Bombus Impatiens Research Articles

Learning to handle flowers increases pollen collection for bees but does not affect pollination success for plants

Abstract Cooperation and conflict are common in plant–pollinator interactions. Flowering plants often entice pollinators to visit by offering floral food rewards, thereby facilitating pollination. However, pollinators such as bees can learn to improve their collection of floral rewards (such as pollen), changing how they interact with the flower’s reproductive organs, which together could reduce pollination success. Consequently, complex flowers that slow pollinator learning might benefit the plant. Yet how pollinator learning and flower complexity interact to affect pollination success is unknown. We therefore asked how differences in complexity of 4 flower types (Phacelia campanularia, Exacum affine, Solanum elaeagnifolium, and Erythranthe guttata) affected learning by pollen-foraging generalist bumble bees (Bombus impatiens) and how learning affected pollen collection and pollen deposition on these flowers. We found that bees generally learned how to efficiently handle more complex flower types more slowly. Bees that required more visits to become efficient foragers collected less pollen, with no effect on pollen deposition. Except for the simplest flower type, learning also involved development of motor routines unique to each flower type. Experienced bees overall collected more pollen, but individual differences in motor routines did not affect pollen collection. Conversely, individual differences in motor routines affected pollen deposition, but there was no overall effect of experience. Thus, even though learning overall benefits the bee, it does not alter female (and potentially male) fitness benefits for the plant. We discuss potential reasons for these patterns and consequences for bee behavior and flower evolution.

Insect Pollinivores of Sorghum bicolor and Plant Traits that Influence Visitation1

Abstract Grasses such as sorghum, (Sorghum bicolor (L.) Moench), are rarely considered as an important food resource for pollinators. Here, we report insects collecting or consuming pollen of sorghum over the 8-wk flowering period of a mapping population in Tifton, GA. We also examine the response of insects to plant morphological traits and disease damage and the ability of bumble bees (Bombus impatiens) to cross-pollinate sorghum. The most numerous insect observed collecting/consuming sorghum pollen was the hover fly Toxomerus politus Say (i.e., the maize calligrapher) followed by honey bees (Apis mellifera L.), lined earwigs (Doru taeniatum Dorhn), lovebugs (Plecia nearctica Hardy), southern carpenter bees (Xylocopa micans Lepeletier), common eastern bumble bees (Bombus impatiens Cresson), exotic stripetails (Allograpta exotica Wiedemann), margined soldier beetles (Chauliognathus marginatus F.), a signal fly (Rivellia sp. Robineau-Desvoidy), and a dusky-winged hover fly (Ocyptamus fuscipennis Say). Maximum flowering occurred at Week 3, which coincided with the maximum number of maize calligraphers, bumble bees, and honey bees observed. A positive linear relationship was seen between the number of flowering plots and the number of maize calligraphers and honey bees. The probability of observing a bee on a panicle increased as plant height increased, whereas the probability of observing a bee or hover fly decreased as plant disease percentage increased. These results suggest that inflorescence abundance, plant height, and plant disease impact bee visitation in sorghum. Furthermore, field-captured bumble bees successfully fertilized sorghum in a greenhouse study, suggesting that bumblebees can pollinate sorghum flowers in field conditions.

Economic foraging in a floral marketplace: asymmetrically dominated decoy effects in bumblebees.

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.

Changes in bumblebee queen gut microbiotas during and after overwintering diapause.

Bumblebees are key pollinators with gut microbiotas that support host health. After bumblebee queens undergo winter diapause, which occurs before spring colony establishment, their gut microbiotas are disturbed, but little is known about community dynamics during diapause itself. Queen gut microbiotas also help seed worker microbiotas, so it is important that they recover post-diapause to a typical community structure, a process that may be impeded by pesticide exposure. We examined how bumblebee queen gut microbiota community structure and metabolic potential shift during and after winter diapause, and whether post-diapause recovery is affected by pesticide exposure. To do so, we placed commercial Bombus impatiens queens into diapause, euthanizing them at 0, 2 and 4 months of diapause. Additionally, we allowed some queens to recover from diapause for 1 week before euthanasia, exposing half to the common herbicide glyphosate. Using whole-community, shotgun metagenomic sequencing, we found that core bee gut phylotypes dominated queen gut microbiotas before, during and after diapause, but that two phylotypes, Schmidhempelia and Snodgrassella, ceased to be detected during late diapause and recovery. Despite fluctuations in taxonomic community structure, metabolic potential remained constant through diapause and recovery. Also, glyphosate exposure did not affect post-diapause microbiota recovery. However, metagenomic assembly quality and our ability to detect microbial taxa and metabolic pathways declined alongside microbial abundance, which was substantially reduced during diapause. Our study offers new insights into how bumblebee queen gut microbiotas change taxonomically and functionally during a key life stage and provides guidance for future microbiota studies in diapausing bumblebees.

Consumption of pollen contaminated with field-realistic concentrations of fungicide causes sublethal effects in Bombus impatiens (Hymenoptera: Apidae) microcolonies.

Bumble bees are declining across the globe. The causes of this decline have been attributed to a variety of stressors, including pesticides. Fungicides are a type of pesticide that has been understudied in the context of bumble bee health. As a result, fungicides are often applied to flowering plants without consideration of pollinator exposure. Recent work demonstrates that fungicides have sublethal effects in bumble bees, but little is known about how much fungicide it takes to cause these sublethal effects. To address this gap in the literature, we fed microcolonies of the common eastern bumble bee (Bombus impatiens CressonHymenoptera: ApidaeHymenoptera: ApidaeHymenoptera: ApidaeHymenoptera: Apidae) pollen contaminated with a range of fungicide concentrations. We chose these concentrations based on the range of fungicide concentrations in pollen and nectar that were reported in the literature. Results revealed that later-stage pupae and newly emerged males are potentially sensitive to fungicide exposure, showing smaller size and reduced fat reserves at intermediate levels of contamination. Compared to the control, intermediated levels of fungicide-contaminated pollen led to increased pupal mortality and delayed male emergence. Contrary to expectations, higher fungicide levels did not exhibit a linear relationship with negative impacts, suggesting nuanced effects. Because body size and emergence timing are important aspects of bumble bee reproductive behavior, results have implications for mating success, potentially disrupting colony development.

Mating frequency estimation and its importance for colony abundance analyses in eusocial pollinators: a case study of Bombus impatiens (Hymenoptera: Apidae).

The genus Bombus (bumble bees) includes approximately 265 species, many of which are in decline in North America and Europe. To estimate colony abundance of bumble bees in natural and agricultural habitats, sibship relationships are often reconstructed from genetic data with the assumption that colonies have 1 monandrous queen. However, some species such as the North American common eastern bumble bee (Bombus impatiens Cresson) can display low levels of polyandry, which may bias estimates of colony abundance based on monandrous sibship reconstructions. To accurately quantify rates of polyandry in wild and commercially mated queens of this species, we empirically estimated mating frequencies using a novel statistical model and genotypes from 730 bees. To genotype individuals, we used a highly polymorphic set of microsatellites on colonies established from 20 wild-caught gynes and 10 commercial colonies. We found multiple fathers in 3 of the wild colonies and 3 of the commercial colonies. This resulted in average effective mating frequencies of 1.075 ± 0.18 and 1.154 ± 0.25 for wild and commercial colonies, respectively. These findings agree with previous reports of low rates of polyandry for B. impatiens. Using a large empirical dataset, we demonstrate that assuming monandry for colony abundance estimation in species that violate this assumption results in an overestimation of the number of colonies. Our results emphasize the importance of studying mating frequencies in social species of conservation concern and economic importance for the accuracy of colony abundance estimation and for understanding their ecology and sociobiology.

The role of landscape factors in shaping bumble bee pathogen loads across regions of the eastern Nearctic

Abstract Pathogens and parasites are drivers of declines in bumble bees. Their levels can be influenced by numerous abiotic and biotic factors, thus managing disease in these bees requires understanding the relative impact of these factors on pathogen loads. We evaluated loads of black queen cell virus (BQCV) and deformed wing virus in bumble bees (Bombus impatiens) and honey bees (Apis mellifera), and loads of parasites Vairimorpha bombi and Crithidia bombi in B. impatiens, from sites varying in habitat type and quality across North Carolina, USA. Pathogen loads were assessed against metrics for land cover, floral quality, bee diversity and weather. Results were compared against similar data from Pennsylvania, USA, and other landscape studies on bumble bees in North America. Spatial variation in pathogen loads was lower in bumble bees in North Carolina than in Pennsylvania, which reduced the power to detect landscape effects. For example, Crithidia was fairly ubiquitous and Vairimorpha was not detected. Data from both states revealed that developed land and honey bees were most consistently positively correlated with viral loads, especially for BQCV, whereas forest and nesting habitat availability were often negatively correlated with loads. Multivariate models only supported a positive relationship between summer floral resources and BQCV loads in North Carolina bumble bees. Together with the broader literature, these data indicate that levels of developed land and honey bees are most associated with increased pathogen loads while floral availability and bee community composition show more context‐dependent effects.

Size-dependent responses of colony-founding bumblebee (Bombus impatiens) queens to exposure to pesticide residues in soil during hibernation

Bumblebees and other key pollinators are experiencing global declines, a phenomenon driven by multiple environmental stressors, including pesticide exposure. While bumblebee queens spend most of their life hibernating underground, no study to date has examined how exposure to pesticide-contaminated soils might affect bumblebee queens during this solitary phase of their lifecycle. We exposed Bombus impatiens queens (n = 303) to soil treated with field-realistic concentrations of two diamide insecticides (chlorantraniliprole and cyantraniliprole) and two fungicides (boscalid and difenoconazole), alone or combined, during a 30-week hibernation period. We found that exposure to boscalid residues in soil doubled the likelihood of queens surviving through the colony initiation period (after successful hibernation) and laying eggs. Our data also revealed complex interactions between pesticide exposure and queen body mass on aspects of colony founding. Among others, exposure to cyantraniliprole led to lethal and sublethal post-hibernation effects that were dependent on queen size, with larger queens showing higher mortality rates, delayed emergence of their first brood, and producing smaller workers. Our results show that effects of pesticide exposure depend on intrinsic traits of bumblebee queen physiology and challenge our understanding of how bees respond to pesticides under environmentally realistic exposure scenarios.

Assessing the attractiveness of native wildflower species to bees (Hymenoptera: Anthophila) in the southeastern United States

Abstract Habitat loss, agricultural intensification, pesticide use, disease and climate change have contributed to the decline of numerous insect groups. Recent government initiatives have recognized the importance of supplementary wildflower plantings to support native bee populations, yet little information exists on the attractiveness of recommended plant species to bees. With the use of network analysis and generalized linear mixed models, we evaluated the attractiveness of 18 wildflower species belonging to 6 families (Apiaceae, Apocynaceae, Asteraceae, Fabaceae, Lamiaceae and Verbenaceae) planted in a randomized block design by conducting timed targeted sweep netting surveys. We also evaluated the floral preferences of four bee species (Apis mellifera, Halictus poeyi/ligatus, Bombus griseocollis and Bombus impatiens) to the planted wildflower species with the use of GLMs. We found that Asclepias tuberosa, Gaillardia pulchella and Verbena hastata attracted the greatest abundance, richness and diversity of native bees compared to the other wildflower species. Collectively, Asclepias tuberosa, Gaillardia pulchella and Verbena hastata attracted 89% of all bee taxa in the study, which exemplifies the importance of including them in a wildflower mixture to attract native bees. While several wildflower species attracted low abundance, richness and diversity of native bees such as Monarda fistulosa, Symphyotrichum pilosum and Eryngium yuccifolium, they are still ecologically important for attracting uncommon or specialist bee species and should be included in seed mixes for this reason. Solution. Our results highlight that each species of native wildflower varies in the abundance, richness and diversity of native bees they attract; this could be considered in the decision‐making of landowners and land managers to promote wildflower plantings to reflect the needs of certain groups of bees of conservation interest.

Herbivore and pollinator body size effects on strawberry fruit quality.

Land use change affects both pollinator and herbivore populations with consequences for crop production. Recent evidence also shows that land use change affects insect traits, with intraspecific body size of pollinators changing across landscape gradients. However, the consequences on crop production of trait changes in different plant interactors have not been well-studied. We hypothesized that changes in body size of key species can be enough to affect crop productivity, and therefore looked at how the field-realistic variation in body size of both an important pollinator, Bombus impatiens (Cresson), and a key pest herbivore, Lygus lineolaris (Palisot), can affect fruit size and damage in strawberry. First, we determined if pests vary in body size along land use gradients as prior studies have documented for pollinators; and second, we tested under controlled conditions how the individual and combined changes in size of an important pollinator and a key herbivore pest affect strawberry fruit production. The key herbivore pest was smaller in landscapes with more natural and semi-natural habitat, confirming that herbivore functional traits can vary along a land use gradient. Additionally, herbivore size, and not pollinator size, marginally affected fruit production-with plants exposed to larger pests producing smaller fruits. Our findings suggest that land use changes at the landscape level affect crop production not just through changes in the species diversity of insect communities that interact with the plant, but also through changes in body size traits.

Field agrochemical exposure impacts locomotor activity in wild bumblebees.

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.

Evaluating Pollination and Weed Control Strategies under Mesotunnel Systems for Organic Muskmelon Production in Iowa

Bacterial wilt of cucurbits, caused by Erwinia tracheiphila, is spread by spotted (Diabrotica undeimpunctata howardi) and striped (Acalymma vittatum) cucumber beetles and results in major losses for US cucurbit (Cucurbitaceae spp.) growers. Organic growers of muskmelon (Cucumis melo) lack reliable control measures against bacterial wilt. During previous field trials in Iowa, USA, a system called mesotunnels, which are 3.5-ft-tall barriers covered with a nylon mesh insect netting, resulted in a higher marketable yield of organic ‘Athena’ muskmelon than low tunnels or noncovered plots. However, satisfactory pollination and weed control are challenging in mesotunnels because the netting covers the crop for most or all of the growing season, and economic feasibility of these systems has not been determined. Consequently, two field trials conducted in Iowa from 2020 to 2022 evaluated strategies to ensure pollination under mesotunnels in commercial-scale plots, assess effectiveness of teff (Eragrostis tef) as a living mulch for weed control in mesotunnel systems, and compare the profitability of the treatment options for organic ‘Athena’ muskmelon. The treatments used during the pollination trial were as follows: full season, in which mesotunnels remained sealed all season and bumble bees (Bombus impatiens) were added at the start of bloom for pollination; open ends, wherein both ends of the tunnels were opened at the start of bloom then reclosed 2 weeks later; and on-off-on, in which nets were removed at the start of bloom and then reinstalled 2 weeks later. The full-season treatment had significantly higher marketable yield than the other treatments in two of three trial years. Plants with the full season and open ends treatments had a bacterial wilt incidence <2.5% across all three years and similar numbers of cucumber beetles, whereas plants with the on-off-on treatment had an average bacterial wilt incidence of 11.0% and significantly more cucumber beetles. The open ends treatment had fewer bee visits to ‘Athena’ muskmelon flowers than the other treatments. In the 2-year (2021–22) weed management trial, treatments applied to the furrow between plastic-mulched rows were as follows: landscape fabric; teff seeded at 4 lb/acre and mowed 3 weeks after seeding; teff seeded at 4 lb/acre and not mowed; a control with bare ground where weeds were mowed 3 weeks after transplanting; and a bare ground control with no mowing. The landscape fabric and mowed teff treatments had statistically similar marketable yield, and mowing appeared to minimize yield losses compared with nonmowed treatments. The landscape fabric had no weeds, followed by mowed teff, mowed bare ground, and nonmowed teff. Nonmowed bare ground had the highest weed biomass. The partial budget and cost-efficiency ratio analysis indicated that the full-season treatment was the most cost-efficient pollination option for mesotunnel systems. An economic analysis of the weed management strategies showed that using teff as a living mulch in the furrows between organic ‘Athena’ muskmelon rows, coupled with timely mowing to suppress its growth, can generate revenue comparable to that of landscape fabric. Our findings suggest that organic ‘Athena’ muskmelon growers in Iowa may gain the greatest yield and soil quality benefits when mesotunnels are kept closed for the entire season, bumble bees are used for pollination, and teff (mowed 3 weeks after seeding) is used to control weeds in the furrows. Further trials integrating these pollination and weed management strategies would help validate a comprehensive approach to organic ‘Athena’ muskmelon production under mesotunnels.

Determining the plant-pollinator network in a culturally significant food and medicine garden in the Great Lakes region.

Understanding the interactions between plants and pollinators within a system can provide information about pollination requirements and the degree to which species contribute to floral reproductive success. Past research has focused largely on interactions within monocultured agricultural systems and only somewhat on wild pollination networks. This study focuses on the culturally significant Three Sisters Garden, which has been grown and tended by many Indigenous peoples for generations in the Great Lakes Region. Here, the plant-pollinator network of the traditional Three Sisters Garden with the inclusion of some additional culturally significant plants was mapped. Important visitors in this system included the common eastern bumble bee, Bombus impatiens Cresson (Hymenoptera: Apidae), and the hoary squash bee, Xenoglossa pruinosa (Say) (Hymenoptera: Apidae), as determined by their abundances and pollinator service index (PSI) values. Understanding the key pollinators in the Three Sisters Garden links biological diversity to cultural diversity through the pollination of culturally significant plants. Further, this information could be of use in supporting Indigenous food sovereignty by providing knowledge about which wild pollinators could be supported to increase fruit and seed set within the Three Sisters Garden. Our findings can also lead to more effective conservation of important wild pollinator species.

Genetic modification of the bee parasite Crithidia bombi for improved visualization and protein localization

Crithidia bombi is a trypanosomatid parasite that infects several species of bumble bees (Bombus spp.), by adhering to their intestinal tract. Crithidia bombi infection impairs learning and reduces survival of workers and the fitness of overwintering queens. Although there is extensive research on the ecology of this host-pathogen system, we understand far less about the mechanisms that mediate internal infection dynamics. Crithidia bombi infects hosts by attaching to the hindgut via the flagellum, and one previous study found that a nectar secondary compound removed the flagellum, preventing attachment. However, approaches that allow more detailed observation of parasite attachment and growth would allow us to better understand factors mediating this host-pathogen relationship. We established techniques for genetic manipulation and visualization of cultured C. bombi. Using constructs established for Crithidia fasciculata, we successfully generated C. bombi cells expressing ectopic fluorescent transgenes using two different selectable markers. To our knowledge, this is the first genetic modification of this species. We also introduced constructs that label the mitochondrion and nucleus of the parasite, showing that subcellular targeting signals can function across parasite species to highlight specific organelles. Finally, we visualized fluorescently tagged parasites in vitro in both their swimming and attached forms, and in vivo in bumble bee (Bombus impatiens) hosts. Expanding our cell and molecular toolkit for C. bombi will help us better understand how factors such as host diet, immune system, and physiology mediate outcomes of infection by these common parasites.

Temperature influences desiccation resistance of bumble bees

Ongoing climate change has increased temperatures and the frequency of droughts in many parts of the world, potentially intensifying the desiccation risk for insects. Because resisting desiccation becomes more difficult at higher temperatures and lower humidity, avoiding water loss is a key challenge facing terrestrial insects. However, few studies have examined the interactive effects of temperature and environmental humidity on desiccation resistance in insects. Such studies on bees (Hymenoptera: Apoidea: Anthophila) are especially rare, despite their ecological and economic importance. Here, we crossed temperature (20, 25, and 30 °C) with humidity (<5, 50, >95 % RH) manipulations and measured time to mortality, water loss rates, and the water content at mortality of bumble bees (Bombus impatiens). We found that both higher temperature and lower humidity increased water loss rates, while warmer temperatures reduced survival time and lower humidity decreased water content at mortality. Additionally, we observed large intraspecific variation in water balance traits between colonies, and larger individuals survived longer and could tolerate more water loss before mortality. This study raises important questions about the mechanisms underpinning water loss in bumble bees and suggests that frequent access to nectar may be especially important for bumble bees’ water balance and survival in a warming and drying climate.

A novel insecticide impairs bumblebee memory and sucrose responsiveness across high and low nutrition.

Wild bees are important pollinators of crops and wildflowers but are exposed to a myriad of different anthropogenic stressors, such as pesticides and poor nutrition, as a consequence of intensive agriculture. These stressors do not act in isolation, but interact, and may exacerbate one another. Here, we assessed whether a field-realistic concentration of flupyradifurone, a novel pesticide that has been labelled as 'bee safe' by regulators, influenced bumblebee sucrose responsiveness and long-term memory. In a fully crossed experimental design, we exposed individual bumblebees (Bombus impatiens) to flupyradifurone at high (50% (w/w)) or low (15% (w/w)) sucrose concentrations, replicating diets that are either carbohydrate rich or poor, respectively. We found that flupyradifurone impaired sucrose responsiveness and long-term memory at both sucrose concentrations, indicating that better nutrition did not buffer the negative impact of flupyradifurone. We found no individual impact of sugar deficiency on bee behaviour and no significant interactions between pesticide exposure and poor nutrition. Our results add to a growing body of evidence demonstrating that flupyradifurone has significant negative impacts on pollinators, indicating that this pesticide is not 'bee safe'. This suggests that agrochemical risk assessments are not protecting pollinators from the unintended consequences of pesticide use.

ECR Spotlight – Kelton Verble and Ellen Keaveny

ECR Spotlight is a series of interviews with early-career authors from a selection of papers published in Journal of Experimental Biology and aims to promote not only the diversity of early-career researchers (ECRs) working in experimental biology but also the huge variety of animals and physiological systems that are essential for the ‘comparative’ approach. Kelton Verble and Ellen Keaveny are authors on ‘A rapid return to normal: temporal gene expression patterns following cold exposure in the bumble bee Bombus impatiens’, published in JEB. Kelton is a PhD candidate in the lab of Dr Jeffrey Lozier at The University of Alabama, USA, investigating the transcriptomics and epigenomics underlying thermal tolerance in bumble bees. Ellen is a PhD candidate in the lab of Michael Dillon at the University of Wyoming, USA, investigating how temperature affects bumble bees at cellular, organismal, population and community levels.

A rapid return to normal: temporal gene expression patterns following cold exposure in the bumble bee Bombus impatiens.

Bumble bees are common in cooler climates and many species likely experience periodic exposure to very cold temperatures, but little is known about the temporal dynamics of cold response mechanisms following chill exposure, especially how persistent effects of cold exposure may facilitate tolerance of future events. To investigate molecular processes involved in the temporal response by bumble bees to acute cold exposure, we compared mRNA transcript abundance in Bombus impatiens workers exposed to 0°C for 75 min (inducing chill coma) and control bees maintained at a constant ambient temperature (28°C). We sequenced the 3' end of mRNA transcripts (TagSeq) to quantify gene expression in thoracic tissue of bees at several time points (0, 10, 30, 120 and 720 min) following cold exposure. Significant differences from control bees were only detectable within 30 min after the treatment, with most occurring at the 10min recovery time point. Genes associated with gluconeogenesis and glycolysis were most notably upregulated, while genes related to lipid and purine metabolism were downregulated. The observed patterns of expression indicate a rapid recovery after chill coma, suggesting an acute differential transcriptional response during recovery from chill coma and return to baseline expression levels within an hour, with no long-term gene expression markers of this cold exposure. Our work highlights the functions and pathways important for acute cold recovery, provides an estimated time frame for recovery from cold exposure in bumble bees, and suggests that cold hardening may be less important for these heterothermic insects.

Insect Flight Energetics and the Evolution of Size, Form, and Function.

Flying insects vary greatly in body size and wing proportions, significantly impacting their flight energetics. Generally, the larger the insect, the slower its flight wingbeat frequency. However, variation in frequency is also explained by differences in wing proportions, where larger-winged insects tend to have lower frequencies. These associations affect the energy required for flight. The correlated evolution of flight form and function can be further defined using a lineage of closely related bee species varying in body mass. The decline in flight wingbeat frequency with increasing size is paralleled by the flight mass-specific metabolic rate. The specific scaling exponents observed can be predicted from the wing area allometry, where a greater increase (hyperallometry) leads to a more pronounced effect on flight energetics, and hypoallometry can lead to no change in frequency and metabolic rate across species. The metabolic properties of the flight muscles also vary with body mass and wing proportions, as observed from the activity of glycolytic enzymes and the phospholipid compositions of muscle tissue, connecting morphological differences with muscle metabolic properties. The evolutionary scaling observed across species is recapitulated within species. The static allometry observed within the bumblebee Bombus impatiens, where the wing area is proportional and isometric, affects wingbeat frequency and metabolic rate, which is predicted to decrease with an increase in size. Intraspecific variation in flight muscle tissue properties is also related to flight metabolic rate. The role of developmental processes and phenotypic plasticity in explaining intraspecific differences is central to our understanding of flight energetics. These studies provide a framework where static allometry observed within species gives rise to evolutionary allometry, connecting the evolution of size, form, and function associated with insect flight.

Pesticide risk during commercial apple pollination is greater for honeybees than other managed and wild bees

Abstract Most pesticide research has focussed on risk to managed honeybees, but other managed and wild bees are also exposed to pesticides. Critically, we know little about the magnitude and sources of risk to honeybees compared with other bees during crop pollination. To compare pesticide exposure and risk across wild and managed bees, we sampled the main bee groups present during bloom in 20 apple orchards, including managed honeybees (Apis mellifera), managed bumblebee workers (Bombus impatiens), wild mining bees (Andrena spp. and Andrena [Melandrena] spp.), bumblebee foundress queens (Bombus impatiens) and eastern carpenter bees (Xylocopa virginica). We screened all bees for 92 pesticides and computed a Risk Quotient using available toxicity data (honeybee LD50s), adjusting for differences in toxicity known to scale with body mass. To gain insight into exposure origin, we compared residues in bees to those in focal orchard apple and dandelion flowers. Nearly all bee samples contained pesticides (95%), with the average contamination level ranging from 7.1 ± 2.8 parts per billion (ppb) in B. impatiens workers to 388.4 ± 146.2 ppb in Andrena. Exposure profiles were similar for all bees except A. mellifera, whose unique exposure profile included high levels of the neonicotinoid insecticide thiamethoxam. All bee groups except wild B. impatiens queens had at least one sample exceeding a US Environmental Protection Agency or European Food Safety Authority exposure level of concern. Apis mellifera experienced significantly greater risk than other bee groups, with 63% and 81% of samples exceeding an acute or chronic exposure level of concern, respectively. Risk to honeybees was driven primarily by high thiamethoxam levels not found in focal orchard flowers and likely originating outside the orchard. Synthesis and applications: We find that pesticide exposure and risk differ between honeybees and other managed and wild bees during apple pollination. Furthermore, pesticide exposure is a landscape‐scale phenomenon and therefore measures to reduce exposure must consider the surroundings beyond focal farms. Limiting orchard sprays, while reducing on‐farm exposures, will not protect far‐foraging bees from off‐farm exposures such as thiamethoxam, which we hypothesize is coming from nearby seed‐treated corn fields planted during apple bloom.