape, comprising a mosaic of post-fire regenerating habitats, and which is a recognized global hotspot for bee diversity. Each community was characterized by a highly skewed species abundance distribution, with a few dominant and many rare bee species, and was consistent with a log series model indicating that a few environmental factors govern the community. Floral community composition, the quantity and quality of forage resources present, and the geographic locality organized bee communities at various levels: (1) The overall structure of the bee community (116 species), as revealed through ordination, was dependent upon nectar resource diversity (defined as the variety of nectar volume-concentration combinations available), the ratio of pollen to nectar energy, floral diversity, floral abundance, and post-fire age. (2) Bee diversity, measured as species richness, was closely linked to floral diversity (especially of annuals), nectar resource diversity, and post-fire age of the habitat. (3) The abundance of the most common species was primarily related to post-fire age, grazing intensity, and nesting substrate availability. Ordination models based on age-characteristic post-fire floral community structure explained 39-50% of overall variation observed in bee community structure. Cluster analysis showed that all the communities shared a high degree of similarity in their species composition (27-59%); however, the geographical location of sites also contributed a smaller but significant component to bee community structure. We conclude that floral resources act in specific and previously unexplored ways to modulate the diversity of the local geographic species pool, with specific disturbance factors, superimposed upon these patterns, mainly affecting the dominant species.
Agricultural conversion is one of the most prevalent anthropogenic uses on the terrestrial earth. Persistence of organisms in such landscapes is thought to be related to species-specific characteristics such as life history traits and dispersal distance. In an agricultural landscape in California, we examined local (farm-level) and landscape variables associated with nesting preferences of native ground-nesting bees. Compared to the known ground-nesting visitors to crops, bee community nesting on farms was depauperate. Further, more abundant and diverse communities of bees were found nesting at farms with patches of natural habitat near by than farms that were far away from natural habitat. Species responded differently to soil conditions created by farming practices, but the variability in nesting bee abundance was lower in farms near natural habitat than farms far from natural habitat. These findings suggest that most bee species are affected adversely but to varying degrees by agricultural intensification, and that natural habitats may buffer against the bee population variability in agricultural landscape. We present source/sink dynamics and resource limitations as possible explanations for the observed patterns.
1. Does the diversity and abundance of one trophic level affect another? Several studies at the landscape level have found a positive relationship between the diversity of floral resources and the diversity and abundance of pollinators. However, little is known about the relationship between these trophic levels on a smaller spatial scale, and the importance of blossom density relative to plant species richness in predicting abundance and richness of different flower visitor groups.
2. This study used a small‐scale approach to investigate how, and if, the diversity and abundance of floral resources in study plots affected the visitation activity of different flower visitor groups. During 201 observation periods between late May and mid‐August 2003, 3682 visits were observed. Bumblebees (60%), muscoids (17%), syrphids (9%), and beetles (5%) were the most abundant flower visitors.
3. Regression analysis was used to investigate the relationship between blossom density and plant species richness with visitation activity, including the probability of presence in plots, the visits within plots, and the visitor richness of the most abundant pollinator groups.
4. The activity of beetles, bumblebees, and muscoids was positively predicted by the variation in blossom density, while syrphid activity was better predicted by plant species richness. Overall, the models for beetles and bumblebees explained much more of the variation in activity compared with the models for the dipterans, and blossom density was a better predictor of both flower visitor richness and activity than was plant species richness.
1. We quantified pollen deposition on the stigma, pollen removal from the anthers and pollen losses in Echium vulgare, visited by workers of Bombus terrestris under controlled conditions. We used dye as a pollen analogue. Bumble-bees were trained to visit a sequence of non-emasculated flowers to estimate pollen carryover and to visit individual flowers to estimate pollen loss.
2. Carryover of pollen grains and dye particles between flowers was similar, which justifies using dye as a pollen analogue. On average 93·8% of the dye particles on the bee were carried over to the next flower. Only a small fraction of the pollen grains was deposited on the stigma (0·15%). A much larger fraction (6·1%) was lost in another way: passively during flight, through grooming or on floral parts other than the stigma. The bees removed 44% of the pollen grains from a fresh flower and 50·3% of this removed pollen adhered to the bee.
3. We predict that, using the parameters mentioned above, during a single visit to a newly opened flower, a bee collects an amount of pollen grains which will bring about 60% geitonogamous self-pollination in the next flower visited. The expected percentage of self-pollination is considerably less if bees visit flowers that have been visited before.
In light of growing concerns over the implications of many conventional agricultural practices, and especially the deep tilling of soils, the Food and Agriculture Organization of the United Nations (FAO), among others, has begun to promote a package of soil conserving practices under the banner of ‘conservation agriculture’. While the title might be novel, its associated practices have long been employed by farmers, and studied by social scientists seeking to understand the reasons for their adoption and non-adoption. This paper reviews and synthesizes this past research in order to identify those independent variables that regularly explain adoption, and thereby facilitate policy prescriptions to augment adoption around the world. While a disaggregated analysis of a subset of commonly used variables reveals some underlying patterns of influence, once various contextual factors (e.g. study locale or method) are controlled, the primary finding of the synthesis is that there are few if any universal variables that regularly explain the adoption of conservation agriculture across past analyses. Given the limited prospect of identifying such variables through further research, we conclude that efforts to promote conservation agriculture will have to be tailored to reflect the particular conditions of individual locales.
The areas of wild land around the edges of agricultural fields are a vital resource for many species. These include insect pollinators, to whom field margins provide both nest sites and important resources (especially when adjacent crops are not in flower). Nesting pollinators travel relatively short distances from the nest to forage: most species of bee are known to travel less than two kilometres away. In order to ensure that these pollinators have sufficient areas of wild land within reach of their nests, agricultural landscapes need to be designed to accommodate the limited travelling distances of nesting pollinators. We used a spatially-explicit modelling approach to consider whether increasing the width of wild strips of land within the agricultural landscape will enhance the amount of wild resources available to a nesting pollinator, and if it would impact differently on pollinators with differing foraging strategies. This was done both by creating field structures with a randomised geography, and by using landscape data based upon the British agricultural landscape. These models demonstrate that enhancing field margins should lead to an increase in the availability of forage to pollinators that nest within the landscape. With the exception of species that only forage within a very short range of their nest (less than 125 m), a given amount of field margin manipulation should enhance the proportion of land available to a pollinator for foraging regardless of the distance over which it normally travels to find food. A fixed amount of field edge manipulation should therefore be equally beneficial for both longer-distance nesting foragers such as honeybees, and short-distance foragers such as solitary bees.
Recent declines of bee species have led to great interest in preserving and promoting bee populations for agricultural and wild plant pollination. Many correlational studies have examined the indirect effects of factors such as landscape context and land management practices and found great variation in bee response. We focus here on the evidence for effects of direct factors (i.e., food resources, nesting resources, and incidental risks) regulating bee populations and then interpret varied responses to indirect factors through their species-specific and habitat-specific effects on direct factors. We find strong evidence for food resource availability regulating bee populations, but little clear evidence that other direct factors are commonly limiting. We recommend manipulative experiments to illuminate the effects of these different factors. We contend that much of the variation in impact from indirect factors, such as grazing, can be explained by the relationships between indirect factors and floral resource availability based on environmental circumstances.
1. Solitary bees are central place foragers returning to their nests several times a day with pollen and nectar to provision their brood cells. They are especially susceptible to landscape changes that lead to an increased spatial separation of suitable nesting sites and flower rich host plant stands. While knowledge of bee foraging ranges is currently growing, quantitative data on the costs of foraging flights are very scarce, although such data are crucial to understand bee population dynamics.
2. In this study, the impact of increased foraging distance on the duration of foraging bouts and on the number of brood cells provisioned per time unit was experimentally quantified in the two pollen specialist solitary bee species Hoplitis adunca and Chelostoma rapunculi. Females nesting at different sites foraged under the same environmental conditions on a single large and movable flowering host plant patch in an otherwise host plant free landscape.
3. The number of brood cells provisioned per time unit by H. adunca was found to decrease by 23%, 31% and 26% with an increase in the foraging distance by 150, 200 and 300 m, respectively. The number of brood cells provisioned by C. rapunculi decreased by 46% and 36% with an increase in the foraging distance by 500 and 600 m, respectively.
4. Contrary to expectation, a widely scattered arrangement of host plants did not result in longer mean duration of a foraging bout in H. adunca compared to a highly aggregated arrangement, which might be due to a reduced flight directionality combined with a high rate of revisitation of already depleted flowers in the aggregated plant arrangement or by a stronger competition and disturbance by other flower visitors.
5. The results of this study clearly indicate that a close neighbourhood of suitable nesting and foraging habitats is crucial for population persistence and thus conservation of endangered solitary bee species.
The spectral representation of stationary Gaussian processes via the Fourier basis provides a computationally efficient specification of spatial surfaces and nonparametric regression functions for use in various statistical models. I describe the representation in detail and introduce the spectralGP package in R for computations. Because of the large number of basis coefficients, some form of shrinkage is necessary; I focus on a natural Bayesian approach via a particular parameterized prior structure that approximates stationary Gaussian processes on a regular grid. I review several models from the literature for data that do not lie on a grid, suggest a simple model modification, and provide example code demonstrating MCMC sampling using the spectralGP package. I describe reasons that mixing can be slow in certain situations and provide some suggestions for MCMC techniques to improve mixing, also with example code, and some general recommendations grounded in experience.
A guide to using S environments to perform statistical analyses providing both an introduction to the use of S and a course in modern statistical methods. The emphasis is on presenting practical problems and full analyses of real data sets.
The rise in the use of statistical models for non‐Gaussian data, such as generalized linear models ( GLM s) and generalized linear mixed models ( GLMM s), is pushing aside the traditional approach of transforming data and applying least‐squares linear models ( LM s). Nonetheless, many least‐squares statistical tests depend on the variance of the sum of residuals, which by the Central Limit Theorem converge to a Gaussian distribution for large sample sizes. Therefore, least‐squares LM s will likely have good performance in assessing the statistical significance of regression coefficients.
Using simulations of count data, I compared GLM approaches for testing whether regression coefficients differ from zero with the traditional approach of applying LM s to transformed data. Simulations assumed that variation among sample populations was either (i) negative binomial or (ii) log‐normal Poisson (i.e. log‐normal variation among populations that were then sampled by a Poisson distribution). I used the simulated data to conduct tests of the hypotheses that regression coefficients differed from zero; I did not investigate statistical properties of the coefficient estimators, such as bias and precision.
For negative binomial simulations whose assumptions closely matched the GLM s, the GLM s were nonetheless prone to type I errors (false positives) especially when there was more than one predictor (independent) variable. After correcting for type I errors, however, the GLM s provided slightly better statistical power than LM s. For log‐normal‐Poisson simulations, both a GLMM and the LM s performed well, but under some simulated conditions the GLM s had high type I error rates, a deadly sin for statistical tests.
These results show that, while GLM s have slight advantages in power when they are properly specified, they can lead to badly wrong conclusions about the significance of regression coefficients if they are mis‐specified. In contrast, transforming data and applying least‐squares linear analyses provide robust statistical tests for significance over a wide range of conditions. Thus, the traditional approach of transforming data and applying LM s is still useful.
In an abandoned field near Ithaca, New York, bee species partitioned the available flower resources, primarily by foraging at different times of the season and by visiting different flower species. Honey bees Apis mellifera concentrated on different flower species than did native bees. Spring-flying native bees included many species adapted to forage on native flower species that bloom before the tree canopy closes. Apis mellifera outcompeted native bees at large clusters of attractive flower species such as apple and the crucifer Barbarea vulgaris. Halictus ligatus was the most common early summer halictine. This species concentrated its foraging on different flowers than did other early summer bees, probably because of differences in innate flower preferences. A profuse bloom of native goldenrods Solidago in late summer was the seasonal peak of resource abundance. Native bees must complete their seasonal cycles and enter diapause before the killing frost. They therefore foraged on early-blooming goldenrod species such as S. juncea. In contrast, honey bees overwinter as active bees within their hives. They were most common on later-blooming goldenrods, where they collected much of the nectar required for overwintering. Competition among foraging bees was probably most important in the spring at this site. Partitioning of flower species and differences in seasonal flight times resulted primarily from differences in innate flower preferences, number of generations per season, and means of overwintering.-Author
Nesting resources structure native bee communities and the availability of suitable nests may enhance population abundance and persistence. Nesting rates of ground-nesting bees have proven challenging to assess due to a lack of standardized methods, We quantified the abundance of ground-nesting native bees using emergence traps over a seven-month study period. We then compared specimens captured in emergence traps with pan- and net-collected specimens. We hypothesized that ground-nesting bees would be highly similar to bees found foraging within our study site. However, the species assemblage of ground-nesting bees collected from emergence traps was significantly dissimilar from the assemblages collected with aerial nets and pan traps, indicating different sampling methods target different components of the species assemblage. We then examined the importance of nesting resources found at each emergence trap on the abundance of ground-nesting bees collected from emergence traps. Quantification of potential nesting resources, such as percent bare soil, has been proposed as a proxy of nesting habitat for ground-nesting guilds. Sloped ground and soil compaction were the most predictive nesting resources at the community-level. Further, spatial distribution of nesting resources within the study landscape also affected nesting rates, although this varied by species. Bees occurred in 85% of emergence traps, with sampling date strongly affecting the number of bees collected. Emergence traps provide a useful method of sampling the ground-nesting native bee community and investigating nesting incidence.
Biotic pollination is an important ecosystem service for the production of many food crops. The supply of pollination is mostly studied at the landscape scale while recent studies on the demand for pollination services provide a global-scale picture based on aggregate national-level data. This paper quantifies both demand and supply of pollination in the European Union (EU) at a relatively high spatial resolution, allowing an analysis of the match between demand and supply. Finally, we evaluate how policies interact with the spatial differences between demand and supply of this ecosystem service.
Habitat loss from urban development threatens native plant populations in many regions of the world. In addition to direct plant mortality, urban intensification potentially impacts pollinator communities and in turn disrupts the pollination mutualisms that are critical to the viability of native plant populations. We placed standardized flowering plant arrays into woodlands along a gradient of increasing urban land use to simultaneously quantify landscape-scale and local-scale effects on pollinators and on reproduction of two spring ephemeral wildflowers (Claytonia virginica and Polemonium reptans) in woodland fragments in the Mid-Atlantic Region of North America. Greater pollinator abundance and associated diversity significantly reduced the degree of pollen limitation, demonstrating that pollinator populations are critical to successful pollination of these plant populations. However, landscape-scale habitat loss did not reduce pollinator abundance or diversity. Habitat loss at the landscape scale therefore does not appear to drive changes in pollination in this woodland system. Rather, local-scale habitat characteristics were more important, with pollinators being more abundant in brighter woodland patches for one plant species, and in larger patches for the other species. Because we found abundant pollinators and adequate pollination even in isolated, urban woodland fragments, our results are encouraging for the conservation of both plants and pollinators in urban landscapes.
In intensive agricultural landscapes, restoration within farms could enhance biodiversity and ecosystem services such as pollination by native pollinators. Although governments and conservation groups are promoting small-scale restoration on working farms, there are few studies that assess whether these practices enhance pollinator communities in restored areas. Further, there is no information on whether floral enhancements will deplete pollinators in adjacent fields by concentrating ambient populations or whether they result in a net increase in abundance in adjacent farm fields. We investigated whether field edges restored with native perennial plants in California's Central Valley agricultural region increased floral abundance and potential bee nesting sites, and native bee and syrphid fly abundance and diversity, in comparison to relatively unmanaged edges. Native bees and syrphid flies collected from flowers were more abundant, species-rich, and diverse at hedgerow sites than in weedy, unmanaged edg...
Variation in the availability of food resources over space and time is a likely driver of how landscape structure and composition affect animal populations. Few studies, however, have directly assessed the spatiotemporal variation in resource availability that arises from landscape pattern, or its effect on populations and population dynamic parameters. We tested the effect of floral resource availability at the landscape scale on the numbers of worker, male, and queen offspring produced by bumble bee, Bombus vosnesenśkii, colonies experimentally placed within complex agricultural-natural landscapes. We quantified flower densities in all land use types at different times of the season and then used these data to calculate spatially explicit estimates of floral resources surrounding each colony. Floral availability strongly correlated with landscape structure, and different regions of the landscape showed distinct seasonal patterns of floral availability. The floral resources available in the landscape surrounding a colony positively affected the number of workers and males it produced. Production was more sensitive to early- than to later-season resources. Floral resources did not significantly affect queen production despite a strong correlation between worker number and queen number among colonies. No landscape produced high floral resources during both the early and late season, and seasonal consistency is likely required for greater queen production. Floral resources are important determinants of colony growth and likely affect the pollination services provided by bumble bees at a landscape scale. Spatiotemporal variation in floral resources across the landscape precludes a simple relationship between resources and reproductive success as measured by queens, but nonetheless likely influences the total abundance of bumble bees in our study region.
Flight patterns of honeybees (Apis mellifera ligustica) were quantified as the bees foraged among artificial flowers for sugar solution (nectar). Bees exhibited considerable directionality on successive flights which minimized repeat visits to flowers and they usually made short flights to nearby flowers, thus minimizing flight time. The change in direction on successive flights between flowers were independent of the number of immediately preceding consecutive rewarding visits but decreased as the number of non-rewarding visits increased. Flight distances were short after visits to rewarding flowers but increased as the number of immediately preceding non-rewarding visits increased. The bees' rate of caloric intake (calories/time) was highest at the floral arrays having the highest density, and it was greater at arrays with clumped nectar-distributions than at those with randomly distributed nectar. These findings are explained in terms of the observed flight patterns.
Human domination of the biosphere has greatly altered ecosystems, often overwhelming their capacity to provide ecosystem services critical to our survival. Yet ecological understanding of ecosystem services is quite limited. Previous work maps the supply and demand for services, assesses threats to them, and estimates economic values, but does not measure the underlying role of biodiversity in providing services. In contrast, experimental studies of biodiversity-function examine communities whose structures often differ markedly from those providing services in real landscapes. A bridge is needed between these two approaches. To develop this research agenda, I discuss critical questions and key approaches in four areas: (1) identifying the important 'ecosystem service providers'; (2) determining the various aspects of community structure that influence function in real landscapes, especially compensatory community responses that stabilize function, or non-random extinction sequences that rapidly erode it; (3) assessing key environmental factors influencing provision of services, and (4) measuring the spatio-temporal scale over which providers and services operate. I show how this research agenda can assist in developing environmental policy and natural resource management plans.
"Changes in flower-visiting insect populations or communities that result from human impacts can be documented by measuring spatial or temporal trends, or by comparing abundance or species composition before and after disturbance. The level of naturally occurring variation in populations and communities over space and time will dictate the sampling effort required to detect human-induced changes. We compiled a set of existing surveys of the bee faunas of natural communities from around the world to examine patterns of abundance and richness. We focused on a subset of these studies to illustrate variation in bee communities among different sites and within sites over different spatial and temporal scales. We used examples from our compilation and other published studies to illustrate sampling approaches that maximize the value of future sampling efforts. Existing studies suggest that bee faunas are locally diverse, highly variable in space and time, and often rich in rare species. All of these attributes indicate that intense sampling among sites and years will be required to differentiate changes due to specific impacts from the natural dynamics of populations and communities. Given the limits on time and funding for studying bees, approaches that maximize information for effort must be sought for future studies.Reliable information on population and community changes may be gleaned from examining 'functional groups' rather than entire faunas. Regardless of the purpose of the study, standardized sampling protocols using replicated designs will increase the value of data. Standardization permits statistical testing of changes in bee populations and communities, and allows for rigorous comparison between studies."
Parallel declines of wild pollinators and pollinator-dependent plants have raised alarms over the loss of pollination services in agroecosystems. A spatially explicit approach is needed to develop specific recommendations regarding the design of agricultural landscapes to sustain wild pollinator communities and the services they provide. I modeled pollination services in agroecosystems using a pair of models: a stochastic individual-based simulation model of wild pollinators, pollinator-dependent plants, and crop pollination; and a set of coupled difference equations designed to capture the nonspatial component of the simulation model. Five spatially explicit models of habitat conversion to crops were simulated, and results for pollination services were compared. Mean-field behavior of the simulation model was in good agreement with analysis of the difference equations. A major feature of the models was the presence of a cusp leading to loss of stability and extinction of pollinators and pollinator-dependent plants beyond a critical amount of habitat loss. The addition of pollen obtained from crop visitation caused a breakdown of the cusp preventing extinction of pollinators, but not of wild pollinator-dependent plants. Spatially restricted foraging and dispersal also altered model outcomes relative to mean-field predictions, in some cases causing extinction under parameter settings that would otherwise lead to persistence. Different patterns of habitat conversion to crops resulted in different levels of pollination services. Most interesting was the finding that optimal pollination services occurred when the size of remnant habitat patches was equal to half the mean foraging and dispersal distance of pollinators and the spacing between remnant patches was equal to the mean foraging and dispersal distance. Conservation of wild pollinators and pollinator-dependent plants in agroecosystems requires careful attention to thresholds in habitat conversion and spatial pattern and scale of remnant habitats. Maximization of pollination services was generally incompatible with conservation of wild pollinator-dependent plants. My prediction is that pollination services will be maximized by providing islands of nesting habitat where interisland distance matches mean foraging distances of wild pollinators.
Many ecosystem services are delivered by organisms that depend on habitats that are segregated spatially or temporally from the location where services are provided. Management of mobile organisms contributing to ecosystem services requires consideration not only of the local scale where services are delivered, but also the distribution of resources at the landscape scale, and the foraging ranges and dispersal movements of the mobile agents. We develop a conceptual model for exploring how one such mobile-agent-based ecosystem service (MABES), pollination, is affected by land-use change, and then generalize the model to other MABES. The model includes interactions and feedbacks among policies affecting land use, market forces and the biology of the organisms involved. Animal-mediated pollination contributes to the production of goods of value to humans such as crops; it also bolsters reproduction of wild plants on which other services or service-providing organisms depend. About one-third of crop production depends on animal pollinators, while 60-90% of plant species require an animal pollinator. The sensitivity of mobile organisms to ecological factors that operate across spatial scales makes the services provided by a given community of mobile agents highly contextual. Services vary, depending on the spatial and temporal distribution of resources surrounding the site, and on biotic interactions occurring locally, such as competition among pollinators for resources, and among plants for pollinators. The value of the resulting goods or services may feed back via market-based forces to influence land-use policies, which in turn influence land management practices that alter local habitat conditions and landscape structure. Developing conceptual models for MABES aids in identifying knowledge gaps, determining research priorities, and targeting interventions that can be applied in an adaptive management context.
Bees are the most important pollinator taxon; therefore, understanding the scale at which they forage has important ecological implications and conservation applications. The foraging ranges for most bee species are unknown. Foraging distance information is critical for understanding the scale at which bee populations respond to the landscape, assessing the role of bee pollinators in affecting plant population structure, planning conservation strategies for plants, and designing bee habitat refugia that maintain pollination function for wild and crop plants. We used data from 96 records of 62 bee species to determine whether body size predicts foraging distance. We regressed maximum and typical foraging distances on body size and found highly significant and explanatory nonlinear relationships. We used a second data set to: (1) compare observed reports of foraging distance to the distances predicted by our regression equations and (2) assess the biases inherent to the different techniques that have been used to assess foraging distance. The equations we present can be used to predict foraging distances for many bee species, based on a simple measurement of body size.
We analysed the dynamics of a plant-pollinator interaction network of a scrub community surveyed over four consecutive years. Species composition within the annual networks showed high temporal variation. Temporal dynamics were also evident in the topology of the network, as interactions among plants and pollinators did not remain constant through time. This change involved both the number and the identity of interacting partners. Strikingly, few species and interactions were consistently present in all four annual plant-pollinator networks (53% of the plant species, 21% of the pollinator species and 4.9% of the interactions). The high turnover in species-to-species interactions was mainly the effect of species turnover (c. 70% in pairwise comparisons among years), and less the effect of species flexibility to interact with new partners (c. 30%). We conclude that specialization in plant-pollinator interactions might be highly overestimated when measured over short periods of time. This is because many plant or pollinator species appear as specialists in 1 year, but tend to be generalists or to interact with different partner species when observed in other years. The high temporal plasticity in species composition and interaction identity coupled with the low variation in network structure properties (e.g. degree centralization, connectance, nestedness, average distance and network diameter) imply (i) that tight and specialized coevolution might not be as important as previously suggested and (ii) that plant-pollinator interaction networks might be less prone to detrimental effects of disturbance than previously thought. We suggest that this may be due to the opportunistic nature of plant and animal species regarding the available partner resources they depend upon at any particular time.
Quantifying and mapping ecosystem services
- C Schulp
- S Lautenbach
- P Verburg
Schulp, C., S. Lautenbach, and P. Verburg. 2014. Quantifying and mapping ecosystem services:
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A new method for studying pollen dispersal using micronized Variation in native bee faunas and its 679 implications for detecting community changes
- R E Stockhouse
- N M Williams
- R L Minckley
- F A Silveira
Stockhouse, R.E. 1976. A new method for studying pollen dispersal using micronized Williams, N. M., R. L. Minckley, and F. A. Silveira. 2001. Variation in native bee faunas and its
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implications for detecting community changes. Conservation Ecology 5: 7.
Wild bees enhance honey bees pollination of hybrid 551 sunflower
- S S Greenleaf
- C Kremen
Greenleaf, S.S. and C. Kremen. 2006. Wild bees enhance honey bees pollination of hybrid
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sunflower. Proceedings of the National Academy of Sciences 103: 13890–13895.
Bees in a Changing World: How land surface phenology, bee community 587 distributions, and pollinator-plant interactions are impacted by urbanization and agriculture
- M Leong
Leong, M. 2014. Bees in a Changing World: How land surface phenology, bee community
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distributions, and pollinator-plant interactions are impacted by urbanization and agriculture.
vegan: Community Ecology Package
- P Solymos
- M H H Stevens
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package version 2.0-10. URL http://CRAN.R-project.org/package=vegan
How efficient are bees in pollinating sunflowers
- F D Parker
Parker, F.D. 1981 How efficient are bees in pollinating sunflowers? Journal of the Kansas
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Notes on the biology of a common 613 sunflower bee, Melissodes (Eumelissodes) agilis Cresson
- F D Parker
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- G E Bohart
Parker, F. D., V. J. Tepedino, and G. E. Bohart. 1981. Notes on the biology of a common
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sunflower bee, Melissodes (Eumelissodes) agilis Cresson. Journal of the New York
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Using Farm Bill programs for pollinator conservation
- M Vaughan
- M Skinner
Vaughan, M. and M. Skinner. 2008. Using Farm Bill programs for pollinator conservation.
Long foraging 689 distances impose high costs on offspring production in solitary bees
- A Zurbuchen
- S Cheesman
- J Klaiber
- A Müler
- S Hein
- S Dorn
Zurbuchen, A., S. Cheesman, J. Klaiber, A. Müler, S. Hein, and S. Dorn. 2010. Long foraging
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distances impose high costs on offspring production in solitary bees. Journal of Animal Ecology
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79: 674-681.