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Over decades, pesticide regulations have cycled between approval and implementation, followed by the discovery of negative effects on nontarget organisms that result in new regulations, pesticides, and harmful effects. This relentless pattern undermines the capacity to protect the environment from pesticide hazards and frustrates end users that need pest management tools. Wild pollinating insects are in decline, and managed pollinators such as honey bees are experiencing excessive losses, which threatens sustainable food security and ecosystem function. An increasing number of studies demonstrate the negative effects of field-realistic exposure to pesticides on pollinator health and fitness, which contribute to pollinator declines. Current pesticide approval processes, although they are superior to past practices, clearly continue to fail to protect pollinator health. In the present article, we provide a conceptual framework to reform cyclical pesticide approval processes and better protect pollinators.

While most models of decision-making assume that individuals assign options absolute values, animals often assess options comparatively, violating principles of economic rationality. Such 'irrational' preferences are especially common when two rewards vary along multiple dimensions of quality and a third, 'decoy' option is available. Bumblebees are models of decision-making, yet whether they are subject to decoy effects is unknown. We addressed this question using bumblebees (Bombus impatiens) choosing between flowers that varied in their nectar concentration and reward rate. We first gave bees a choice between two flower types, one higher in concentration and the other higher in reward rate. Bees were then given a choice between these flowers and either a 'concentration' or 'rate' decoy, designed to be asymmetrically dominated on each axis. The rate decoy increased bees' preference in the expected direction, while the concentration decoy did not. In a second experiment, we manipulated choices along two single reward dimensions to test whether this discrepancy was explained by differences in how concentration versus reward rate were evaluated. We found that low-concentration decoys increased bees' preference for the medium option as predicted, whereas low-rate decoys had no effect. Our results suggest that both low- and high-value flowers can influence pollinator preferences in ways previously unconsidered. [ABSTRACT FROM AUTHOR]

Novel insecticides are continuously being developed for application in response to increased legal restriction of previously developed insecticides and resistance in target insects. These novel insecticides, such as flupyradifurone (FPF), remain relatively untested on non‐target organisms, including bumblebees. Further, existing tests on honeybees tend to focus on adult mortality and thus sub‐lethal effects, such as impacts on reproductive output, are neglected, despite their importance for population‐level impacts.To address if the novel insecticide FPF has sub‐lethal effects on bumblebee reproduction and behaviour, we established microcolonies and chronically exposed them to field‐realistic concentrations over a 14‐day period.We found that exposure to FPF reduced the bumblebees' reproductive output in terms of the number of larvae produced and the mean mass of each larval instar. FPF‐treated bees also stored less sucrose and constructed fewer honeypots. However, adult bumblebee mortality was similar between control and FPF‐exposed microcolonies.Our results show that field‐realistic FPF exposure leads to increased larval mortality and/or delayed larval development, as well as reduced nectar storage, without affecting adult mortality.Policy implications. Insecticides that impair bumblebee reproduction can have long‐term population‐level consequences, even if adult bees do not experience increased mortality. Despite this fact, sub‐lethal effects, such as impacts on reproduction, are not mandatorily assessed within the regulatory process. Our findings highlight the importance of determining sub‐lethal effects of pesticides across developmental stages, as well as using pollinator species other than honeybees within the regulatory process. [ABSTRACT FROM AUTHOR]

Agricultural intensification has been identified as one of the key causes of global insect biodiversity losses. These losses have been further linked to the widespread use of agrochemicals associated with modern agricultural practices. Many of these chemicals are known to have negative sublethal effects on commercial pollinators, such as managed honeybees and bumblebees, but less is known about the impacts on wild bees. Laboratory‐based studies with commercial pollinators have consistently shown that pesticide exposure can impact bee behavior, with cascading effects on foraging performance, reproductive success, and pollination services. However, these studies typically assess only one chemical, neglecting the complexity of real‐world exposure to multiple agrochemicals and other stressors. In the summer of 2020, we collected wild‐foraging workers of the common eastern bumblebee, Bombus impatiens, from five squash (Cucurbita) agricultural sites (organic and conventional farms), selected to represent a range of agrochemical, including neonicotinoid insecticide, use. For each bee, we measured two behaviors relevant to foraging success and previously shown to be impacted by pesticide exposure: sucrose responsiveness and locomotor activity. Following behavioral testing, we used liquid chromatography–tandem mass spectrometry (LC–MS/MS) chemical analysis to detect and quantify the presence of 92 agrochemicals in each bumblebee. Bees collected from our sites did not vary in pesticide exposure as expected. While we found a limited occurrence of neonicotinoids, two fungicides (azoxystrobin and difenoconazole) were detected at all sites, and the pesticide synergist piperonyl butoxide (PBO) was present in all 123 bees. We found that bumblebees that contained higher levels of PBO were less active, and this effect was stronger for larger bumblebee workers. While PBO is unlikely to be the direct cause of the reduction in bee activity, it could be an indicator of exposure to pyrethroids and/or other insecticides that we were unable to directly quantify, but which PBO is frequently tank‐mixed with during pesticide applications on crops. We did not find a relationship between agrochemical exposure and bumblebee sucrose responsiveness. To our knowledge, this is the first evidence of a sublethal behavioral impact of agrochemical exposure on wild‐foraging bees. [ABSTRACT FROM AUTHOR]

Foraging theory assumes that animals assess value based on objective payoffs; however, animals often evaluate rewards comparatively, forming expectations based on recent experience. This form of evaluation may be particularly relevant for nectar foragers such as bumblebees, where individuals can visit thousands of flowers daily that vary in nectar quality. While many animals, including bees, demonstrate reference-based evaluation in experimental contexts, it is unclear whether this occurs in the wild. Here, we asked how daily experience with wildflower nectar influenced wild bumblebees' reward evaluation. We measured the daily nectar concentration of bee-visited wildflowers (Penstemon spp.), before presenting foragers with conspecific flowers filled with a range of artificial nectar concentrations. We recorded bees' acceptance of artificial nectar, the probability of subsequent visits to flowers on the same plant, and residence time. While bees had a minimum threshold of nectar acceptability that was unaffected by experience, when there was higher-concentration environmental nectar, they were less likely to accept lower-quality rewards on manipulated plants. Bees also visited more flowers and stayed longer on plants with higher-concentration nectar. This study shows evidence for both absolute and reference-based evaluation in wild bees and points towards differences between bees' behavior in lab- and wild-foraging contexts. [ABSTRACT FROM AUTHOR]

Neonicotinoid insecticides can have sub‐lethal effects on bees which has led to calls from conservationists for a global ban. In contrast, agrochemical companies argue that neonicotinoids do not harm honeybees at field‐realistic levels. However, the focus on honeybees neglects the potential impact on other bee species. We conducted a meta‐analysis to assess whether field‐realistic neonicotinoid exposure has sub‐lethal effects on non‐Apis bees. We extracted data from 53 papers (212 effects sizes) and found that it largely consisted of two genera: bumblebees (Bombus) and mason bees (Osmia), highlighting a substantial taxonomic knowledge gap. Neonicotinoid exposure negatively affected reproductive output across all bees and impaired bumblebee colony growth and foraging. Neonicotinoids also reduced Bombus, but not Osmia, individual development (growth and body size). Our results suggest that restrictions on neonicotinoids should benefit bee populations and highlight that the current regulatory process does not safeguard pollinators from the unwanted consequences of insecticide use. [ABSTRACT FROM AUTHOR]

Bumblebees are important pollinators of agricultural crops and wildflowers, but many species are in decline. Neonicotinoid insecticides are the most commonly used insecticide globally and can have negative sublethal effects on bumblebee colony growth and reproduction. Individual bumblebees can visit hundreds to thousands of flowers a day to forage for their colony. As such, they are a model species for studying optimal foraging, and small impairments to an individual's foraging decisions may have compounding effects on the colony's nutritional intake. We exposed bumblebees (Bombus impatiens) to an acute, field-realistic dose of the neonicotinoid insecticide imidacloprid, before allowing them to forage on an artificial floral array. We found that neonicotinoid-exposed bumblebees made suboptimal foraging decisions, as they were more likely to visit flowers located further apart than control bees.This indicates that for a given flower patch, individual bees exposed to a neonicotinoid will likely use more energy and forage less efficiency than unexposed bees, although further studies that directly measure energetic cost are required to confirm this. Given the robust and growing body of evidence demonstrating negative sublethal effects of neonicotinoids on bees, sublethal assessments on non-Apis bees should be made mandatory within the regulatory process. [ABSTRACT FROM AUTHOR]

Females and males often face different sources of selection, resulting in dimorphism in morphological, physiological, and even cognitive traits. Sex differences are often studied in respect to spatial cognition, yet the different ecological roles of males and females might shape cognition in multiple ways. For example, in dietary generalist bumblebees (Bombus), the ability to learn associations is critical to female workers, who face informationally rich foraging scenarios as they collect nectar and pollen from thousands of flowers over a period of weeks to months to feed the colony. While male bumblebees likely need to learn associations as well, they only forage for themselves while searching for potential mates. It is thus less clear whether foraging males would benefit from the same associative learning performance as foraging females. In this system, as in others, cognitive performance is typically studied in lab-reared animals under captive conditions, which may not be representative of patterns in the wild. In the first test of sex and species differences in cognition using wild bumblebees, we compared the performance of Bombus vancouverensis nearcticus (formerly bifarius) and Bombus vosnesenskii of both sexes on an associative learning task at Sierra Nevada (CA) field sites. Across both species, we found that males and females did not differ in their ability to learn, although males were slower to respond to the sucrose reward. These results offer the first evidence from natural populations that male bumblebees may be equally as able to learn associations as females, supporting findings from captive colonies of commercial bees. The observed interspecific variation in learning ability opens the door to using the Bombus system to test hypotheses about comparative cognition. [ABSTRACT FROM AUTHOR]

Learning associations between food-related stimuli and nutrients allows foragers to collect resources efficiently. In turn, the nutrients that foragers consume can themselves affect learning performance, through innate preferences for pre-ingestive stimuli, as well as post-ingestive reinforcement. Bees are insect models of learning and memory, yet the vast majority of this research concerns nectar (carbohydrate) rather than pollen (protein/lipid) rewards, despite the fact that many bees collect both simultaneously. We asked how one component of pollen surface chemistry, a free fatty acid (oleic acid), affected bees' performance in a nectar-learning task. We found that ingestion of oleic acid enhanced visual learning, likely through positive post-ingestive reinforcement. This was supported by our finding that although bees did not prefer to consume the oleic acid solution, its ingestion both decreased motor activity and increased survival. These results are a step towards understanding how nutritionally complex floral rewards may affect cognitive processes that underlie pollination mutualisms. [ABSTRACT FROM AUTHOR]

Abstract: Understanding how animals perceive, learn and remember stimuli is critical for understanding both how cognition is shaped by natural selection, and how ecological factors impact behaviour. However, the majority of studies on cognition involve captive animals in laboratory settings. While controlled settings are required to accurately measure aspects of cognition, they may not yield realistic estimates of learning performance in natural environments. Wild bees offer a useful system in which to study cognitive ecology and comparative cognition more broadly: they encompass around 20,000 species globally, varying in characteristics such as life‐history strategy, degree of sociality and dietary specialization. Yet, the limited number of protocols currently available for studying insect cognition has restricted research to a few commercially available bee species, in almost exclusively laboratory settings. We present a protocol (Free‐Moving Proboscis Extension Response [FMPER]) to measure wild bees’ colour preferences, learning performance and memory. We first used laboratory‐reared bumblebees Bombus impatiens to establish that FMPER yielded results consistent with learning theory. We then successfully tested wild honeybees Apis mellifera in the laboratory and Bombus vosnesenskii at field sites. Free‐Moving Proboscis Extension Response is straightforward to implement, is low cost, and may be readily adapted to other flower‐visiting insects. We believe it will be useful to a broad range of evolutionary biologists, behavioural ecologists and pollination ecologists interested in measuring cognitive performance in the wild and across a broader range of species. [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]

How foragers cope with complexity in both needs and resources is a major question in behavioral ecology. When faced with nutritionally diverse resources, or when foraging for offspring with divergent nutritional needs, animals must meet the challenge of how to structure their foraging bouts, including what resources to forage for and in what order (how) to collect them. We investigated how nutritional variation in resources and requirements shapes the structure of bumble bee foraging bouts. Bumble bee workers collect 2 nutritionally distinct resources for consumers with different nutritional needs, floral nectar (largely carbohydrates) for their own needs and that of larvae, and pollen (largely protein) that is used primarily by larvae. We maintained colonies of the Eastern Bumble Bee (Bombus impatiens) in the laboratory on either protein-rich or protein-limited diets and assessed bees' foraging bout structure on artificial flowers that offered low, medium, or high ratios of pollen to nectar. We analyzed bout structure using both traditional floral constancy metrics as well as hierarchical Bayesian analyses. Bees from pollen-satiated colonies responded to variation in floral pollen:nectar ratios, tending to collect pollen consecutively when nectar volumes were high. In contrast, foragers from pollen-limited colonies were relatively insensitive to floral reward ratio, tending to collect pollen in long runs regardless of nectar volume. We discuss the implications of these findings for the pollination services that bees provide plants. [ABSTRACT FROM AUTHOR]

Ever since Karl von Frisch's Nobel Prize-winning work in the early 1900s, bees have served as an important model system for the study of learning, memory and foraging behaviour. Bees can learn about floral features including colour, scent, texture and electrostatic charge, and show surprisingly sophisticated forms of learning. However, nearly every study of bee cognition and foraging to date has used a sole reward: nectar, most often in the form of a simple sucrose solution. Plants also offer a number of other rewards to pollinators, the most prevalent being pollen that bees collect as their primary source of protein. Indeed, a significant proportion of angiosperm species are nectarless, rewarding bees with pollen alone. Surprisingly, whether free-flying bees can learn visual features based solely on floral pollen rewards is unknown. Here we show that bees can learn to associate multiple floral features with a pure pollen reward. Furthermore, these associations are remembered long term, comparable to bees' memory for nectar associations. These findings raise new questions about bee learning and the evolutionary history between plants and bee pollinators. [ABSTRACT FROM AUTHOR]

Bees are vital pollinators of crops and wildflowers and as such, wild bee declines threaten food security and functioning ecosystems. One driver of bee declines is the use of systemic insecticides, such as commonly used neonicotinoids. However, rising pest resistance to neonicotinoids, and restrictions on their use in the EU, has increased the demand for replacement insecticides to control crop pests. Flupyradifurone is a novel systemic insecticide that is thought to be relatively 'bee safe' although it can be present in the nectar and pollen of bee-attractive crops. Bumblebees rely on learning to forage efficiently, and thus detriments to learning performance may have downstream consequences on their ability to forage. While neonicotinoids negatively influence bumblebee learning and memory, whether this is also the case for their replacements is unclear. Here, we exposed bumblebees (Bombus impatiens) to an acute, field-realistic dose of flupyradifurone before training them to learn either an olfactory or colour association. We found that flupyradifurone impaired bumblebees' learning and memory performance in both olfactory and visual modalities. Flupyradifurone-treated bees were also less motivated to feed. Given the similarity between the detriments to cognition found here and those previously reported for neonicotinoids, this implies that these insecticides may have similar sub-lethal effects on bees. Restrictions on neonicotinoid use are therefore unlikely to benefit bees if novel insecticides like flupyradifurone are used as an alternative, highlighting that current agrochemical risk assessments are not protecting bees from the unwanted consequences of pesticide use. Sub-lethal assessments on non- Apis bees should be made mandatory in agrochemical regulation to ensure that novel insecticides are indeed 'bee safe'. [Display omitted] • Flupyradifurone is a novel insecticide that could replace neonicotinoids. • Flupyradifurone impaired bumblebee olfactory and colour learning. • Bumblebees exposed to flupyradifurone were also less motivated to feed. • Policy makers should incorporate sub-lethal effects on bumblebees into risk assessment. [ABSTRACT FROM AUTHOR]

Across the animal kingdom, many animals build structures. One especially diverse example is that of nest building by birds. It remains unclear, however, what birds know or whether they learn about the structural aspects of the material with which they build a nest. Here we tested whether nest-building male zebra finches would choose the appropriate type of material when building in a novel situation. They did do this: males provided with a nestbox with either a small or a large entrance hole and with nest material of two types (‘long’ and ‘short’) chose the type of material that was appropriate for the box in which they built. Additionally, the birds' material use improved with experience: males building in nestboxes with small entrances became less choosy in their material choice as they became more skilled at inserting material of either length into their nestbox. The birds, therefore, first chose the appropriate materials for the nestbox in which they were building but then modified their handling skills so as to make use of all of the available material. How the cognitive abilities used in this nest-building context compare with those used in solving other physical problems such as tool use tasks is not yet clear. [Copyright &y& Elsevier]

Highlights: [•] It is not always clear why nest-building birds choose the materials they do. [•] We compared nest and food colour preferences in male zebra finches. [•] Birds preferred blue nest material, but had no food colour preferences. [•] It appears that nest material colour preferences are specific to that context. [Copyright &y& Elsevier]

Nests are built by nearly all bird species and can be extremely varied in their structural characteristics, both within and among species. As with a number of other avian behaviours, it seems plausible that early learning might be important in producing adult nest-building behaviour. To examine whether preferences that adults have for nest materials are related to their early-life experience, we experimentally manipulated the colour of the nest in which Zebra Finch pairs built and raised chicks. We then tested these chicks at maturity to determine whether they preferred the colour of the nest from which they had fledged or preferred the same colour as their father. We also examined the overall structure of nests that fathers and their sons built to determine whether the nest a male builds resembles that from which he hatched. When males and females naïve to building were paired as adults and tested for their nest material preferences, they did not prefer the colour of their natal nest. When these males were re-paired and their preference tested a second time, the majority then preferred the colour that their father had preferred (which was also the colour preferred by most of the males). The structural components of a male's nest did not resemble the nest built by his father, but neither did his father's nests resemble each other. We found no evidence that the experience of the nest from which a bird fledges influences his preferences for the colour of nest material or the structure of his first nest. [ABSTRACT FROM AUTHOR]

Whether learning plays a role in nest building in birds is largely unknown. Here we investigated whether the colour of nest materials used to build a first nest affected the subsequent nest material choices made by male zebra finches when building a second nest. Males were tested for their preference for green or brown nest material and then were provided with either their preferred or nonpreferred colour with which to build their first nest. The success of this nesting attempt was manipulated such that half of the breeding pairs had their eggs removed, while the other half were allowed to keep their eggs and fledge chicks. Males were then retested to determine which colour of nest material they now preferred. Males had strong initial preferences for one or other of the two colours of nest material. Males that built a nest with their preferred colour of nest material continued to prefer that colour after nesting, regardless of their breeding success in that nest. However, of the males that built a nest with material of their nonpreferred colour, those that raised and fledged chicks from it subsequently preferred that colour of material for their second nest, while males that suffered a failed breeding attempt did not. Thus breeding experience can influence decisions relating to nest material choice in nest construction in the zebra finch. [Copyright &y& Elsevier]

Intensive agriculture is reliant on pesticides to control crop pests, but these chemicals can have negative environmental consequences. This has resulted in repeated calls for pesticide risk assessments to be modified to better protect ecosystem services such as pollination. However, the pesticide licensing process is complex, and consequently there is often confusion between risk assessments where the environmental impact of pesticide use is considered, and risk management where licensing decisions are made. Using bees as a case study, we provide a roadmap for how pesticides are licensed for use in the European Union. By outlining the regulatory process, we highlight key data gaps that need to be addressed to generate a holistic approach to environmental risk assessment. Such an approach is vital to protect pollinators and wildlife more broadly from the unintended consequences of pesticide use. Siviter et al. outline the regulatory process by which pesticides are licenced in the European Union, and highlight gaps that need to be addressed to generate a holistic approach to environmental risk assessment. [ABSTRACT FROM AUTHOR]

While most models of decision-making assume that individuals assign options absolute values, animals often assess options comparatively, violating principles of economic rationality. Such 'irrational' preferences are especially common when two rewards vary along multiple dimensions of quality and a third, 'decoy' option is available. Bumblebees are models of decision-making, yet whether they are subject to decoy effects is unknown. We addressed this question using bumblebees ( Bombus impatiens ) choosing between flowers that varied in their nectar concentration and reward rate. We first gave bees a choice between two flower types, one higher in concentration and the other higher in reward rate. Bees were then given a choice between these flowers and either a 'concentration' or 'rate' decoy, designed to be asymmetrically dominated on each axis. The rate decoy increased bees' preference in the expected direction, while the concentration decoy did not. In a second experiment, we manipulated choices along two single reward dimensions to test whether this discrepancy was explained by differences in how concentration versus reward rate were evaluated. We found that low-concentration decoys increased bees' preference for the medium option as predicted, whereas low-rate decoys had no effect. Our results suggest that both low- and high-value flowers can influence pollinator preferences in ways previously unconsidered.

Judgement bias, or 'optimism' and 'pessimism', has been demonstrated across many taxa, yet the cognitive mechanisms underlying this behaviour remain unclear. In an optimism paradigm, animals are trained to an association, and, if given a positive experience, behave more favourably towards 'ambiguous' stimuli. We tested whether this effect could be explained by changes to stimulus response gradients by giving bees a task where their response was tested across a wider gradient of stimuli than typically tested. In line with previous work, we found that bees given a positive experience demonstrated judgement bias, being more likely to visit ambiguous stimuli. However, bees were also less likely to visit a stimulus on the other side of the rewarded stimulus (S+), and as such had a shifted stimulus response curve, showing a diminished peak shift response. In two follow-up experiments we tested the hypothesis that our manipulation altered bees' stimulus response curves via changes to the peak shift response by reducing peak shift in controls. We found that, in support of our hypothesis, elimination of peak shift also eliminated differences between treatments. Our results point towards a cognitive explanation of 'optimistic' behaviour in non-human animals and offer a new paradigm for considering emotion-like states., Competing Interests: We declare we have no competing interests., (© 2023 The Authors.)

While classic models of animal decision-making assume that individuals assess the absolute value of options, decades of research have shown that rewards are often evaluated relative to recent experience, creating incentive contrast effects. Contrast effects are often assumed to be purely sensory, yet consumer and experimental psychology tell us that label-based expectations can affect value perception in humans and rodents. However, this has rarely been tested in non-model systems. Bumblebees forage on a variety of flowers that vary in their signals and rewards and show contrast when rewards are lowered. We manipulated bees' expectations of stimulus quality, before downshifting the reward to induce incentive contrast. We found that contrast effects were not solely driven by experience with a better reward, but also influenced by experience with associated stimuli. While bees' initial response did not differ between treatments, individuals were faster to accept the lower-quality reward when it was paired with a novel stimulus. We explored the boundaries of these label-based expectations by testing bees along a stimulus gradient and found that expectations generalized to similar stimuli. Such reference-dependent evaluations may play an important role in bees' foraging choices, with the potential to impact floral evolution and plant community dynamics.

Species' cognitive traits are shaped by their ecology, and even within a species, cognition can reflect the behavioural requirements of individuals with different roles. Social insects have a number of discrete roles (castes) within a colony and thus offer a useful system to determine how ecological requirements shape cognition. Bumblebee queens are a critical point in the lifecycle of their colony, since its future success is reliant on a single individual's ability to learn about floral stimuli while finding a suitable nest site; thus, one might expect particularly adept learning capabilities at this stage. I compared wild Bombus vosnesenskii queens and workers on their ability to learn a colour association and found that queens performed better than workers. In addition, queens of another species, B. insularis, a cuckoo species with a different lifecycle but similar requirements at this stage, performed equally well as the non-parasitic queens. To control for differences in foraging experience, I then repeated this comparison with laboratory-based B. impatiens and found that unmated queens performed better than workers. These results add to the body of work on how ecology shapes cognition and opens the door to further research in comparative cognition using wild bees.

Neonicotinoid pesticides can have a multitude of negative sublethal effects on bees. Understanding their impact on wild populations requires accurately estimating the dosages bees encounter under natural conditions. This is complicated by the possibility that bees might influence their own exposure: two recent studies found that bumblebees ( Bombus terrestris ) preferentially consumed neonicotinoid-contaminated nectar, even though these chemicals are thought to be tasteless and odourless. Here, we used Bombus impatiens to explore two elements of these reported preferences, with the aim of understanding their ecological implication and underlying mechanism. First, we asked whether preferences persisted across a range of realistic nectar sugar concentrations, when measured at a series of time points up until 24 h. Second, we tested whether bees' neonicotinoid preferences were driven by an ability to associate their post-ingestive consequences with floral stimuli such as colour, location or scent. We found no evidence that foragers preferred to consume neonicotinoid-containing solutions, despite finding effects on feeding motivation and locomotor activity in line with previous work. Bees also did not preferentially visit floral stimuli previously paired with a neonicotinoid-containing solution. These results highlight the need for further research into the mechanisms underlying bees' responses to these pesticides, critical for determining how neonicotinoid-driven foraging preferences might operate in the real world for different bee species., Competing Interests: The authors have no competing interests., (© 2020 The Authors.)

Neonicotinoid pesticides can impair bees' ability to learn and remember information about flowers, critical for effective foraging. Although these effects on cognition may contribute to broader effects on health and performance, to date they have largely been assayed in simplified protocols that consider learning in a single sensory modality, usually olfaction. Given that real flowers display a variety of potentially useful signals, we assessed the effects of acute neonicotinoid exposure on multimodal learning in free-flying bumblebees. We found that neonicotinoid consumption differentially impacted learning of floral stimuli, impairing scent, but not colour, learning. These findings raise questions about the mechanisms by which pesticides might differentially impair sensory systems, with implications for how neonicotinoids affect multiple aspects of bee ecology.

Pollen plays a dual role as both a gametophyte and a nutritional reward for pollinators. Although pollen chemistry varies across plant species, its functional significance in pollination has remained obscure, in part because little is known about how floral visitors assess it. Bees rely on pollen for protein, but whether foragers evaluate its chemistry is unclear, as it is primarily consumed by larvae. We asked whether the chemical composition of pollen influences bumblebees' foraging behaviour. Using putatively sweet and bitter pollen blends, we found that chemical composition influenced two aspects of bee behaviour relevant to plant fitness: the amount of pollen collected and the likelihood of subsequently visiting a visually similar flower. These findings offer a new perspective on the nutritional ecology of plant-pollinator interactions, as they show that pollen's taste may mediate its collection and transfer., (© 2016 The Author(s).)

Bees are model organisms for the study of learning and memory, yet nearly all such research to date has used a single reward, nectar. Many bees collect both nectar (carbohydrates) and pollen (protein) on a single foraging bout, sometimes from different plant species. We tested whether individual bumblebees could learn colour associations with nectar and pollen rewards simultaneously in a foraging scenario where one floral type offered only nectar and the other only pollen. We found that bees readily learned multiple reward-colour associations, and when presented with novel floral targets generalized to colours similar to those trained for each reward type. These results expand the ecological significance of work on bee learning and raise new questions regarding the cognitive ecology of pollination., (© 2015 The Author(s).)