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1. Resource pulses, narrow periods of high resource availability, can elicit strong behavioural responses across diverse taxa. Mass‐flowering agricultural crops are an example of a resource pulse that insect pollinators exploit. However, the underlying mechanism behind changes in pollinator behaviour associated with mass‐flowering crops is still relatively unexplored. 2. The present study quantified the behavioural response of bumble bees, an important wild pollinator, to commercial cranberry bloom, an important mass‐flowering crop in Wisconsin, U.S.A. Over a 2‐year period, foraging trip duration was measured using radio frequency identification at 14 farms situated across landscape contexts, ranging from high to low natural area (woodland amount). Using transect surveys, floral resource abundance at a landscape scale was estimated. 3. It was found that bumble bees were highly sensitive to temporal changes in landscape‐level resource abundance associated with the onset of cranberry bloom, during which they decreased foraging trip duration by 22% and increased the number of foraging trips during bloom by 24% on average relative to the period before and after bloom. This phenomenon was consistent across colonies, individual bees, and landscape contexts, despite a higher abundance of flowers in low woodland landscapes. Bumble bee colonies growing in low‐ and high‐woodland landscapes exhibited a similar performance. 4. As mass‐flowering crops are probably a factor influencing bumble bee foraging behaviour in agricultural regions, investigations should continue into how variable resource landscapes, particularly those offering resource pulses, affect wild pollinators and the pollination services they provide. [ABSTRACT FROM AUTHOR]

Foraging is an essential process for mobile animals, and its optimization serves as a foundational theory in ecology and evolution; however, drivers of foraging are rarely investigated across landscapes and seasons. Using a common bumblebee species from the western United States (Bombus vosnesenskii), we ask whether seasonal decreases in food resources prompt changes in foraging behavior and space use. We employ a unique integration of population genetic tools and spatially explicit foraging models to estimate foraging distances and rates of patch visitation for wild bumblebee colonies across three study regions and two seasons. By mapping the locations of 669 wild-caught individual foragers, we find substantial variation in colony-level foraging distances, often exhibiting a 60-fold difference within a study region. Our analysis of visitation rates indicates that foragers display a preference for destination patches with high floral cover and forage significantly farther for these patches, but only in the summer, when landscape-level resources are low. Overall, these results indicate that an increasing proportion of long-distance foraging bouts take place in the summer. Because wild bees are pollinators, their foraging dynamics are of urgent concern, given the potential impacts of global change on their movement and services. The behavioral shift toward long-distance foraging with seasonal declines in food resources suggests a novel, phenologically directed approach to landscape-level pollinator conservation and greater consideration of late-season floral resources in pollinator habitat management. [ABSTRACT FROM AUTHOR]

Bee population declines are linked to the reduction of nutritional resources due to land-use intensification, yet we know little about the specific nutritional needs of many bee species. Pollen provides bees with their primary source of protein and lipids, but nutritional quality varies widely among host-plant species. Therefore, bees might have adapted to assess resource quality and adjust their foraging behavior to balance nutrition from multiple food sources. We tested the ability of two bumble bee species, Bombus terrestris and Bombus impatiens, to regulate protein and lipid intake. We restricted B. terrestris adults to single synthetic diets varying in protein:lipid ratios (P:L). The bees over-ate protein on low-fat diets and over-ate lipid on high-fat diets to reach their targets of lipid and protein, respectively. The bees survived best on a 10:1 P:L diet; the risk of dying increased as a function of dietary lipid when bees ate diets with lipid contents greater than 5:1 P:L. Hypothesizing that the P:L intake target of adult worker bumble bees was between 25:1 and 5:1, we presented workers from both species with unbalanced but complementary paired diets to determine whether they self-select their diet to reach a specific intake target. Bees consumed similar amounts of proteins and lipids in each treatment and averaged a 14:1 P:L for B. terrestris and 12:1 P:L for B. impatiens. These results demonstrate that adult worker bumble bees likely select foods that provide them with a specific ratio of P:L. These P:L intake targets could affect pollen foraging in the field and help explain patterns of host-plant species choice by bumble bees. [ABSTRACT FROM AUTHOR]

The aim of this study was to investigate the effects of Elado-dressed winter oilseed rape (OSR, 10 g clothianidin & 2 g beta-cyfluthrin/kg seed) on the development, reproduction and behaviour of large earth bumble bees ( Bombus terrestris) as part of a large-scale monitoring field study in Northern Germany, where OSR is usually cultivated at 25-33 % of the arable land. Both reference and test sites comprised 65 km in which no other crops attractive to pollinating insects were present. Six study locations were selected per site and 10 bumble bee hives were placed at each location. At each site, three locations were directly adjacent to OSR fields and three locations were situated 400 m distant from the nearest OSR field. The development of colonies was monitored from the beginning of OSR flowering in April until June 2014. Pollen from returning foragers was analysed for its composition. An average of 44 % of OSR pollen was found in pollen loads of bumble bees indicating that OSR was a major resource for the colonies. At the end of OSR flowering, hives were transferred to a nature reserve until the end of the study. Colony development in terms of hive weight and the number of workers showed a typical course with no statistically significant differences between the sites. Reproductive output was comparatively high and not negatively affected by the exposure to treated OSR. In summary, Elado-dressed OSR did not cause any detrimental effects on the development or reproduction of bumble bee colonies. [ABSTRACT FROM AUTHOR]

Nutritional deficits may be one factor contributing to the ongoing decline of wild and managed bees. As a consequence, interest in understanding the effect of floral resource availability on nutritional intake - and subsequently bee health and performance - has increased. However, the proximate mechanisms underlying bee foraging choices are still poorly understood. We do, for instance, not yet know whether and how bees assess the nutritional quality of pollen or whether they use information on pollen quality to adjust foraging patterns. In a previous study, we showed that Bombus terrestris workers use their sense of taste to discriminate pollen differing in concentration and, thus, nutrient content. We now investigated whether they translate this ability into foraging behavior by observing microcolonies offered the choice between two feeders containing different pollen types and concentrations and, thus, qualities. We examined whether bumblebees used direct (chemotactile) or indirect (olfactory) cues to assess pollen quality and whether they individually assessed pollen quality or relied on larval feedback. By changing pollen quality after 21 days, we further tested whether information on pollen quality was subsequently associated with other (faster assessable) cues (e.g., odor) or continuously re-assessed. We found that bees always preferred pollen of higher quality and individually assessed quality based on chemotactile cues. Moreover, changing pollen quality resulted in rapid equivalent changes in foraging behavior, even without feedback from larvae. Consequently, individual bumblebee foragers continuously re-assess pollen quality to provide their colonies with high-quality food. Significance statement: The ongoing decline of wild and managed bees is likely driven by several factors with nutritional deficits being one of them. Because bees are amongst the most important pollinators, interest in understanding the effect of floral resource availability on nutritional intake and, subsequently, bee health and performance has increased. However, the proximate mechanisms underlying bee foraging choices are still poorly understood. In this study, we investigated how bumblebees differentiate between different pollen qualities by observing microcolonies offered the choice between different diets. We found that bees always preferred pollen of higher quality and individually assessed quality based on chemotactile cues. This finding increases our understanding of how social bees assess nutritional quality and make appropriate foraging choices. [ABSTRACT FROM AUTHOR]

Stress has complex effects on learning and memory, depending on both the type of stress and when the animal experiences it. Honeybees and bumblebees are agriculturally important pollinators for whom the effects of stress are extremely relevant. These pollinators are often transported long distances during which colonies experience severe physical disturbance, causing stress to individuals prior to their release for pollination. Under natural foraging conditions, bees are excellent at learning about the flowers they forage from, including associations between floral stimuli and rewards. However, it is not clear how stress might affect bees’ abilities to learn and remember floral features. Here we address the effects of acute stress on learning and memory in the bumble bee, Bombus impatiens . Using the Proboscis Extension Response (PER) protocol, we look at stress effects on learning and memory in three experiments. After being trained to a conditioned stimulus, we addressed: (1) the effect of 24 h and (2) 30 min of stress on the recall of this learned association and (3) the effect of stress on subsequently learning an association. We found that 24 h of stress after learning appeared to improve memory recall, and there was a trend in the same direction for 30 min of stress. However, bees that were stressed prior to learning an association did not differ from unstressed bees in their ability to learn or remember an association. Our finding that stress has effects on memory in the bumblebee could have implications for their use in behavioral experiments and for promoting the success of both managed and wild populations. [ABSTRACT FROM AUTHOR]

Ants on flowers can disrupt pollination by consuming rewards or harassing pollinators, but it is difficult to disentangle the effects of these exploitative and interference forms of competition on pollinator behavior. Using highly rewarding and quickly replenishing artificial flowers that simulate male or female function, we allowed bumblebees (Bombus impatiens) to forage (1) on flowers with or without ants (Myrmica rubra) and (2) on flowers with or without ant scent cues. Bumblebees transferred significantly more pollen analogue both to and from ant-free flowers, demonstrating that interference competition with ants is sufficient to modify pollinator foraging behavior. Bees also removed significantly less pollen analogue from ant-scented flowers than from controls, making this the first study to show that bees can use ant scent to avoid harassment at flowers. Ant effects on pollinator behavior, possibly in addition to their effects on pollen viability, may contribute to the evolution of floral traits minimizing ant visitation. [ABSTRACT FROM AUTHOR]

Abstract: The effects of floral herbivores on floral traits may result in alterations in pollinator foraging behaviour and subsequently influence plant reproductive success. Fed-upon plants may have evolved mechanisms to compensate for herbivore-related decreased fecundity. We conducted a series of field experiments to determine the relative contribution of floral herbivores and pollinators to female reproductive success in an alpine herb, Pedicularis gruina, in two natural populations over two consecutive years. Experimental manipulations included bagging, hand supplemental, geitonogamous pollination, and simulated floral herbivory. Bumblebees not only avoided damaged flowers and plants but also decreased successive visits of flowers in damaged plants, and the latter may reduce the level of geitonogamy. Although seed set per fruit within damaged plants was higher than that in intact plants, total seed number in damaged plants was less than that in intact plants, since floral herbivory-mediated pollinator limitation led to a sharp reduction of fruit set. Overall, the results suggest that resource reallocation within inflorescences of damaged plants may partially compensate for a reduction in seed production. Additionally, a novel finding was the decrease in successive within-plant bumblebee visits following floral herbivory. This may increase seed quantity and quality of P. gruina since self-compatible species exhibit inbreeding depression. The patterns of compensation of herbivory and its consequences reported in this study give an insight into the combined effects of interactions between floral herbivory and pollination on plant reproductive fitness. [Copyright &y& Elsevier]

Metals in soil are known to negatively affect the health of many groups of organisms, but it is unclear whether they can affect plant-pollinator interactions, and whether pollinators that visit plants growing on contaminated soils are at risk of ingesting potentially toxic resources. We address whether the presence of metals in nectar alters foraging behavior by bumblebees by manipulating nectar with one of two common soil contaminants (Al or Ni) in flowers of Impatiens capensis (Balsaminaceae). While the presence of Al in nectar did not influence foraging patterns by bumblebees, flowers containing Ni nectar solutions were visited for shorter time periods relative to controls, and discouraged bees from visiting nearby Ni-contaminated flowers. However, because bumblebees still visited these flowers, they likely ingested a potentially toxic resource. Our findings suggest that soil metals could cascade to negatively affect pollinators in metal contaminated environments. [Copyright &y& Elsevier]

In modelling bumble bee foraging, net rate of energetic intake has been suggested as the appropriate currency. The foraging behaviour of honey bees is better predicted by using efficiency, the ratio of energetic gain to expenditure, as the currency. We re-analyse several studies of bumble bee foraging and show that efficiency is as good a currency as net rate in terms of predicting behaviour. We suggest that future studies of the foraging of bumble bees should be designed to distinguish between net rate and efficiency maximizing behaviour in an attempt to discover which is the more appropriate currency. [ABSTRACT FROM AUTHOR]

Seed production may be limited because flowers do not get enough suitable pollen or because plants lack the resources to make seeds. We used replicated plantings to test factors that influence effects of bumblebee behavior on pollen limitation, as measured by the difference in seed set between hand- and naturally pollinated flowers, of Penstemon digitalis in patches of four to 41 flowering individuals. Seed set per flower was 376% higher in the largest as compared with the smallest Penstemon patches. This positive density dependence reflects activity of long-tongued bees, which (1) have higher effective density as patch size increases, (2) visit greater proportions of plants as patch size increases, and (3) visit smaller proportions of flowers per visited plant as patch size increases. Our results suggest that economics of flight and maneuverability of large, long-tongued bumblebees lead them to transfer more pollen between than within Penstemon plants in large patches. Density of smaller, short-tongued bumblebees was not positively associated with Penstemon seed set, but these bees may be important pollinators at low plant densities. Our experimental system indicates a clear positive relationship between activity of effective pollinators and seed set in a species capable of pollinating itself. [ABSTRACT FROM AUTHOR]

An environmental study of pollinator effectiveness of Bumblebees in Agapanthus compared to other insects in the region of Tasmania is presented. White and blue naturalized flowers (Agapanthus) of the area with pollen loads of 9-10 specimens for each of Bumblebees and other pollinator insects were considered for the study. From the product of the number of flower visits, the percentage of visits that involved stigmatic contact, and the mean number of Agapanthus pollen grains per insect type, the pollinator effectiveness score was calculated for each insect type. The study concluded Bumblebee workers/drones were the most common visitors to flowers of Agapanthus, followed by honeybees and other insects.

1. Little is known about the pollinators of rare plants, which is cause for concern given that pollination is essential for the long-term survival of most plant species. The aim of this study was to determine the probable pollinators of three species of rare arable weed: red hemp-nettle Galeopsis angustifolia, small-flowered catchfly Silene gallica and spreading hedge-parsley Torilis arvensis. Species of arable weed are among those suffering the greatest declines in the UK. 2. Five field sites were chosen, two of which were sampled in 2 years. Visitation and pollen transport webs were constructed for the entire plant–pollinator community at each site. Visitation webs described the frequency with which each insect species visited each plant species. Pollen transport webs quantified which insect species transported the pollen of which plant species. 3. A wide range of insect species visited the three plant species. A pollinator importance index was calculated that combined information on both the relative abundance of each insect carrying the pollen of the rare plant and its pollen fidelity. Using this method Galeopsis angustifolia was most likely to be pollinated by Bombus pascuorum at one site and Sphaerophoria scripta at another. Silene gallica was also likely be pollinated by Sphaerophoria scripta. 4. The pollinator fauna of the three plant species varied considerably across their geographical range, but less from one year to the next. 5. Synthesis and applications. All three species of rare plant were linked to other plant species in the community by shared pollinators. In many cases these other plant species constituted the primary food sources for the shared pollinators. Therefore, the long-term survival of rare plant populations is likely to depend on the more common plant species in the community. We recommend that management of the rare plants studied here should also include the protection and management of populations of some of the more common plant species in their respective communities. [ABSTRACT FROM AUTHOR]

Pollinators collect resources that are patchy, since flowers are usually aggregated on several spatial scales. Empirical studies have established that pollinators almost invariably visit a smaller proportion of flowers as patch size increases. This has not been adequately explained. Here I present data on the payoff curve achieved by bumblebees, Bombus lapidarius, when visiting patches containing different numbers of inflorescences, and use the marginal value theorem to predict the optimum duration of stay within patches. The data demonstrate that visiting a declining proportion of inflorescences as patch size increases is an optimal strategy, if we assume that bees are attempting to maximise their rate of reward acquisition. I argue that this occurs because searching for the remaining unvisited inflorescences is easier in a small patch. On large patches, bees visited more inflorescences per patch than predicted (although still visiting a declining proportion). I suggest that this may occur because bees are using simple departure rules which result in near-optimal behaviour. I show that a departure rule based on two successive encounters with empty inflorescences closely predicts observed behaviour. [ABSTRACT FROM AUTHOR]

I examined the response of bumblebees to two kinds of spatial variation and two kinds of temporal variation in nectar levels. The spatial variation involved differences in reward value among plant species and differences in nectar availability among patches of flowers of a single species. The temporal variation involved changes in nectar availability over a season and between years. In all cases the response of bees was measured by differences in bee density or abundance. The response of bees of different plant species was determined by the density of foraging bees on the flowers of each species at a given time (bee:flower ratio = B/F). I examined several groups of plant species whose members overlapped in their flowering periods and in their bumblebee visitors. Within a group, the relative magnitudes of the B/F's for species indicate the degree to which their flowers are preferred by bees. If bees are optimal foragers, preference should reflect the per-flower reward value of species. I characterized the potential reward value of species by their nectar production rates (NPR's). The bees' preference for species was found to be very similar to the relative magnitudes of their NPR's. I also considered whether including the time and energy costs of foraging on the different species would produce a better predictor of preference. While these factors did affect the absolute reward value of species they did not influence their reward value relative to one another. Foraging costs are most likely to have no effect on preference when species have similar floral morphologies. To assess the response of bees to differences in patch quality I bagged some of the inflorescenes in an experimental patch. Following this reduction in the number of flowers, the number of bees in the patch decreased proportionately, producing a B/F very similar to that of a control patch. When the bagging was removed the number of bees increased such that the B/F was again similar to that of the control patch. In general these results provide further support for optimal foraging in bumblebees. I examined the response of bees to temporal variation in nectar levels to determine whether resource limitation for bumblebees change over time. Seasonal changes in the abundances of three bumblebee species were compared with seasonal changes in the total nectar production by the plants each bee visited. Preference factors were used to convert floral production to nectar production. In general, seasonal production and utilization were closely matched, indicating no major change in nectar limitation. The cumulative numbers of bees seen during each of two summer seasons (1974 and 1975) were very similar. This indicates that the resource base, which had not changed between years, was capable of supporting only a limited number of bees. During 1975 a natural species removal experiment occurred which allowed an assessment of competition among bees. Apis mellifera, which had been well represented in 1974, was absent. The bumblebees of short and medium tongue length with which honeybees overlapped in resource used exhibited competitive release. Their abundances increased in 1975 resulting in near-perfect density compensation. [ABSTRACT FROM AUTHOR]

The presence of yeasts in pollen and floral nectar is rather the norm than the exception. Due to the metabolic activities of yeasts, sugar and amino acid composition of nectar often drastically change and may negatively impact the nutritional value of nectar for pollinators and hence insect fitness. On the other hand, the presence of yeasts in floral nectar may also increase its nutritional value due to yeast's probiotic effect and the release of yeast's metabolites. In this study, we investigated whether the presence of defined flower‐ and insect‐associated yeasts affected individual and colony fitness of the bumble bee pollinator Bombus terrestris. Specifically, we tested whether the presence of yeasts in nectar affected bumble bee foraging behavior and nectar consumption, individual growth and colony development, larval and queen mortality, and mating success. Quantitative analyses of sugar and amino acid profiles showed that nectar yeasts significantly affected the chemical composition of nectar. However, dual‐choice experiments indicated that yeast inoculation did not significantly affect foraging behavior or consumption rates. Nest development, on the other hand, was significantly affected by the presence of yeasts, but effects largely depended on species identity, with Candida bombiphila, Metschnikowia gruessii, and Rhodotorula mucilaginosa having the largest positive impact on colony growth. Interestingly, the effects at the colony level were more pronounced than at the individual level. In vitro growth tests further showed that yeasts have the potential to suppress the growth of the bumble bee gut pathogen Crithidia bombi. Overall, these results demonstrate that nectar‐inhabiting yeasts can have diverse effects on bumble bee fitness and therefore may mediate plant–pollinator mutualisms. [ABSTRACT FROM AUTHOR]