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Abstract The microbial composition of stored food can influence its stability and the microbial species consumed by the organism feeding on it. Many bee species store nectar and pollen in provisions constructed to feed developing offspring. Yet, whether microbial composition is determined by the pollen types within provisions, variation between bee species at the same nesting sites, or geographic distance was unclear. Here, we sampled two species of cooccurring cavity nesting bees in the genus Osmia at 13 sites in California and examined the composition of pollen, fungi, and bacteria in provisions. Pollen composition explained 15% of variation in bacterial composition and ∼30% of variation in fungal composition, whereas spatial distance among sites explained minimal additional variation. Symbiotic microbe genera Ascosphaera, Sodalis, and Wolbachia showed contrasting patterns of association with pollen composition, suggesting distinct acquisition and transmission routes for each. Comparing provisions from both bee species comprised of the same pollens points to environmental acquisition rather than bee species as a key factor shaping the early stages of the bee microbiome in Osmia. The patterns we observed also contrast with Apilactobacillus-dominated provision microbiome in other solitary bee species, suggesting variable mechanisms of microbial assembly in stored food among bee species. 

ABSTRACT Microbial environmental transmission among individuals plays an important role in shaping the microbiomes of many species. Despite the importance of the microbiome for host fitness, empirical investigations on environmental transmission are scarce, particularly in systems where interactions across multiple trophic levels influence symbiotic dynamics. Here, we explore microbial transmission within insect microbiomes, focusing on solitary bees. Specifically, we investigate the environmental transmission hypothesis, which posits that solitary bees acquire and deposit their associated microbiota from and to their surroundings, especially flowers. Using experimental setups, we examine the transmission dynamics ofApilactobacillus micheneri, a fructophilic and acidophilic bacterium, between the solitary beeOsmia lignaria(Megachilidae) and the plantPhacelia tanacetifolia(Boraginaceae). Our results demonstrate that bees not only acquire bacteria from flowers but also deposit these microbes onto uninoculated flowers for other bees to acquire them, supporting a bidirectional microbial exchange. We therefore find empirical support for the environmental transmission hypothesis, and we discuss the multitrophic dependencies that facilitate microbial transmission between bees and flowers. 

Abstract Host–microbe interactions underlie the development and fitness of many macroorganisms, including bees. Whereas many social bees benefit from vertically transmitted gut bacteria, current data suggests that solitary bees, which comprise the vast majority of species diversity within bees, lack a highly specialized gut microbiome. Here we examine the composition and abundance of bacteria and fungi throughout the complete life cycle of the ground-nesting solitary bee Anthophora bomboides standfordiana. In contrast to expectations, immature bee stages maintain a distinct core microbiome consisting of Actinobacterial genera (Streptomyces, Nocardiodes) and the fungus Moniliella spathulata. Dormant (diapausing) larval bees hosted the most abundant and distinctive bacteria and fungi, attaining 33 and 52 times their initial copy number, respectively. We tested two adaptive hypotheses regarding microbial functions for diapausing bees. First, using isolated bacteria and fungi, we found that Streptomyces from brood cells inhibited the growth of multiple pathogenic filamentous fungi, suggesting a role in pathogen protection during overwintering, when bees face high pathogen pressure. Second, sugar alcohol composition changed in tandem with major changes in fungal abundance, suggesting links with bee cold tolerance or overwintering biology. We find that A. bomboides hosts a conserved core microbiome that may provide key fitness advantages through larval development and diapause, which raises the question of how this microbiome is maintained and faithfully transmitted between generations. Our results suggest that focus on microbiomes of mature or active insect developmental stages may overlook stage-specific symbionts and microbial fitness contributions during host dormancy. 

Abstract For most animals, the microbiome is key for nutrition and pathogen defence, and is often shaped by diet. Corbiculate bees, including honey bees, bumble bees, and stingless bees, share a core microbiome that has been shaped, at least in part, by the challenges associated with pollen digestion. However, three species of stingless bees deviate from the general rule of bees obtaining their protein exclusively from pollen (obligate pollinivores) and instead consume carrion as their sole protein source (obligate necrophages) or consume both pollen and carrion (facultative necrophages). These three life histories can provide missing insights into microbiome evolution associated with extreme dietary transitions. Here, we investigate, via shotgun metagenomics, the functionality of the microbiome across three bee diet types: obligate pollinivory, obligate necrophagy, and facultative necrophagy. We find distinct differences in microbiome composition and gene functional profiles between the diet types. Obligate necrophages and pollinivores have more specialized microbes, whereas facultative necrophages have a diversity of environmental microbes associated with several dietary niches. Our study suggests that necrophagous bee microbiomes may have evolved to overcome cellular stress and microbial competition associated with carrion. We hypothesize that the microbiome evolved social phenotypes, such as biofilms, that protect the bees from opportunistic pathogens present on carcasses, allowing them to overcome novel nutritional challenges. Whether specific microbes enabled diet shifts or diet shifts occurred first and microbial evolution followed requires further research to disentangle. Nonetheless, we find that necrophagous microbiomes, vertebrate and invertebrate alike, have functional commonalities regardless of their taxonomy. 

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 (Andrenaspp. andAndrena [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) inB. impatiensworkers to 388.4 ± 146.2 ppb inAndrena. Exposure profiles were similar for all bees exceptA. mellifera, whose unique exposure profile included high levels of the neonicotinoid insecticide thiamethoxam.All bee groups except wildB. impatiensqueens had at least one sample exceeding a US Environmental Protection Agency or European Food Safety Authority exposure level of concern.Apis melliferaexperienced 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. 

Abstract Bee–fungus associations are common, and while most studies focus on entomopathogens, emerging evidence suggests that bees associate with a variety of symbiotic fungi that can influence bee behavior and health. Here, we review nonpathogenic fungal taxa associated with different bee species and bee-related habitats. We synthesize results of studies examining fungal effects on bee behavior, development, survival, and fitness. We find that fungal communities differ across habitats, with some groups restricted mostly to flowers (Metschnikowia), while others are present almost exclusively in stored provisions (Zygosaccharomyces). Starmerella yeasts are found in multiple habitats in association with many bee species. Bee species differ widely in the abundance and identity of fungi hosted. Functional studies suggest that yeasts affect bee foraging, development, and pathogen interactions, though few bee and fungal taxa have been examined in this context. Rarely, fungi are obligately beneficial symbionts of bees, whereas most are facultative bee associates with unknown or ecologically contextual effects. Fungicides can reduce fungal abundance and alter fungal communities associated with bees, potentially disrupting bee–fungi associations. We recommend that future study focus on fungi associated with non-honeybee species and examine multiple bee life stages to document fungal composition, abundance, and mechanistic effects on bees. 

Abstract Floral nectar is frequently colonised by microbes. However, nectar microbial communities are typically species‐poor and dominated by few cosmopolitan genera. One hypothesis is that nectar constituents may act as environmental filters. We tested how five non‐sugar nectar compounds as well as elevated sugar impacted the growth of 12 fungal and bacterial species isolated from nectar, pollinators, and the environment. We hypothesised that nectar isolated microbes would have the least growth suppression. Additionally, to test if nectar compounds could affect the outcome of competition between microbes, we grew a subset of microbes in co‐culture across a subset of treatments. We found that some compounds such as H₂O₂suppressed microbial growth across many but not all microbes tested. Other compounds were more specialised in the microbes they impacted. As hypothesised, the nectar specialist yeastMetschnikowia reukaufiiwas unaffected by most nectar compounds assayed. However, many non‐nectar specialist microbes remained unaffected by nectar compounds thought to reduce microbial growth. Our results show that nectar chemistry can influence microbial communities but that microbe‐specific responses to nectar compounds are common. Nectar chemistry also affected the outcome of species interactions among microbial taxa, suggesting that non‐sugar compounds can affect microbial community assembly in flowers. 

Theoretical frameworks of terrestrial community assembly often focus on single trophic levels (e.g. plants) without considering how complex interdependencies across different trophic levels influence assembly mechanisms. Yet, when multiple trophic levels are considered (e.g. plant–pollinator, plant–microbe interactions) the focus is typically on network analyses at local spatial scales. As spatial variation in biodiversity (β‐diversity) is increasingly being recognized for its relevance in understanding community assembly and conservation, considering how β‐diversity at one trophic level may be influenced by assembly processes that alter abundance and composition of interacting communities at a different trophic level (multitrophic dependency) is critical. Here, we build on single trophic level community assembly frameworks to explore the assembly processes affecting β‐diversity in multitrophic communities comprising flowering plants, their bee pollinators, and the corresponding bee‐gut microbiota to better understand the importance of multitrophic dependency in community assembly. Using distance‐based redundancy analysis and variation partitioning, we investigated community assembly processes across three interconnected trophic levels in two ecological regions in southern California: the Santa Monica Mountains and three islands of the Channel Island Archipelago. We found that the deterministic effects of multitrophic dependency are stronger on directly connected trophic levels than on indirectly connected trophic levels (i.e. flowers explain bee communities and bees explain bee‐gut bacteria communities, but flowers weakly explain variation in bee‐gut bacteria communities). We also found notable regional variation, where multitrophic dependency was weaker on the Channel Islands as ecological drift was more pronounced. Our results suggest that integrating the influence of multitrophic dependency on community assembly is important for elucidating drivers of β‐diversity and that multitrophic dependency can be determined by the regional context in which β‐diversity is measured. Taken together, our results highlight the importance of considering multiscale perspectives – both multitrophic and multiregional – in community assembly to fully elucidate assembly processes. 

Wild bees are vital for the pollination of native plants and crops, providing essential ecosystem services. Climate change is known to impact biodiversity and species distributions, but insects adapted to desert ecosystems may exhibit unique physiological, behavioral, and evolutionary responses. The desert pallid bee (C. pallida), a solitary bee native to the arid southwestern United States and northern Mexico, primarily forages on yellow palo verde (P. microphylla), blue palo verde (P. florida), and desert ironwood (O. tesota). This study used MaxEnt to estimate the current and projected geographical overlap of suitable habitats for C. pallida and its host plants. Here, we used MaxEnt to estimate the current and forecasted overlapping geographically suitable habitat of C. pallida with all three host plants. We forecasted potential environmentally suitable areas for each species to the year 2040 using the current distribution model and climate projections with moderate CO2 levels. We found a continued spatial alignment in the suitable area of the bee and its host plants with a 70% increase in the range overlap area, though shifted to higher average altitudes and a slight northern expansion. These findings may provide insight to stakeholders on the conservation needs of desert-dwelling pollinators.