Density-dependent effects of ants on selection for bumble bee pollination in Polemonium viscosum

Mutualisms are commonly exploited by cheater species that usurp rewards without providing reciprocal benefits. Yet most studies of selection between mutualist partners ignore interactions with third species and consequently overlook the impact of cheaters on evolution in the mutualism. Here, we explicitly investigate how the abundance of nectar-thieving ants (cheaters) influences selection in a pollination mutualism between bumble bees and the alpine skypilot, Polemonium viscosum. As suggested in past work with this species, bumble bees accounted for most of the seed production (78% +/- 6% [mean +/- SE]) in our high tundra study population and, in the absence of ants, exerted strong selection for large flowers. We tested for indirect effects of ant abundance on seed set through bumble bee pollination services (pollen delivery and pollen export) and a direct effect through flower damage. Ants reduced seed set per flower by 20% via flower damage. As ant density increased within experimental patches, the rate of flower damage rose, but pollen delivery and export did not vary significantly, showing that indirect effects of increased cheater abundance on pollinator service are negligible in this system. To address how ants affect selection for plant participation in the pollination mutualism we tested the impact of ant abundance on selection for bumble bee-mediated pollination. Results show that the impact of ants on fitness (seed set) accruing under bumble bee pollination is density dependent in P. viscosum. Selection for bumble bee pollination declined with increasing ant abundance in experimental patches, as predicted if cheaters constrain fitness returns of mutualist partner services. We also examined how ant abundance influences selection on flower size, a key component of plant investment in bumble bee pollination. We predicted that direct effects of ants would constrain bumble bee selection for large flowers. However, selection on flower size was significantly positive over a wide range of ant abundance (20-80% of plants visited by ants daily). Although high cheater abundance reduces the fitness returns of bumble bee pollination, it does not completely eliminate selection for bumble bee attraction in P. viscosum.     

Herbivory-mediated pollinator limitation: negative impacts of induced volatiles on plant-pollinator interactions

Although induced plant responses to herbivory are well studied as mechanisms of resistance, how induction shapes community interactions and ultimately plant fitness is still relatively unknown. Using a wild tomato, Solanum peruvianum, native to the Peruvian Andes, we evaluated the disruption of pollination as a potential ecological cost of induced responses. More specifically, we tested the hypothesis that metabolic changes in herbivore-attacked plants, such as the herbivore-induced emission of volatile organic compounds (VOCs), alter pollinator behavior and consequentially affect plant fitness. We conducted a series of manipulative field experiments to evaluate the role of herbivore-induced vegetative and floral VOC emissions as mechanisms by which herbivory affects pollinator behavior. In field surveys and bioassays in the plants' native habitat, we found that real and simulated herbivory (methyl jasmonate application) reduced attractiveness of S. peruvianum flowers to their native pollinators. We show that reduced pollinator preference, not resource limitation due to leaf tissue removal, resulted in reduced seed set. Solitary bee pollinators use floral plant volatiles, emitted in response to herbivory or methyl jasmonate treatment, as cues to avoid inflorescences on damaged plants. This herbivory-induced pollinator limitation can be viewed as a general cost of induced plant responses as well as a specific cost of herbivory-induced volatile emission.     

Modeling scale up of anthropogenic impacts from individual pollinator behavior to pollination systems

Understanding how anthropogenic disturbances affect plant–pollinator systems has important implications for the conservation of biodiversity and ecosystem functioning. Previous laboratory studies show that pesticides and pathogens, which have been implicated in the rapid global decline of pollinators over recent years, can impair behavioral processes needed for pollinators to adaptively exploit floral resources and effectively transfer pollen among plants. However, the potential for these sublethal stressor effects on pollinator–plant interactions at the individual level to scale up into changes to the dynamics of wild plant and pollinator populations at the system level remains unclear. We developed an empirically parameterized agent-based model of a bumblebee pollination system called SimBee to test for effects of stressor-induced decreases in the memory capacity and information processing speed of individual foragers on bee abundance (scenario 1), plant diversity (scenario 2), and bee–plant system stability (scenario 3) over 20 virtual seasons. Modeling of a simple pollination network of a bumblebee and four co-flowering bee-pollinated plant species indicated that bee decline and plant species extinction events could occur when only 25% of the forager population showed cognitive impairment. Higher percentages of impairment caused 50% bee loss in just five virtual seasons and system-wide extinction events in less than 20 virtual seasons under some conditions. Plant species extinctions occurred regardless of bee population size, indicating that stressor-induced changes to pollinator behavior alone could drive species loss from plant communities. These findings indicate that sublethal stressor effects on pollinator behavioral mechanisms, although seemingly insignificant at the level of individuals, have the cumulative potential in principle to degrade plant–pollinator species interactions at the system level. Our work highlights the importance of an agent-based modeling approach for the identification and mitigation of anthropogenic impacts on plant–pollinator systems.     

Geographic structure of pollinator communities, reproductive assurance, and the evolution of self-pollination

Reproductive assurance is often invoked as an explanation for the evolution of self-fertilization in plants. However, key aspects of this hypothesis have received little empirical support. In this study, I use geographic surveys of pollinator communities along with functional studies of floral trait variation to examine the role of pollination ecology in mating system differentiation among populations and subspecies of the annual plant Clarkia xantiana. A greenhouse experiment involving 30 populations from throughout the species' range indicated that variation in two floral traits, herkogamy and protandry, was closely related to levels of autofertility and that trait variation was partitioned mainly among populations. Emasculation experiments in the field showed that autonomous selfing confers reproductive assurance by elevating fruit and seed production. Surveys of pollinator communities across the geographic range of the species revealed that bee pollinator abundance and community composition differed dramatically between populations of the outcrossing subspecies xantiana and the selfing subspecies parviflora despite their close proximity. Specialist bee pollinators of Clarkia were absent from selfing populations, but they were the most frequent visitors to outcrossing populations. Moreover, within the outcrossing subspecies xantiana, there was a close correspondence between specialist abundance and population differentiation in herkogamy, a key mating system trait. This spatial covariation arose, in part, because geographically peripheral populations had reduced herkogamy, higher autofertility, and lower pollinator abundance compared to central populations of xantiana. Finally, I detected strong spatial structure to bee communities both across the range of the species and within the outcrossing subspecies. In both cases, spatial structure was stronger for specialist bees compared to generalist bees, and pollinator communities varied in parallel with population variation in herkogamy. These results provide evidence that mating system differentiation parallels spatial variation in pollinator abundance and community composition at both broad and more restricted spatial scales, consistent with the hypothesis that pollinator abundance and reproductive assurance are important drivers of plant mating system evolution.     

Bridging the generation gap in plants: pollination, parental fecundity, and offspring demography

Despite extensive study of pollination and plant reproduction on the one hand, and of plant demography on the other, we know remarkably little about links between seed production in successive generations, and hence about long-term population consequences of variation in pollination success. We bridged this "generation gap" in Ipomopsis aggregata, a long-lived semelparous wildflower that is pollinator limited, by adding varying densities of seeds to natural populations and following resulting plants through their entire life histories. To determine whether pollen limitation of seed production constrains rate of population growth in this species, we sowed seeds into replicated plots at a density that mimics typical pollination success and spacing of flowering plants in nature, and at twice that density to mimic full pollination. Per capita offspring survival, flower production, and contribution to population increase (lambda) did not decline with sowing density in this experiment, suggesting that typical I. aggregata populations freed from pollen limitation will grow over the short term. In a second experiment we addressed whether density dependence would eventually erase the growth benefits of full pollination, by sowing a 10-fold range of seed densities that falls within extremes estimated for the natural "seed rain" that reaches the soil surface. Per capita survival to flowering and age at flowering were again unaffected by sowing density, but offspring size, per capita flower production, and lambda declined with density. Such density dependence complicates efforts to predict population dynamics over the longer term, because it changes components of the life history (in this case fecundity) as a population grows. A complete understanding of how constraints on seed production affect long-term population growth will hinge on following offspring fates at least through flowering of the first offspring generation, and doing so for a realistic range of population densities.     

Reproductive Success Increases with Local Density of Conspecif ics in a Desert Mustard (Lesquerella fendleri)

We investigated the effects of plant density on reproduction for an insect-pollinated desert mustard (Lesquerella fendleri [Brassicaceae]). Individual reproductive success, as measured by seeds per fruit, proportion of flowers setting fruit, and total seed production, increased with the density of conspecifics within 1 m. However, including the density of conspecifics at greater distances (1–3 m) did not significantly increase the amount of variation in reproductive success explained by the regression model. This implies that processes occurring on a scale of 1 m or less have important effects on reproduction. Total seed production also was greater for high-density plants than for otherwise similar plants with a low-density of conspecifics. We argue that increased pollinator visitation is the most likely cause of this facilitation and that investigations of the effects of rarity on reproductive success should directly consider density along with more commonly used attributes such as population size and fragmentation.     

Pollinator effectiveness varies with experimental shifts in flowering time

The earlier flowering times exhibited by many plant species are a conspicuous sign of climate change. Altered phenologies have caused concern that species could suffer population declines if they flower at times when effective pollinators are unavailable. For two perennial wildflowers, Tradescantia ohiensis and Asclepias incarnata, we used an experimental approach to explore how changing phenology affects the taxonomic composition of the pollinator assemblage and the effectiveness of individual pollinator taxa. After finding in the previous year that fruit set varied with flowering time, we manipulated flowering onset in greenhouses, placed plants in the field over the span of five weeks, and measured pollinator effectiveness as the number of seeds produced after a single visit to a flower. The average effectiveness of pollinators and the expected rates of pollination success were lower for plants of both species flowering earlier than for plants flowering at historical times, suggesting there could be reproductive costs to earlier flowering. Whereas for A. incarnata, differences in average seed set among weeks were due primarily to changes in the composition of the pollinator assemblage, the differences for T. ohiensis were driven by the combined effects of compositional changes and increases over time in the effectiveness of some pollinator taxa. Both species face the possibility of temporal mismatch between the availability of the most effective pollinators and the onset of flowering, and changes in the effectiveness of individual pollinator taxa through time may add an unexpected element to the reproductive consequences of such mismatches.     

Collapse of a pollination web in small conservation areas

A suspected global decline in pollinators has heightened interest in their ecological significance. In a worst-case scenario, the decline of generalist pollinators is predicted to trigger cascades of linked declines among the multiple specialist plant species to which they are linked, but this has not been documented. I studied a portion of a pollination web involving a generalist pollinator, the oil-collecting bee Rediviva peringueyi, and a community of oil-secreting plants. Across 27 established conservation areas located in the Cape Floral Region, I found substantial variation in the bees' occurrence in relation to soil type and the successional stage of the vegetation. Anthropogenic declines were detectable against this background of naturally occurring variation: R. peringueyi was absent from small conservation areas (< 385 ha) in an urban matrix. In the absence of the bee, seed set failed in six specialist plant species that are pollinated only by R. peringueyi but remained high in a pollination generalist, which had replacement pollinators. The findings are consistent with theoretical predictions of the importance of generalist pollinators in maintaining the structure of pollination webs.     

Effect of pollinator abundance on self-fertilization and gene flow: application to GM Canola.

Cross-pollination from fields of transgenic crops is of great public concern. Although cross-pollination in commercial canola (Brassica napus) fields has been empirically measured, field trials are expensive and do not identify the causes of cross-pollination. Therefore, theoretical models can be valuable because they can provide estimates of cross-pollination at any given site and time. We present a general analytical model of field-to-field gene flow due to the following competing mechanisms: the wind, bees, and autonomous pollination. We parameterize the model for the particular case of field-to-field cross-pollination of genetically modified (GM) canola via the wind and via bumble bees (Bombus spp.) and honey bees (Apis mellifera). We make extensive use of the large data set of bee densities collected during the recent U.K. Farm Scale Evaluations. We predict that canola approaches almost full seed set without pollinators and that autonomous pollination is responsible for > or = 25% of seed set, irrespective of pollinator abundance. We do not predict the relative contribution of bees vs. the wind in landscape-scale gene flow in canola. However, under model assumptions, we predict that the maximum field-to-field gene flow due to bumble bees is 0.04% and 0.13% below the current EU limit for adventitious GM presence for winter- and spring-sown canola, respectively. We predict that gene flow due to bees is approximately 3.1 times higher at 20% compared to 100% male-fertility, and due to the wind, 1.3 times higher at 20% compared to 100% male-fertility, for both winter- and spring-sown canola. Bumble bee-mediated gene flow is approximately 2.7 times higher and wind-mediated gene flow approximately 1.7 times lower in spring-sown than in winter-sown canola, regardless of the degree of male-sterility. The model of cross-pollination due to the wind most closely predicted three previously published observations: field-to-field gene flow is low; gene flow increases with the proportion of plants that are male-sterile; and gene flow is higher in winter- than in spring-sown canola. Our results therefore suggest that the wind, not bees, is the main vector of long-distance gene flow in canola.     

Imperfect Replacement of Native Species by Non‐Native Species as Pollinators of Endemic Hawaiian Plants

Native plant species that have lost their mutualist partners may require non‐native pollinators or seed dispersers to maintain reproduction. When natives are highly specialized, however, it appears doubtful that introduced generalists will partner effectively with them. We used visitation observations and pollination treatments (experimental manipulations of pollen transfer) to examine relationships between the introduced, generalist Japanese White‐eye (Zosterops japonicus) and 3 endemic Hawaiian plant species (Clermontia parviflora, C. montis‐loa, and C. hawaiiensis). These plants are characterized by curved, tubular flowers, apparently adapted for pollination by curve‐billed Hawaiian honeycreepers. Z. japonicus were responsible for over 80% of visits to flowers of the small‐flowered C. parviflora and the midsize‐flowered C. montis‐loa. Z. japonicus‐visited flowers set significantly more seed than did bagged flowers. Z. japonicus also demonstrated the potential to act as an occasional Clermontia seed disperser, although ground‐based frugivory by non‐native mammals likely dominates seed dispersal. The large‐flowered C. hawaiiensis received no visitation by any birds during observations. Unmanipulated and bagged C. hawaiiensis flowers set similar numbers of seeds. Direct examination of Z. japonicus and Clermontia morphologies suggests a mismatch between Z. japonicus bill morphology and C. hawaiiensis flower morphology. In combination, our results suggest that Z. japonicus has established an effective pollination relationship with C. parviflora and C. montis‐loa and that the large flowers of C. hawaiiensis preclude effective visitation by Z. japonicus. Remplazo Imperfecto de Especies Nativas por Especies No‐Nativas como Polinizadores de Plantas Endémicas de Hawaii     

Evolution of polyploidy and the diversification of plant-pollinator interactions

One of the major mechanisms of plant diversification has been the evolution of polyploid populations that differ from their diploid progenitors in morphology, physiology, and environmental tolerances. Recent studies have indicated that polyploidy may also have major effects on ecological interactions with herbivores and pollinators. We evaluated pollination of sympatric diploid and tetraploid plants of the rhizomatous herb Heuchera grossulariifolia (Saxifragaceae) along the Selway and Salmon Rivers of northern Idaho, USA, during four consecutive years. Previous molecular and ecological analyses had indicated that the tetraploid populations along these two river systems are independently derived and differ from each other in multiple traits. In each region, we evaluated floral visitation rate by all insect visitors, pollination efficacy of all major visitors, and relative contribution of all major pollinators to seed set. In both regions, diploid and tetraploid plants attracted different suites of floral visitors. Most pollination was attributable to several bee species and the moth Greya politella. Lasioglossum bees preferentially visited diploid plants at Lower Salmon but not at Upper Selway, queen Bombus centralis preferentially visited tetraploids at both sites, and worker B. centralis differed between sites in their cytotype preference. Hence, diploid and autotetraploid H. grossulariifolia plants act essentially as separate ecological species and may experience partial reproductive isolation through differential visitation and pollination by their major floral visitors. Overall the results, together with recent results from other studies, suggest that the repeated evolution of polyploidy in plants may contribute importantly to the structure and diversification of ecological interactions in terrestrial communities.     

Fine-scale flower-visiting behavior revealed by using a high-speed camera

Most estimations of the pollination efficiency of insects have been based on observation by the naked human eye. However, insect behaviors are often too rapid to analyze sufficiently this way. Here we demonstrate the use of high-speed cameras to analyze the fine-scale behaviors of Macroglossum pyrrhosticta, Xylocopa appendiculata, and Papilio dehaanii when visiting Clerodendrum trichotomum. The fine-scale nectar drinking time, number of contacts with anthers and/or stigmas, and frequencies of body part contact with anthers and/or stigmas differed significantly among pollinator species. Pollination efficiency was not equal among pollinators. In addition, M. pyrrhosticta made the least number of contacts with anthers and/or stigmas even though it showed the highest visitation frequency. These results demonstrate that when examined from the viewpoint of rapid visitation behaviors, pollination dynamics differ among pollinator species, and flower visits and pollination rates are not equal.     

Nectar secondary compounds affect self-pollen transfer: implications for female and male reproduction

Pollen movement within and among plants affects inbreeding, plant fitness, and the spatial scale of genetic differentiation. Although a number of studies have assessed how plant and floral traits influence pollen movement via changes in pollinator behavior, few have explored how nectar chemical composition affects pollen transfer. As many as 55% of plants produce secondary compounds in their nectar, which is surprising given that nectar is typically thought to attract pollinators. We tested the hypothesis that nectar with secondary compounds may benefit plants by encouraging pollinators to leave plants after visiting only a few flowers, thus reducing self-pollen transfer. We used Gelsemium sempervirens, a plant whose nectar contains the alkaloid gelsemine, which has been shown to be a deterrent to foraging bee pollinators. We found that high nectar alkaloids reduced the total and proportion of self-pollen received by one-half and one-third, respectively. However, nectar alkaloids did not affect female reproduction when we removed the potential for self-pollination (by emasculating all flowers on plants). We then tested the assumption that self-pollen in combination with outcrossed pollen depresses seed set. We found that plants were weakly self-compatible, but self-pollen with outcrossed pollen did not reduce seed set relative to solely outcrossed flowers. Finally, an exponential model of pollen carryover suggests that high nectar alkaloids could benefit plants via increased pollen export (an estimate of male function), but only when pollinators were efficient and abundant and plants had large floral displays. Results suggest that high nectar alkaloids may benefit plants via increased pollen export under a restricted set of ecological conditions, but in general, the costs of high nectar alkaloids in reducing pollination balanced or outweighed the benefits of reducing self-pollen transfer for estimates of female and male reproduction.     

Herbivory reduces plant interactions with above- and belowground antagonists and mutualists

Herbivores affect plants through direct effects, such as tissue damage, and through indirect effects that alter species interactions. Interactions may be positive or negative, so indirect effects have the potential to enhance or lessen the net impacts of herbivores. Despite the ubiquity of these interactions, the indirect pathways are considerably less understood than the direct effects of herbivores, and multiple indirect pathways are rarely studied simultaneously. We placed herbivore effects in a comprehensive community context by studying how herbivory influences plant interactions with antagonists and mutualists both aboveground and belowground. We manipulated early-season aboveground herbivore damage to Cucumis sativus (cucumber, Cucurbitaceae) and measured interactions with subsequent aboveground herbivores, root-feeding herbivores, pollinators, and arbuscular mycorrhizal fungi (AMF). We quantified plant growth and reproduction and used an enhanced pollination treatment to determine if plants were pollen limited. Increased herbivory reduced interactions with both antagonists and mutualists. Plants with high levels of early herbivory were significantly less likely to suffer leaf damage later in the summer and tended to be less attacked by root herbivores. Herbivory also reduced pollinator visitation, likely due to fewer and smaller flowers, and reduced AMF colonization. The net effect of herbivory on plant growth and reproduction was strongly negative, but lower fruit and seed production were not due to reduced pollinator visits, because reproduction was not pollen limited. Although herbivores influenced interactions between plants and other organisms, these effects appear to be weaker than the direct negative effects of early-season tissue loss.     

Reduced pollinator service and elevated pollen limitation at the geographic range limit of an annual plant

Mutualisms are well known to influence individual fitness and the population dynamics of partner species, but little is known about whether they influence species distributions and the location of geographic range limits. Here, we examine the contribution of plant-pollinator interactions to the geographic range limit of the California endemic plant Clarkia xantiana ssp. xantiana. We show that pollinator availability declined from the center to the margin of the geographic range consistently across four years of study. This decline in pollinator availability was caused to a greater extent by variation in the abundance of generalist rather than specialist bee pollinators. Climate data suggest that patterns of precipitation in the current and previous year drove variation in bee abundance because of its effects on cues for bee emergence in the current year and the abundance of floral resources in the previous year. Experimental floral manipulations showed that marginal populations had greater outcross pollen limitation of reproduction, in parallel with the decline in pollinator abundance. Although plants are self-compatible, we found no evidence that autonomous selfing contributes to reproduction, and thus no evidence that it alleviates outcross pollen limitation in marginal populations. Furthermore, we found no association between the distance to the range edge and selfing rate, as estimated from sequence and microsatellite variation, indicating that the mating system has not evolved in response to the pollination environment at the range periphery. Overall, our results suggest that dependence on pollinators for reproduction may be an important constraint limiting range expansion in this system.     

Habitat-specific impacts of multiple consumers on plant population dynamics

Multiple consumers often attack seeds, seedlings, and adult plants, but their population-level consequences remain uncertain. We examined how insect and small mammal consumers influenced the demography and abundance of the perennial shrub, bush lupine (Lupinus arboreus). In grassland and dune habitats we established replicate experimental lupine populations in 81-m2 plots that were either protected from, or exposed to, herbivorous voles and granivorous mice (via fencing) and/or root feeding insects (via insecticide treatment). Populations were initiated with transplanted seedlings in 1999 and 2000. We followed the demography of these cohorts, subsequent generations, and the seed bank for 5.5 years. Voles and insects killed many seedlings in dune (1999 only) and grassland (1999 and 2000) habitats. After 2000, insects and voles had minimal effects on seedling or adult survival. Seed predation by granivorous mice, however, greatly depressed seedling recruitment, resulting in lower adult lupine abundance in control plots vs. those protected from rodents. In grasslands, initial effects of voles and insects on seedling survival produced large differences among treatments in adult plant density and the cumulative number of seeds produced throughout the experiment. Differences among grassland populations in seed rain, however, had little influence on the magnitude of seedling recruitment into this habitat. Instead, recruitment out of a preexisting seed bank compensated for the lack of seed production in populations exposed to consumers. Shading by dense adults in plots protected from consumers limited seedling establishment within these populations. Although differences among populations in cumulative seed rain did not influence adult establishment, populations protected from consumers accumulated substantially larger seed banks than controls. These results illustrate how density dependence, habitat-specific seed dynamics, and particular demographic impacts of consumers interact to shape plant population responses to consumers.     

Pollination in Dianthus deltoides (Caryophyllaceae): Effects of Habitat Fragmentation on Visitation and Seed Set

Abstract: I analyse the effects of habitat fragmentation on the pollination success of a perennial, butterfly-pollinated, caryophyllaceous herb, the maiden pink, Dianthus deltoides L. The study was conducted in July 1986 and July 1987 at two different sites in southwest Sweden, an undisturbed "mainland" site and a fragmented site consisting of "habitat islands" within a heavily utilized agricultural area The fragmented area had a lower diversity and abundance of both flowering plants and flower-visiting insects. Dianthus flowers received fewer visits in the fragmented area than in the mainland area, and the seed set was much lower. Hand pollination increased seed set up to 4.1 times in the fragmented area, but no significant differences were found between hand-pollinated and control flowers at the mainland site. There were no differences between the two sites in standing crop of nectar, ovule number per flowers, or seed set of bagged flowers, band-pollinated flowers, and hand-pollinated fertilized flowers Thus, the difference in natural seed set between the two sites can be explained by differences in pollinator service.     

Evaluating the Quality of Citizen‐Scientist Data on Pollinator Communities

Abstract: Concerns about pollinator declines have grown in recent years, yet the ability to detect changes in abundance, taxonomic richness, and composition of pollinator communities is hampered severely by the lack of data over space and time. Citizen scientists may be able to extend the spatial and temporal extent of pollinator monitoring programs. We developed a citizen‐science monitoring protocol in which we trained 13 citizen scientists to observe and classify floral visitors at the resolution of orders or super families (e.g., bee, wasp, fly) and at finer resolution within bees (superfamily Apoidea) only. We evaluated the protocol by comparing data collected simultaneously at 17 sites by citizen scientists (observational data set) and by professionals (specimen‐based data set). The sites differed with respect to the presence and age of hedgerows planted to improve habitat quality for pollinators. We found significant, positive correlations among the two data sets for higher level taxonomic composition, honey bee (Apis mellifera) abundance, non‐Apis bee abundance, bee richness, and bee community similarity. Results for both data sets also showed similar trends (or lack thereof) in these metrics among sites differing in the presence and age of hedgerows. Nevertheless, citizen scientists did not observe approximately half of the bee groups collected by professional scientists at the same sites. Thus, the utility of citizen‐science observational data may be restricted to detection of community‐level changes in abundance, richness, or similarity over space and time, and citizen‐science observations may not reliably reflect the abundance or frequency of occurrence of specific pollinator species or groups.     

Manuscript in preparation for <Emphasis Type="Italic">Behavioral Ecology and Sociobiology</Emphasis> Bumble bee pollen foraging regulation: role of pollen quality,storage levels,and odor

The regulation of protein collection through pollen foraging plays an important role in pollination and in the life of bee colonies that adjust their foraging to natural variation in pollen protein quality and temporal availability. Bumble bees occupy a wide range of habitats from the Nearctic to the Tropics in which they play an important role as pollinators. However, little is known about how a bumble bee colony regulates pollen collection. We manipulated protein quality and colony pollen stores in lab-reared colonies of the native North American bumble bee, Bombus impatiens. We debut evidence that bumble bee colony foraging levels and pollen storage behavior are tuned to the protein quality (range tested: 17–30% protein by dry mass) of pollen collected by foragers and to the amount of stored pollen inside the colony. Pollen foraging levels (number of bees exiting the nest) significantly increased by 55%, and the frequency with which foragers stored pollen in pots significantly increased by 233% for pollen with higher compared to lower protein quality. The number of foragers exiting the nest significantly decreased (by 28%) when we added one pollen load equivalent each 5 min to already high intranidal pollen stores. In addition, pollen odor pumped into the nest is sufficient to increase the number of exiting foragers by 27%. Foragers directly inspected pollen pots at a constant rate over 24 h, presumably to assess pollen levels. Thus, pollen stores can act as an information center regulating colony-level foraging according to pollen protein quality and colony need. An erratum to this article can be found at     

Detecting Insect Pollinator Declines on Regional and Global Scales

Recently there has been considerable concern about declines in bee communities in agricultural and natural habitats. The value of pollination to agriculture, provided primarily by bees, is >$200 billion/year worldwide, and in natural ecosystems it is thought to be even greater. However, no monitoring program exists to accurately detect declines in abundance of insect pollinators; thus, it is difficult to quantify the status of bee communities or estimate the extent of declines. We used data from 11 multiyear studies of bee communities to devise a program to monitor pollinators at regional, national, or international scales. In these studies, 7 different methods for sampling bees were used and bees were sampled on 3 different continents. We estimated that a monitoring program with 200–250 sampling locations each sampled twice over 5 years would provide sufficient power to detect small (2–5%) annual declines in the number of species and in total abundance and would cost U.S.$2,000,000. To detect declines as small as 1% annually over the same period would require >300 sampling locations. Given the role of pollinators in food security and ecosystem function, we recommend establishment of integrated regional and international monitoring programs to detect changes in pollinator communities. Detección de Declinaciones de Insectos Polinizadores a Escalas Regional y Global