Search

A key question in pollination biology is that of how pollinators identify and choose foraging patches. Several approaches have been employed for this, including field studies and large greenhouse flight chambers. Most methods used to date are limited, though, by reliance on a combination of artificial flowers, large spatial scales, or lack of spatially distinct floral patches. To address this issue, we designed and tested a y-maze flight arena and tested it using the bumblebee Bombus impatiens and canola plants. Our results indicate that the system is not biased by environmental conditions, or by an innate "handedness" of Bombus impatiens. We found that bees made all the expected patch choices when presented with soil, plants without flowers, or plants with flowers. This new method is important and useful as it allows researchers to ask questions of both plant health and insect behavior and the chamber system is modular allowing for simple changes to the setup to focus on different questions. Highlights: • Y-maze flight arena was designed to evaluate foraging behavior on intact plants. • No evidence of side preference in individual bees. • Behaviors observed in the y-maze appear to correspond to behaviors observed in other settings. [ABSTRACT FROM AUTHOR]

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]

When foraging in their natural environment, many animals readily complement their personal knowledge with additional social information. To balance the costs and benefits of copying others, animals have to discern situations in which it is more advantageous to use social rather than personal information. Here, we used foraging bumblebees (Bombus terrestris) in a controlled laboratory setting and showed that the difficulty of a foraging task affects how the bees weight the 2 types of information. We used artificial flowers to devise easy and difficult discriminatory tasks, and tested the influence of floral and social cues on decision making. When facing an easy discrimination task, foraging bees were likely to rely on personal information and were only marginally affected by social information. By contrast, they prioritized social over personal information when flower discrimination was difficult and therefore the probability of making errors was higher. In summary, bees are able to use social and personal information to optimize foraging decisions in a flexible way. [ABSTRACT FROM AUTHOR]

In their natural environments, most animals must learn about multiple kinds of rewards, both within and across contexts. Despite this, the majority of research on animal learning involves a single reward type. For example, bees are an important model system for the study of cognition and its ecological consequences, but nearly all research to date on their learning concerns a single reward, nectar (carbohydrates), even though foragers often simultaneously collect pollen (protein). Features of learning under more ecologically realistic conditions involving multiple reward types are thus largely unexplored. To address this gap, we compared performance on a colour-learning task when floral surrogates offered bumblebees, Bombus impatiens , a single type of floral reward versus multiple, nutritionally distinct rewards. In one experiment, bees learned a floral association with nectar either alone or while simultaneously collecting pollen. In a reciprocal experiment, bees learned a floral association with pollen either alone or while simultaneously collecting nectar. Bees that collected pollen while learning about nectar did not suffer any detriment to learning which flower colour offered nectar. However, this was not the case for the reciprocal task: collecting nectar impaired bees' ability to learn and remember associations between floral colour and pollen. Our findings offer new insight into how bees learn in relation to ecologically realistic rewards and how cognitive constraints may shape their behaviour under ecologically realistic foraging scenarios. [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]

Picture-object correspondence provides an alternate method of investigating delayed matching by providing a cue (picture) which may be spontaneously perceived as similar but different from a corresponding target. Memory for, and corresponding choice of, a target corresponding to a cue could be facilitated by the use of a picture. Bumblebees have been found to both easily differentiate images from corresponding objects but also spontaneously perceive a similarity between the two. Herein, an approach was designed to test the possible use of picture cues to signal reward in a delayed matching task. Target choice preference corresponding to picture cues was tested among three bumblebee (Bombus impatiens) colonies using photograph cues (presented prior to target stimuli) corresponding to one of four target stimuli. Photograph cues were the only predictor of corresponding target reward, presented in stable locations. Rewarded and unrewarded tests show a choice preference significantly higher than chance for targets matching the cue. Results suggest that bumblebees can learn to use picture cues in a delayed matching task. Furthermore, experience, conditions of reward inconsistency and location, are discussed as possible contributing factors to learning in a delayed matching task. [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]

Bee species interactions can benefit plant pollination through synergistic effects and complementary effects, or can be of detriment to plant pollination through competition effects by reducing visitation by effective pollinators. Since specific bee interactions influence the foraging performance of bees on flowers, they also act as drivers to regulate the assemblage of flower visitors. We selected squash (Cucurbita pepo L.) and its pollinators as a model system to study the foraging response of honey bees to the occurrence of bumble bees at two types of sites surrounded by a high amount of natural habitats (≥ 58% of land cover) and a low amount of natural habitats (≤ 12% of land cover) in a highland agricultural ecosystem in China. At the individual level, we measured the elapsed time from the departure of prior pollinator(s) to the arrival of another pollinator, the selection of honey bees for flowers occupied by bumble bees, and the length of time used by honey bees to explore floral resources at the two types of sites. At the community level, we explored the effect of bumble bee visitation on the distribution patterns of honey bees on squash flowers. Conclusively, bumble bee visitation caused an increase in elapsed time before flowers were visited again by a honey bee, a behavioral avoidance by a newly-arriving honey bee to select flowers occupied by bumble bees, and a shortened length of time the honey bee takes to examine and collect floral resources. The number of overall bumble bees on squash flowers was the most important factor explaining the difference in the distribution patterns of honey bees at the community level. Furthermore, decline in the number of overall bumble bees on the squash flowers resulted in an increase in the number of overall honey bees. Therefore, our study suggests that bee interactions provide an opportunity to enhance the resilience of ecosystem pollination services against the decline in pollinator diversity. [ABSTRACT FROM AUTHOR]

The learning capacities of males and females may differ with sex-specific behavioural requirements. Bumblebees provide a useful model system to explore how different lifestyles are reflected in learning abilities, because their (female but sterile) workers and males engage in fundamentally different behaviour routines. Bumblebee males, like workers, embark on active flower foraging but in contrast to workers they have to trade off their feeding with mate search, potentially affecting their abilities to learn and utilize floral cues efficiently during foraging. We used a serial colour-learning task with freely flying males and workers to compare their ability to flexibly learn visual floral cues with reward in a foraging scenario that changed over time. Male bumblebees did not differ from workers in both their learning speed and their ability to overcome previously acquired associations, when these ceased to predict reward. In all foraging tasks we found a significant improvement in choice accuracy in both sexes over the course of the training. In both sexes, the characteristics of the foraging performance depended largely on the colour difference of the two presented feeder types. Large colour distances entailed fast and reliable learning of the rewarding feeders whereas choice accuracy on highly similar colours improved significantly more slowly. Conversely, switching from a learned feeder type to a novel one was fastest for similar feeder colours and slow for highly different ones. Overall, we show that behavioural sex dimorphism in bumblebees did not affect their learning abilities beyond the mating context. We discuss the possible drivers and limitations shaping the foraging abilities of males and workers and implications for pollination ecology. We also suggest stingless male bumblebees as an advantageous alternative model system for the study of pollinator cognition. [ABSTRACT FROM AUTHOR]

In contrast to the wealth of knowledge concerning sucrose-rewarded learning, the question of whether bees learn when they collect pollen from flowers has been little addressed. The nutritional value of pollen varies considerably between species, and it may be that bees learn the features of flowers that produce pollen best suited to the dietary requirements of their larvae. It is still unknown, however, whether a non-ingestive reward pathway for pollen learning exists, and how foraging bees sense differences between pollen types. Here we adopt a novel experimental approach testing the learning ability of bees with pollen rewards. Bumblebees were reared under controlled laboratory conditions. To establish which pollen rewards are distinguishable, individual bees were given the choice of collecting two types of pollen, diluted to varying degrees with indigestible a- cellulose. Bees preferentially collected a particular pollen type, but this was not always the most concentrated sample. Preferences were influenced by the degree of similarity between samples and also by the period of exposure, with bees more readily collecting samples of lower pollen concentration after five trials. When trained differentially, bees were able to associate an initially less-preferred contextual colour with the more concentrated sample, whilst their pollen preferences did not change. Successful learning of contextual cues seems to maintain pollen foraging preferences over repeated exposures, suggesting that fast learning of floral cues may preclude continuous sampling and evaluation of alternative reward sources, leading to constancy in pollen foraging. [ABSTRACT FROM AUTHOR]

If the cognitive performance of animals reflects their particular ecological requirements, how can we explain appreciable variation in learning ability amongst closely related individuals (e.g. foraging workers within a bumble bee colony)? One possibility is that apparent 'errors' in a learning task actually represent an alternative foraging strategy. In this study we investigate the potential relationship between foraging 'errors' and foraging success among bumble bee ( Bombus terrestris) workers. Individual foragers were trained to choose yellow, rewarded flowers and ignore blue, unrewarded flowers. We recorded the number of errors (visits to unrewarded flowers) each bee made during training, then tested them to determine how quickly they discovered a more profitable food source (either familiar blue flowers, or novel green flowers). We found that error prone bees discovered the novel food source significantly faster than accurate bees. Furthermore, we demonstrate that the time taken to discover the novel, more profitable, food source is positively correlated with foraging success. These results suggest that foraging errors are part of an 'exploration' foraging strategy, which could be advantageous in changeable foraging environments. This could explain the observed variation in learning performance amongst foragers within social insect colonies. [ABSTRACT FROM AUTHOR]

Given that many pollinators have exhibited dramatic declines related to habitat destruction, an improved understanding of pollinator resource collection across human-altered landscapes is essential to conservation efforts. Despite the importance of bumble bees (Bombus spp.) as global pollinators, little is known regarding how pollen collection patterns vary between individuals, colonies, and landscapes. In this study, Vosnesensky bumble bees (Bombus vosnesenskii Radoszkowski) were collected from a range of human-altered and natural landscapes in northern California. Extensive vegetation surveys and Geographic Information System (GIS)-based habitat classifications were conducted at each site, bees were genotyped to identify colony mates, and pollen loads were examined to identify visited plants. In contrast to predictions based on strong competitive interactions, pollen load composition was significantly more similar for bees captured in a shared study region compared with bees throughout the research area but was not significantly more similar for colony mates. Preference analyses revealed that pollen loads were not composed of the most abundant plant species per study region. The majority of ranked pollen preference lists were significantly correlated for pairwise comparisons of colony mates and individuals within a study region, whereas the majority of pairwise comparisons of ranked pollen preference lists between individuals located at separate study regions were uncorrelated. Results suggest that pollen load composition and foraging preferences are similar for bees throughout a shared landscape regardless of colony membership. The importance of native plant species in pollen collection is illustrated through preference analyses, and we suggest prioritization of specific rare native plant species for enhanced bumble bee pollen collection. [ABSTRACT FROM AUTHOR]

Background: Bumblebees use information provided inadvertently by conspecifics when deciding between different flower foraging options. Such social learning might be explained by relatively simple associative learning mechanism: the bee may learn to associate conspecifics with nectar or pollen reward through previous experience of foraging jointly. However, in some studies, observers were guided by choices of ‘demonstrators’ viewed through a screen, so no reward was given to the observers at the time of seeing other bees’ flowers choice and no demonstrator bee was present at the moment of decision. This behaviour, referred to observational conditioning, implies an additional associative step as the positive value of conspecific is transferred to the associated flower. Here we explore the role of demonstrator movement, and the distance between observers and demonstrators that is required for observation conditioning to take place. Methodology/Principal Findings: We identify the conditions under which observational conditioning occurs in the widespread European species Bombus terrestris. The presence of artificial demonstrator bees leads to a significant change in individual colour preference toward the indicated colour if demonstrators were moving and observation distance was limited (15 cm), suggesting that observational conditioning could only influence relatively short-range foraging decisions. In addition, the movement of demonstrators is a crucial factor for observational conditioning, either due to the more life-like appearance of moving artificial bees or an enhanced detectability of moving demonstrators, and an increased efficiency at directing attention to the indicated flower colour. Conclusion: Bumblebees possess the capacity to learn the quality of a flower by distal observation of other foragers’ choices. This confirms that social learning in bees involves more advanced processes than simple associative learning, and indicates that observational conditioning might be widespread in pollinating insects, raising intriguing questions for the underlying mechanisms as well as the spread of social information in pollinator-plant interactions. [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]

Highlights: [•] Bombus terrestris foragers produce larger amounts of octyl esters compared to non-foragers. [•] Octyl esters are also known as a signal to sterility that reduces aggression. [•] Ester production is positively correlated with worker foraging efforts. [•] Foraging, potential fecundity and aggression are interlinked in B. terrestris workers. [Copyright &y& Elsevier]

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]

Despite the ubiquity and the importance of interspecific interactions among flower visitors, few studies have examined their effects on the realized feeding niches of visitor species in a community context. To evaluate the community-wide effects of interactions among flower visitors, I have examined changes in the flower utilization patterns of each visitor species at several sites where the component of the visitor's community differed. Specifically, I compared the flower preferences and foraging habits (legitimate foraging vs. primary nectar robbing vs. secondary nectar robbing) of five bumble bee species in flower patches consisting of Trifolium pratense L. (red clover) and T. repens L. (white clover) on Hokkaido Island, Japan. I also examined the nectar production and standing crops of each flower species to evaluate the exploitation competition based on nectar. The bumble bee species exhibited different flower utilization patterns among sites. At sites where the long-tongued Bombus diversus tersatus was common and the exotic short-tongued B. terrestris was rare, B. diversus tersatus visited red clover (long-tubed flowers) exclusively, whereas medium-tongued B. pseudobaicalensis and short-tongued B. hypocrita sapporoensis and B. hypnorum koropokkrus preferentially visited white clover (short-tubed flowers). Conversely, at sites where the long-tongued bee was rare, four other species frequently visited red clover in different modes: B. pseudobaicalensis visited legitimately, B. hypocrita sapporoensis and B. terrestris visited as primary nectar robbers, and B. hypnorum koropokkrus visited as a secondary nectar robber. The presence or absence of resource exploitation by the long-tongued species and the interaction between primary and secondary nectar robbers via robbing holes was the major ecological sources of these differences. Diverse effects of interactions among flower visitors played important roles in shaping pattern of plant and flower visitor interactions. [ABSTRACT FROM AUTHOR]

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]

The capacity to discriminate between choice options is crucial for a decision-maker to avoid unprofitable options. The physical properties of rewards are presumed to be represented on context-dependent, nonlinear cognitive scales that may systematically influence reward expectation and thus choice behavior. In this study, we investigated the discrimination performance of free-flying bumblebee workers ( Bombus impatiens) in a choice between sucrose solutions with different concentrations. We conducted two-alternative free choice experiments on two B. impatiens colonies containing some electronically tagged bumblebees foraging at an array of computer-automated artificial flowers that recorded individual choices. We mimicked natural foraging conditions by allowing uncertainty in the probability of reward delivery while maintaining certainty in reward concentration. We used a Bayesian approach to fit psychometric functions, relating the strength of preference for the higher concentration option to the relative intensity of the presented stimuli. Psychometric analysis was performed on visitation data from individually marked bumblebees and pooled data from unmarked individuals. Bumblebees preferred the more concentrated sugar solutions at high stimulus intensities and showed no preference at low stimulus intensities. The obtained psychometric function is consistent with reward evaluation based on perceived concentration contrast between choices. We found no evidence that bumblebees reduce reward expectations upon experiencing non-rewarded visits. We compare psychometric function parameters between the bumblebee B. impatiens and the flower bat Glossophaga commissarisi and discuss the relevance of psychophysics for pollinator-exerted selection pressures on plants. [ABSTRACT FROM AUTHOR]

Animals collecting food from renewable resource patches scattered in space often establish small foraging areas to which they return faithfully. Such area fidelity offers foraging advantages through selection of profitable patches, route minimization, and regular circuit visits to these patches (“trapline foraging”). Resource distribution under field conditions may often vary in time, however, especially when competitors suddenly vanish and a number of patches become available for their neighbors. Previous studies suggested that site-faithful foragers of bumble bees quickly respond to such unexpected events by readjusting their foraging areas, although it is not clear how much their foraging performance was improved, beyond the simple relaxation of competitive pressure, or how they manifest such flexibility while persistently using certain foraging areas or paths. Here, we conducted indoor flight-cage experiments with bumble bees and found that a bee, when encountering a loss of its competitor, improved its foraging performance to a greater extent than expected from a simple relaxation of competitive pressure by increasing the size of its foraging area. Moreover, bees with better-established traplines achieved greater foraging areas after the loss of competitors, suggesting that bees do not necessarily need to “sample” neighboring patches to monitor temporal changes in environments. We discuss how periodical returns and route memory associated with accurate reward values could allow inherently conservative trapliners to make flexible adjustments, by effectively monitoring their circumstances and quickly readjusting to detected changes. [ABSTRACT FROM AUTHOR]

Bumblebees tend to face their nest over a limited range of compass directions when learning the nest's location on departure and finding it on their approach after foraging. They thus obtain similar views of the nest and its surroundings on their learning and return flights. How do bees coordinate their flights relative to nest-based and compass-based reference frames to get such similar views? We show, first, that learning and return flights contain straight segments that are directed along particular compass bearings, which are independent of the orientation of a bee's body. Bees are thus free within limits to adjust their viewing direction relative to the nest, without disturbing flight direction. Second, we examine the coordination of nest-based and compass-based control during likely information gathering segments of these flights: loops during learning flights and zigzags on return flights. We find that bees tend to start a loop or zigzag when flying within a restricted range of compass directions and to fly towards the nest and face it after a fixed change in compass direction, without continuous interactions between their nest-based and compass-based directions of flight. A preferred trajectory of compass-based flight over the course of a motif, combined with the tendency of the bees to keep their body oriented towards the nest automatically narrows the range of compass directions over which bees view the nest. Additionally, the absence of interactions between the two reference frames allows loops and zigzags to have a stereotyped form that can generate informative visual feedback. [ABSTRACT FROM AUTHOR]

Copyright of Journal of Economic Entomology is the property of Oxford University Press / USA and its content may not be copied or emailed to multiple sites or posted to a listserv without the copyright holder's express written permission. However, users may print, download, or email articles for individual use. This abstract may be abridged. No warranty is given about the accuracy of the copy. Users should refer to the original published version of the material for the full abstract. (Copyright applies to all Abstracts.)

While there has been considerable research on the behavioral processes that underlie animals' ability respond to shifting rewards, it remains unclear how animals coordinate multiple processes over time. To investigate this, we compared the behavior of honeybees ( Apis mellifera) and bumblebees ( Bombus impatiens), in an open-ended search task. Bees were given brief access to a high-quality food source, which then became non-rewarding. Then, over an extended period, we examined (1) bees' tendency to persist at the depleted site, (2) their tendency to return to a different low-quality food source where they had been foraging previously, (3) their tendency to return to the hive, and (4) how previous reward history influenced their tendency to shift among these options. Compared to bumblebees, honeybees were much slower to abandon the depleted site and were much more likely to make trips to the hive while bumblebees were much more likely to return to the familiar low-quality site. These observed species differences are interpreted in terms of evolved individual and social differences between these species. We show evidence of well-studied behavioral processes such as extinction, negative contrast effects, and reliance on a social group, and provide, for the first time, a picture of how these processes interact with one another as part of a common sequential decision-making process. [ABSTRACT FROM AUTHOR]

Conspecifics are usually considered competitors negatively affecting food intake rates. However, their presence can also inform about resource quality by providing inadvertent social information. Few studies have investigated whether foragers perceive conspecifics as informers or competitors. Here, we experimentally tested whether variation in the density of demonstrators ('none', 'low' and 'high'), whose location indicated flower profitability, affected decision-making of bumble-bees Bombus terrestris. Bumble-bees foraged on either 'simple' (two colours) or 'complex' (four colours) artificial floral communities. We found that conspecifics at low density may be used as sources of information in first flower choices, whereas they appeared as competitors over the whole foraging sequence. Low conspecific densities improved foragers' first-visit success rate in the simple environment, and decreased time to first landing, especially in the complex environment. High conspecific densities did not affect these behavioural parameters, but reduced flower constancy in both floral communities, which may alter the efficiency of pollinating visits. These results suggest that the balance of the costs and benefits of conspecific presence varies with foraging experience, floral community and density. Spatio-temporal scales could thus be an important determinant of social information use. This behavioural flexibility should allow bumble-bees to better exploit their environment. [ABSTRACT FROM AUTHOR]

In the absence of predators, pollinators can often maximize their foraging success by visiting the most rewarding flowers. However, if predators use those highly rewarding flowers to locate their prey, pollinators may benefit from changing their foraging preferences to accept less rewarding flowers. Previous studies have shown that some predators, such as crab spiders, indeed hunt preferentially on the most pollinator-attractive flowers. In order to determine whether predation risk can alter pollinator preferences, we conducted laboratory experiments on the foraging behavior of bumble bees ( Bombus impatiens) when predation risk was associated with a particular reward level (measured here as sugar concentration). Bees foraged in arenas containing a choice of a high-reward and a low-reward artificial flower. On a bee's first foraging trip, it was either lightly squeezed with forceps, to simulate a crab spider attack, or was allowed to forage safely. The foragers' subsequent visits were recorded for between 1 and 4 h without any further simulated attacks. Compared to bees that foraged safely, bees that experienced a simulated attack on a low-reward artificial flower had reduced foraging activity. However, bees attacked on a high-reward artificial flower were more likely to visit low-reward artificial flowers on subsequent foraging trips. Forager body size, which is thought to affect vulnerability to capture by predators, did not have an effect on response to an attack. Predation risk can thus alter pollinator foraging behavior in ways that influence the number and reward level of flowers that are visited. [ABSTRACT FROM AUTHOR]

Background: Accurate estimates of movement behavior and distances travelled by animals are difficult to obtain, especially for small-bodied insects where transmitter weights have prevented the use of radio-tracking. Methodology/Principal Findings: Here, we report the first successful use of micro radio telemetry to track flight distances and space use of bumblebees. Using ground surveys and Cessna overflights in a Central European rural landscape mosaic we obtained maximum flight distances of 2.5 km, 1.9 km and 1.3 km for Bombus terrestris (workers), Bombus ruderatus (worker), and Bombus hortorum (young queens), respectively. Bumblebee individuals used large areas (0.25-43.53 ha) within one or a few days. Habitat analyses of one B. hortorum queen at the landscape scale indicated that gardens within villages were used more often than expected from habitat availability. Detailed movement trajectories of this individual revealed that prominent landscape structures (e.g. trees) and flower patches were repeatedly visited. However, we also observed long (i.e. .45 min) resting periods between flights (B. hortorum) and differences in flower-handling between bumblebees with and without transmitters (B. terrestris) suggesting that the current weight of transmitters (200 mg) may still impose significant energetic costs on the insects. Conclusions/Significance: Spatio-temporal movements of bumblebees can now be tracked with telemetry methods. Our measured flight distances exceed many previous estimates of bumblebee foraging ranges and suggest that travelling long distances to food resources may be common. However, even the smallest currently available transmitters still appear to compromise flower handling performance and cause an increase in resting behavior of bees. Future reductions of transmitter mass and size could open up new avenues for quantifying landscape-scale space use of insect pollinators and could provide novel insights into the behavior and requirements of bumblebees during critical life stages, e.g. when searching for mates, nest locations or hibernation sites. [ABSTRACT FROM AUTHOR]

the distribution of food resources shifts over time, central place foragers are likely to be repeatedly faced with the question of when to abandon a forage site that is declining in value and to subsequently search elsewhere. Although there has been a great deal of investigation into how individual foragers allocate time between exploration and exploitation, few studies have sought to explore this issue within a larger functional context. We take a comparative approach to this problem by examining decision making in individual honeybees and bumblebees as they responded to a downshift in food reward. Our results show not only that honeybees and bumblebees have significantly divergent strategies with regards to abandoning a food source that is declining in value but also in terms of the subsequent tendency to seek an alternative food source. We interpret these results in terms of both biological and social distinctions between these species and highlight how group-level characteristics are likely to shape the evolutionarily derived foraging strategies of individual animals. [ABSTRACT FROM AUTHOR]

Preference for certain types of flowers in bee species may be an adaptation for efficient foraging, and they often prefer flowers whose shape fits their mouthparts. However, it is unclear whether such flowers are truly beneficial for them. We address this issue by experimentally measuring foraging efficiency of bumblebees, the volume of sucrose solution consumed over handling time (µL/second), using long-tongued Bombus diversus Smith and short-tongued B. honshuensis Tkalcu that visit Clematis stans Siebold et Zuccarini. The corolla tube length of C. stans decreases during a flowering period, and male flowers are longer than female flowers. Long- and short-tongued bumblebees frequently visited longer and shorter flowers, respectively. Based on these preferences, we hypothesized that bumblebee foraging efficiency is higher when visiting flowers that show a good morphological fit between the proboscis and the corolla tube. Foraging efficiency of bumblebees was estimated using flowers for which nectar quality and quantity were controlled, in an experimental enclosure. We show that 1) the foraging efficiency of B. diversus was enhanced when visiting younger, longer flowers, and that 2) the foraging efficiency of B. honshuensis was higher when visiting shorter female flowers. This suggests that morphological correspondence between insects and flowers is important for insect foraging efficiency. However, in contradiction to our prediction, 3) short-tongued bumblebees B. honshuensis sucked nectar more efficiently when visiting younger, longer flowers, and 4) there was no significant difference in the foraging efficiency of B. diversus between flower sexes. These results suggest that morphological fit between the proboscis and the corolla tube is not a sole determinant of foraging efficiency. Bumblebees may adjust their sucking behavior in response to available rewards, and competition over rewards between bumblebee species might change visitation patterns in the wild. Thus, the determinants of foraging efficiency and visitation frequency for bee pollinators may be more complex than previously thought. [ABSTRACT FROM AUTHOR]

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]

Brain plasticity is a common phenomenon across animals and in many cases it is associated with behavioral transitions. In social insects, such as bees, wasps and ants, plasticity in a particular brain compartment involved in multisensory integration (the mushroom body) has been associated with transitions between tasks differing in cognitive demands. However, in most of these cases, transitions between tasks are age-related, requiring the experimental manipulation of the age structure in the studied colonies to distinguish age and experience-dependent effects. To better understand the interplay between brain plasticity and behavioral performance it would therefore be advantageous to study species whose division of labor is not age-dependent. Here, we focus on brain plasticity in the bumblebee Bombus occidentalis, in which division of labor is strongly affected by the individual’s body size instead of age. We show that, like in vertebrates, body size strongly correlates with brain size. We also show that foraging experience, but not age, significantly correlates with the increase in the size of the mushroom body, and in particular one of its components, the medial calyx. Our results support previous findings from other social insects suggesting that the mushroom body plays a key role in experience-based decision making. We also discuss the use of bumblebees as models to analyze neural plasticity and the association between brain size and behavioral performance. Copyright © 2010 S. Karger AG, Basel [ABSTRACT FROM AUTHOR]

The article presents a study which investigates the effect of Bacillus thuringiensis (Bt) on foraging behaviour, reproduction, and survival in bumblebee Bombus terrestis (L.) using two Bt strains and laboratory microcolonies. It reveals that the strains did not affect the insects' foraging behaviour, reproduction, and survival, but they have detrimental affects when put in feeding sugar water. It concludes that the strains are safe to the bees, although they have damaging effects in some cases.

A review of two articles "Foraging Bumble Bees Weigh the Reliability of Personal & Social Information" by A. S. Dunlap, M. E Nielsen, A. Dornhaus from periodical Current Biology and "Copy-When-Uncertain: Bumblebees Rely on Social Information When Rewards are Highly Variable" by M. Smolla, S. Alem and L. Chittka from periodical Biology Letters is presented.

Commercial bumblebee colonies are important crop pollinators. Here we assess whether application of artificial foraging recruitment pheromone can increase foraging activity in Bombus terrestris colonies on a relevant timescale for commercial pollination. We measured bee traffic from the nest to a foraging arena, which is correlated with foraging activity under natural recruitment conditions. During continuous pheromone exposure bee traffic increased by 1.5 to 3.6 times, and this increase lasted up to 105?minute. Repeated 20 minute exposures of a colony to recruitment pheromone, with at least 30 minutes intermissions, triggered consistent traffic increases over a four week period. We conclude that artificial recruitment pheromone can reliably boost bee traffic leaving previously inactive colonies. This method could improve foraging activity and pollination in greenhouse colonies, especially young colonies reluctant to start foraging after introduction to the crop. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The effect of nectar robbing on plant fitness is poorly understood and restricted to a few plant species. Furthermore, the available studies generally evaluate the effects of nectar robbing on female fitness, disregarding the male component. Here we measured the effects of the nectar-robbing bumblebees on male (measured as pollen analogue flow distance) and female (measured as seed production) reproductive success in the insect-dependent Polygala vayredae, a narrow endemic species from the pre-Pyrenees (Spain). Intense nectar robbing by bumblebees significantly reduced the nectar available to legitimate pollinators in the studied population, and this reduction affected both male and female fitness. Significant differences were observed in fluorescent dye dispersion between robbed and non-robbed flowers within the population. Fluorescent dyes from non-robbed flowers were dispersed to larger distances and over a larger number of flowers when compared with robbed ones. Moreover, significant differences were observed in both fruit set and seed ovule ratios between the two groups, with non-robbed flowers presenting higher reproductive outcomes. However, no effect on seed weight was detected among treatments. The data obtained suggest that in this species, nectar robbing has important indirect and negative effects on plant fecundity, through both male and female functions, due to a modification in the foraging behaviour of legitimate visitors. [ABSTRACT FROM AUTHOR]

We lack a mechanism that links wing wear with mortality in foraging social insects. This study tests the hypothesis that wing wear strongly degrades foraging flight performance, thereby providing a biomechanical explanation for the wing wear - mortality relationship. We examine the effect of simulated wing wear - wing area reduction and asymmetry - on the flight behaviour of bumble bee (Bombus flavifrons Cresson, 1863) workers moving between vertically oriented flowers spaced 30 cm apart and arranged in a two-dimensional horizontal grid. Flight behaviour was measured in three dimensions as total flying distance, mean velocity, variability of velocity, maximum acceleration, maximum deceleration, percentage of time spent accelerating, and displacement from a straight line path between flowers. Loss of wing area had surprisingly little effect on flight behaviour. Viewed multivariately, bees with low asymmetry and low loss of mean area, or with high asymmetry and high loss of mean area, differed from the other three treatment groups. When bees were burdened with both high asymmetry and high loss of wing area, their between-flower flight path was less direct. Overall, flight behaviour of bumble bees was highly resilient to major changes in wing area and asymmetry in this simple foraging environment. The wing wear-associated causes of increased mortality remain elusive. Il n’existe pas de mécanisme pour relier l’usure des ailes à la mortalité chez les insectes sociaux qui font des quêtes alimentaires. Notre étude teste l’hypothèse selon laquelle l’usure des ailes réduit considérablement la performance des vols de recherche de nourriture, ce qui fournit une explication biomécanique de la relation entre l’usure des ailes et la mortalité. Nous examinons l’effet d’une usure simulée des ailes - par réduction de la taille et asymétrie des ailes - sur le comportement de vol d’ouvrières de bourdon (Bombus flavifrons Cresson, 1863) qui se déplacent entre des fleurs à orientation verticale placées à 30 cm les unes des autres selon une grille horizontale bidimensionnelle. Le comportement vol a été évalué en trois dimensions d’après les variables suivantes: distance totale de vol, vitesse moyenne, variabilité de la vitesse, accélération maximale, décélération maximale, % du temps passé à l’accélération et déviation du trajet de la ligne droite entre les fleurs. La perte de surface alaire a étonnamment peu d’effet sur le comportement de vol. Dans une perspective multidimensionnelle, les abeilles qui possèdent une asymétrie et une perte de surface moyenne faibles et celles qui ont une asymétrie et une perte de surface moyenne fortes diffèrent des trois autres groupes expérimentaux. Lorsque les abeilles sont affligées à la fois par une forte asymétrie et une perte importante de surface alaire, leur trajet de vol entre les fleurs est moins direct. En somme, le comportement de vol des bourdons est très résilient aux changements importants de surface alaire et d’asymétrie dans cet environnement simple de recherche de nourriture. Les causes de mortalité accrue reliées à l’usure des ailes restent toujours difficiles à cerner. [ABSTRACT FROM AUTHOR]

Foraging bumblebees scent mark flowers with hydrocarbon secretions. Several studies have found these scent marks act as a repellent to bee foragers. This was thought to minimize the risk of visiting recently depleted flowers. Some studies, however, have found a reverse, attractive effect of scent marks left on flowers. Do bees mark flowers with different scents, or could the same scent be interpreted differently depending on the bees’ previous experience with reward levels in flowers? We use a simple experimental design to investigate if the scent marks can become attractive when bees forage on artificial flowers that remain rewarding upon the bees’ return after having depleted them. We contrast this with bees trained in the more natural scenario where revisits to recently emptied flowers are unrewarding. The bees’ association between scent mark and reward value was tested with flowers scent marked from the same source. We find that the bees’ experience with the level of reward determines how the scent mark is interpreted: the same scent can act as both an attractant and a repellent. How experience and learning influence the interpretation of the meaning of chemical signals deposited by animals for communication has rarely been investigated. [ABSTRACT FROM AUTHOR]

The spatial distribution of foraging bumblebees, Bombus terrestris L. (Hymenoptera: Apidae), was studied in a greenhouse planted with two cultivars of tomato, Lycopersicum esculentum Mill. (Solanaceae), in two patches. In both patches, bumblebee densities per square meter were measured on plots, and the results showed that their densities were nearly similar. The densities of available flowers, their pollen production, and availability also were measured. Our results showed that, although the cultivars greatly differed in flower density, flower morphology, and pollen production, their pollen availability (i.e., pollen actually collected by bumblebees per square meter) was approximately the same. Therefore, the mean quantities of pollen collected per bumblebee were similar in each patch. Knowing that bumblebees do not visit different varieties randomly, our results suggest that the major factor affecting the bumblebee distribution among patches was the density of available resource. Results are discussed both from an applied point of view and in relation to the assumptions of the ideal free distribution theory. [ABSTRACT FROM AUTHOR]

We examined the effects of floral reward level and spatial arrangement on the propensity of bumble bees to exhibit flower constancy. In three separate experiments, we compared the flower constancy of bees on dimorphic arrays of blue and yellow flowers that differed either in reward concentration, reward volume, or inter-flower distance. Overall, flower choice patterns varied among bees, ranging from random selection to complete constancy. When flowers contained greater reward volumes and were spaced farther apart, bees showed less flower constancy and more moves to closely neighbouring flowers. Changes in reward concentration had no effect on flower constancy; however, more dilute rewards produced shorter flight times between flowers. In addition, there was a strong positive relationship between degree of flower constancy and net rate of energy gain when flowers were spaced farther apart, indicating that constant bees were more economic foragers than inconstant bees. Together, these results support the view that the flower constancy of pollinators reflects an economic foraging decision. [ABSTRACT FROM AUTHOR]

Bumblebees can avoid recently depleted flowers by responding to repellent scent-marks deposited on flower corollas by previous visitors. It has previously been suggested that avoidance of visited flowers for a fixed period would be a poor strategy, since different plant species vary greatly in the rate at which they replenish floral rewards. In this study, we examined the duration of flower repellency after an initial bumblebee visit, using wild bumblebees ( Bombus lapidarius, B. pascuorum and B. terrestris) foraging on four different plant species ( Lotus corniculatus, Melilotus officinalis, Phacelia tanacetifolia and Symphytum officinale). We constructed a model to predict flower visitation following an initial visit, based on the nectar secretion pattern of the different plant species, the insect visitation rate per flower, and the search and handling times of bumblebees foraging on the plant species in question. The model predicts an optimal duration of flower avoidance which maximises the rate of reward acquisition for all bees. However, this optimum may be open to cheating. For two plant species, the evolutionary stable strategy (ESS) is a shorter duration of flower avoidance than the optimum. We found the duration of flower avoidance was markedly different among flower species and was inversely related to nectar secretion rates. The predicted ESSs for each plant species were close to those observed, suggesting that the key parameters influencing bumblebee behaviour are those included in the model. We discuss how bees may alter the duration of their response to repellent scents, and other factors that affect flower re-visitation. [ABSTRACT FROM AUTHOR]

The return of a successful bumblebee forager stimulates nestmates to leave the nest and search for food. Here we investigate the mechanisms by which this happens. Successful Bombus terrestris foragers perform irregular runs in their nest, often lasting for several minutes. Run duration is at its maximum when food has just been discovered. Running likely serves to distribute a pheromone, since the information flow between "runners" and "recruits" can be disrupted by eliminating air exchange, while leaving other potential means of communication intact. In addition, nectar stores in the nest may be monitored continuously. A sudden influx of nectar into the nest also causes measurable increases in forager activity. The implications of bumblebee recruitment behavior for the evolution of communication in bees are discussed. [ABSTRACT FROM AUTHOR]

Examines the performance of social species of bumblebee, Bombus bimaculatus, and solitary species of carpenter bee, Xylocopa virginica, in studies in which bees had to learn to discriminate between rewarding and non-rewarding flowers that differed in color. Similar levels of overnight memory retention in both species; Suggestion that social bees have better learning capabilities compared with solitary bees.

Summary. Foraging bumblebees (Bombus vosnesenskii) deposit a substance on rewarding flowers which assists in discrimination between rewarding and non-rewarding flowers in a controlled laboratory environment. Discrimination occurs while the bee is on a flower; workers probe rewarding flowers as well as empty ones that have rewarded in the recent past, but they do not probe flowers that have had no reward. Recognition is not the result of honey contamination left on the flower by the bee during feeding. The deposit is only slightly soluble in water or ethyl alcohol but is very soluble in pentane. [ABSTRACT FROM AUTHOR]

Summary. 1. When presented with a patch of 3 different grass species' seeds, a Pogonomyrmex rugosus colony harvested a mix of all species available. Individually marked workers, however, specialized on a single species. These individual specializations hold for at least several days but can also be shifted rapidly. Workers of a similar species, Veromessor pergandei, also appear to specialize to seed species, taking larger than average seeds of one species from a mixed seed patch when seeds of a larger species were available and being collected by colony-mates in that same patch. 2. Seed choice in both ants is not a function of worker size; this variable explains <4% of the variance in seed size harvested by either ant species. Foraging specializations may enhance individual and colony-wide foraging efficiency in a manner similar to that of "majoring" in bumblebees. [ABSTRACT FROM AUTHOR]

Declines in bumblebee populations have led to investigations into potential causes – including agrochemical effects on bumblebee physiology. The indirect effects of agrochemicals (i.e. behavior modulation) have been postulated, but rarely directly tested. Olfactory information is critical in mediating bumblebee-floral interactions. As agrochemicals emit volatiles, they may indirectly modify foraging behavior. We tested the effects of olfactory contamination of floral odor by agrochemical scent on foraging activity of Bombus impatiens using two behavioral paradigms: localization of food within a maze and forced-choice preference. The presence of a fungicide decreased bumblebees’ ability to locate food within a maze. Additionally, bumblebees preferred to forage in non-contaminated feeding chambers when offered a choice between control and either fertilizer- or fungicide-scented chambers. [ABSTRACT FROM AUTHOR]