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Flowers act as multisensory billboards to pollinators by using a range of sensory modalities such as visual patterns and scents. Different floral organs release differing compositions and quantities of the volatiles contributing to floral scent, suggesting that scent may be patterned within flowers. Early experiments suggested that pollinators can distinguish between the scents of differing floral regions, but little is known about how these potential scent patterns might influence pollinators.We show that bumblebees can learn different spatial patterns of the same scent, and that they are better at learning to distinguish between flowers when the scent pattern corresponds to a matching visual pattern. Surprisingly, once bees have learnt the spatial arrangement of a scent pattern, they subsequently prefer to visit novel unscented flowers that have an identical arrangement of visual marks, suggesting that multimodal floral signals may exploit the mechanisms by which learnt information is stored by the bee. [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]

Summary Iridescence is a form of structural coloration, produced by a range of structures, in which hue is dependent on viewing angle [ 1–4 ]. One of these structures, the diffraction grating, is found both in animals (for example, beetles [ 2 ]) and in plants (on the petals of some animal pollinated flowers [ 5 ]). The behavioral impacts of floral iridescence and its potential ecological significance are unknown [ 6–9 ]. Animal-pollinated flowers are described as “sensory billboards” [ 10 ], with many floral features contributing to a conspicuous display that filters prospective pollinators. Yet floral iridescence is more subtle to the human eye than that of many animal displays because the floral diffraction grating is not perfectly regular [ 5–9 ]. This presents a puzzle: if the function of petals is to attract pollinators, then flowers might be expected to optimize iridescence to increase showiness. On the other hand, pollinators memorize floral colors as consistent advertisements of reward quality, and iridescence might corrupt flower color identity. Here we tested the trade-off between flower detectability and recognition, requiring bumblebees ( Bombus terrestris ) to identify artificial flowers that varied in pigmentation and degree of iridescence. We find that iridescence does increase target detectability but that “perfect” iridescence (produced by an artificial diffraction grating) corrupts target identity and bees make many mistakes. However, “imperfect” floral iridescence does not lead to mistaken target identity, while still benefitting flower detectability. We hypothesize that similar trade-offs might be found in the many naturally “imperfect” iridescence-producing structures found in animal-animal, as well as other plant-animal, interactions. [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]

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]

Highlights: [•] Anthocyanins, dominated by delphanin, were more concentrated in purple- than in yellow-flowered Iris lutescens. [•] Purple and yellow flowers could be visually discriminated between by bumbleebees. [•] The overall chemical composition of scent was not significantly different between morphs. [•] The scent of both morphs was dominated by terpenoids. [•] Color-scent correlation was demonstrated based on their shared biosynthetic pathway. [Copyright &y& Elsevier]

Visual and olfactory cues are the first interface between flowers and their visitors and are adaptations to facilitate successful pollination. Initial responses to and associative learning of multimodal cues by flower visitors are based on the perception of colours and volatiles. In this study, we tested how visual, olfactory and multimodal stimuli affect the behaviour of bumblebees ( Bombus terrestris) and correlated the properties of 28 stimuli in relation to the insects' sensory equipment to these behaviours. Initial responses and associative learning were recorded using a radio frequency identification system, tracking the visitation sequences of individual bumblebees to artificial flowers treated with naturally occurring pigments and/or volatiles. The salience of the stimuli was evaluated as the colour contrast to the background and as electroantennogram responses. The main finding was that both initial responses and learning performance were positively correlated to the salience of the stimuli, suggesting that salience is a key feature of flower cues in the interactions with insects. The salience of compound stimuli consisting of two or more pigments and/or volatiles was largely additively determined by the saliences of individual compounds. Potentially, the valence of the stimuli may interfere with the positive relationship between salience and behaviour, which is indicated by our results, too. The salience of multimodal cues depends on the species-specific equipment of visual and olfactory receptors and thus enables flowers to be advertising for some but rather inconspicuous for other flower visitors. [ABSTRACT FROM AUTHOR]

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]

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]

Plants often attract pollinators with floral displays composed of visual, olfactory, tactile and gustatory stimuli. Since pollinators' responses to each of these stimuli are usually studied independently, the question of why plants produce multi-component floral displays remains relatively unexplored. Here we used signal detection theory to test the hypothesis that complex displays reduce a pollinator's uncertainty about the floral signal. Specifically, we asked whether one component of the floral display, scent, improved a bee's certainty about the value of another component, color hue. We first trained two groups of bumble bees (Bombus impatiens Cresson) to discriminate between rewarding and unrewarding artificial flowers of slightly different hues in the presence vs absence of scent. In a test phase, we presented these bees with a gradient of floral hues and assessed their ability to identify the hue rewarded during training. We interpreted the extent to which bees' preferences were biased away from the unrewarding hue (`peak shift') as an indicator of uncertainty in color discrimination. Our data show that the presence of an olfactory signal reduces uncertainty regarding color: not only was color learning facilitated on scented flowers but also bees showed a lower amount of peak shift in the presence of scent. We explore potential mechanisms by which scent might reduce uncertainty about color, and discuss the broader significance of our results for our understanding of signal evolution. [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]

It has been argued that trichromatic bees with photoreceptor spectral sensitivity peaks in the ultraviolet (UV), blue and green areas of the spectrum are blind to long wavelengths (red to humans). South American temperate forests (SATF) contain a large number of human red-looking flowers that are reported to be visited by the bumblebee Bombus dahibomli. In the present study, B. dahlbomii's spectral sensitivity was measured through electroretinogram (ERG) recordings. No extended sensitivity to long wavelengths was found in B. dahlbomii. The spectral reflectance curves from eight plant species with red flowers were measured. The color loci occupied by these flowers in the bee color space was evaluated using the receptor noise-limited model. Four of the plant species have pure red flowers with low levels of chromatic contrast but high levels of negative L-receptor contrast. Finally, training experiments were performed in order to assess the role of achromatic cues in the detection and discrimination of red targets by B. dahlbomii. The results of the training experiments suggest that the bumblebee relies on achromatic contrast provided by the L-receptor to detect and discriminate red targets. These findings are discussed in the context of the evolutionary background under which the relationship between SATF species and their flower visitors may have evolved. [ABSTRACT FROM AUTHOR]

1. Most pollination biologists have used the collective pollinator visits to a plant as the measure of its pollinator attraction. However, we know very little about how many returns by the same individuals compose these visits, and how far each visitor travels after leaving the plant. Such behavioural aspects of individual pollinators are essential to understand the patterns of pollen flow among plants. 2. We observed plant visits by tagged bumble bees Bombus diversus in a field population of Cirsium purpuratum. By dissecting the collective visitation data into visits made by individual foragers, we addressed how ‘visitor density’ (number of individuals that visited a plant per 2 h) and ‘individual visitation rate’ (number of visits made by each individual per 2 h) are related to floral display size (number of flowering heads on a plant) and local flower density (number of flowering heads on neighbouring plants). We also tracked individual bees to determine how display size and local flower density of a plant influences its relative position in a bee's foraging area. 3. Plants attracted both regular visitors (bees that visited a plant more than three times per 2 h) and occasional visitors (bees that visited a plant fewer than four times per 2 h). Densities of both types of visitors increased with floral display size, whereas only occasional visitor's density increased with local flower density. 4. Individual bees preferred to visit central plants within their own foraging areas, plants with larger displays, and plants with lower local flower density. However, these preferences were independent from one another. Plants with large displays were not necessarily chosen by a bee as the centre of its own foraging area. On the other hand, plants with high local flower density were often located near the centre of a bee's foraging area. 5. The observed pollinator movements have implications for pollen flow in the plant population. Plants with larger displays probably experience greater mate diversity by attracting more occasional visitors, but they also assure matings with particular plants by increasing returns from regular visitors. [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]

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]

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]

The article offers the author's insights on the studies that showed the distinct patterns in flowers that has electric surface charge which enables the sensory hairs of bumble bees to discriminate artificaial flowers. Topics mentioned include the accumulation of charge by bees that aid in pollination, the judging of the bee that a flower has been visited by another bee, and the detection through mechanosensory neurons and the use of laser Doppler vibrometer to measure the movements.

Pollinators often sequentially visit 1 flower type while bypassing other equally rewarding flower types, behavior known as flower constancy. One explanation for flower constancy is that pollinators use the search image of a specific flower type to efficiently find flowers because they are often cryptic. The so-called search image hypothesis predicts that temporal specialization to 1 flower type by an individual pollinator declines as flower conspicuousness increases because if flowers are conspicuous, pollinators can divide their attention among different flower types. To test this prediction, we investigated visitation sequences of individual bumble bees (Bombus ignitus) to a patch of blue and yellow artificial flowers. We used different flower sizes and interflower distances as independent determinants of flower visual size and noisiness of the background and, thus, of flower crypsis. Flower color selectivity and flower search time decreased and subsequently increased with visual size of the nearest flowers, with a minimum of approximately 15° of the visual angle. Within-bout constancy, measured as the tendency of flying to the same flower types in succession compared with the expectation of a given flower selectivity within a bout, increased with visual size of the nearest flowers. Flower selectivity and within-bout constancy did not differ among arrays that shared the same floral visual angle, indicating that visual size, rather than flight distance or flower size itself, had a substantial effect on flower choices. Our results suggest that flower visual size and background noisiness affect flower crypsis and constancy, providing support for the search image hypothesis.We experimentally show that small flower sizes and background noisiness increase short-time flower specialization by bumble bees. This is in agreement with the food searching behavior of some vertebrates that limit their search to one type of inconspicuous food while ignoring other types that provide similar benefit. Our findings indicate that this behavior, termed search image foraging, affects flower choices by bees and suggest a universal tendency to form search images in food seeking animals. [ABSTRACT FROM PUBLISHER]