Drought stress influences foraging preference of a solitary bee on two wildflowers.

BACKGROUND AND AIMS: Pollinators provide critical ecosystem services, maintaining biodiversity and benefiting global food production. However, plants, pollinators, and their mutualistic interactions may be affected by drought, which has increased in severity and frequency under climate change. Using two annual, insect-pollinated wildflowers (Phacelia campanularia and Nemophila menziesii), we asked how drought impacts floral traits and foraging preferences of a solitary bee (Osmia lignaria) and explore potential implications for plant reproduction. METHODS: In greenhouses, we experimentally subjected plants to drought to induce water stress, as verified by leaf water potential. To assess the impact of drought on floral traits, we measured flower size, floral display size, nectar volume, and nectar sugar concentration. To explore how drought-induced effects on floral traits affected bee foraging preferences, we performed choice trials. Individual female bees were placed into foraging arenas with two conspecific plants, one droughted and one non-droughted, and were allowed to forage freely. KEY RESULTS: We determined that P. campanularia is more drought-tolerant than N. menziesii based on measures of turgor loss point, and confirmed that droughted plants were more drought-stressed than non-droughted plants. For droughted plants of both species, floral display size was reduced, and flowers were smaller and produced less, more-concentrated nectar. We found that bees preferred non-droughted flowers of N. menziesii. However, bee preference for non-droughted P. campanularia flowers depended on the time of day and was detected only in the afternoon. CONCLUSIONS: Our findings indicate that bees prefer visiting non-droughted flowers, likely reducing pollination success for drought-stressed plants. Lack of preference for non-droughted P. campanularia flowers in the morning may reflect the higher drought tolerance of this species. This work highlights the potentially intersecting, short-term physiological and pollinator behavioral responses to drought and suggests that such responses may reshape plant-pollinator interactions, ultimately reducing reproductive output for less drought-tolerant wildflowers.

Bacterial and Fungal Symbionts in Pollen Provisions of a Native Solitary Bee in Urban and Rural Environments.

Among insects, symbionts such as bacteria and fungi can be linked to their physiology and immature development, and in some cases are part of a defense system against parasites and diseases. Current bacterial and fungal symbiont associations in solitary bees are understudied, especially in the Pacific Northwest region of the USA. We collected pollen provisions from the native spring-foraging solitary bee, Osmia lignaria Say, across two distinct foraging periods over 2 years at 22 sites along an urban-to-rural gradient in western Washington. We then used next-generation sequencing to identify bacterial and fungi within pollen provisions and assessed the effect of their richness and diversity on O. lignaria larval development success and adult emergence. We detected a significantly positive relationship between bacterial diversity in pollen with O. lignaria larval developmental success, and higher bacterial richness and diversity during the later foraging period. Fungal generic richness and diversity decreased with increasing plant richness. Although neither was associated with O. lignaria developmental success, we did detect Ascosphaera spp. known to be pathogenic to O. lignaria and other bee species. Neither bacterial or fungal richness or diversity was affected by site type when classified as urban or rural. This study provides new information on bacterial and fungal symbionts present in pollen provisions of a native solitary bee when foraging across urban and rural areas of the Pacific Northwest.

Native solitary bee reproductive success depends on early season precipitation and host plant richness.

Spring-emerging bees depend upon the synchronized bloom times of angiosperms that provide pollen and nectar for offspring. The emergence of such bees and bloom times are linked to weather but can be phenologically mismatched, which could limit bee developmental success. However, it remains unclear how such phenologically asynchrony could affect spring-emerging pollinators, and especially for those that forage over a relatively short time period. We examined the relationship between weather and host plant selection on the native spring-foraging solitary bee, Osmia lignaria, across 3 years at urban and rural sites in and around Seattle, Washington, USA. We used community science weather data to test the effects of precipitation, wind, and temperature on O. lignaria oviposition and developmental success. We also collected pollen data over two distinct foraging periods, early and late spring, and used Next-Generation Sequencing to identify plant genera from pollen. Among the weather variables, precipitation during the early foraging period adversely affected larval developmental success and adult bee emergence success, but not oviposition. Using DNA metabarcoding, we observed that increases in the number of plant genera in pollen increased adult emergence in both foraging periods, but not oviposition or larval development. We also observed that foraging bees consistently visited certain genera during each foraging period, especially Acer, Salix, and Rubus. However, pollen collected by O. lignaria over different years varied in the number of total genera visited, highlighting the importance of multi-year studies to ascertain bee foraging preferences and its link to developmental success.

Wildfire severity influences offspring sex ratio in a native solitary bee.

Although ecological disturbances can have a strong influence on pollinators through changes in habitat, virtually no studies have quantified how characteristics of wildfire influence the demography of essential pollinators. Nevertheless, evaluating this topic is critical for understanding how wildfire is linked to pollinator population dynamics, particularly given recent changes in wildfire frequency and severity in many regions of the world. In this study, we measured the demographic response of the blue orchard bee (Osmia lignaria) across a natural gradient of wildfire severity to assess how variation in wildfire characteristics influenced reproductive output, offspring sex ratio, and offspring mass. We placed nest blocks with a standardized number and sex ratio of pre-emergent adult bees across the wildfire gradient, finding some evidence for a positive but highly variable relationship between reproductive output and fire severity surrounding the nest site at both local (100 m) and landscape (750 m) scales. In addition, the production of female offspring was > 10% greater at nest sites experiencing the greatest landscape-scale fire severity relative to the lowest-severity areas. The finding that blue orchard bees biased offspring production towards the more expensive offspring sex with increasing fire severity shows a functional response to changes in habitat quality through increased density of flowering plants. Our findings indicate that burned mixed-conifer forest provides forage for the blue orchard bee across a severity gradient, and that the increase in floral resources that follows high-severity fire leads females to shift resource allocation to the more costly sex when nesting.

Body and Wing Allometries Reveal Flight-Fecundity Tradeoff in Response to Larval Provisioning in Osmia lignaria (Hymenoptera: Megachilidae).

Variation in body size has important implications for physical performance and fitness. For insects, adult size and morphology are determined by larval growth and metamorphosis. Female blue orchard bees, Osmia lignaria, (Say) provision a finite quantity of food to their offspring. In this study, we asked how provision-dependent variation in size changes adult morphology. We performed a diet manipulation in which some larvae were starved in the final instar and some were given unlimited food. We examined the consequences on adult morphology in two ways. First, allometric relationships between major body regions (head, thorax, abdomen) and total body mass were measured to determine relative growth of these structures. Second, morphometrics that are critical for flight (wing area, wing loading, and extra flight power index) were quantified. Head and thorax mass had hyperallometric relationships with body size, indicating these parts become disproportionately large in adults when larvae are given copious provisions. However, abdominal mass and wing area increased hypoallometrically with body size. Thus, large adults had disproportionately lighter abdomens and smaller wing areas than smaller adults. Though both males and females followed these general patterns, allometric patterns were affected by sex. For flight metrics, small adults had reduced wing loading and an increased extra flight power index. These results suggest that diet quantity alters development in ways that affect the morphometric trait relationships in adult O. lignaria and may lead to functional differences in performance.

Cross-infectivity of honey and bumble bee-associated parasites across three bee families.

Recent declines of wild pollinators and infections in honey, bumble and other bee species have raised concerns about pathogen spillover from managed honey and bumble bees to other pollinators. Parasites of honey and bumble bees include trypanosomatids and microsporidia that often exhibit low host specificity, suggesting potential for spillover to co-occurring bees via shared floral resources. However, experimental tests of trypanosomatid and microsporidial cross-infectivity outside of managed honey and bumble bees are scarce. To characterize potential cross-infectivity of honey and bumble bee-associated parasites, we inoculated three trypanosomatids and one microsporidian into five potential hosts - including four managed species - from the apid, halictid and megachilid bee families. We found evidence of cross-infection by the trypanosomatids Crithidia bombi and C. mellificae, with evidence for replication in 3/5 and 3/4 host species, respectively. These include the first reports of experimental C. bombi infection in Megachile rotundata and Osmia lignaria, and C. mellificae infection in O. lignaria and Halictus ligatus. Although inability to control amounts inoculated in O. lignaria and H. ligatus hindered estimates of parasite replication, our findings suggest a broad host range in these trypanosomatids, and underscore the need to quantify disease-mediated threats of managed social bees to sympatric pollinators.

Comparison of the Chemical Compositions of the Cuticle and Dufour's Gland of Two Solitary Bee Species from Laboratory and Field Conditions.

Species-specific biochemistry, morphology, and function of the Dufour's gland have been investigated for social bees and some non-social bee families. Most of the solitary bees previously examined are ground-nesting bees that use Dufour's gland secretions to line brood chambers. This study examines the chemistry of the cuticle and Dufour's gland of cavity-nesting Megachile rotundata and Osmia lignaria, which are species managed for crop pollination. Glandular and cuticular lipid compositions were characterized and compared to each other and according to the nesting experience of adult females. Major lipid classes found were hydrocarbons, free fatty acids, and wax esters. Many components were common to the cuticle and Dufour's glands of each species, yet not identical in number or relative composition. Wax esters and fatty acids were more prevalent in Dufour's glands of M. rotundata than on cuticles. Wax esters were more abundant on cuticles of O. lignaria than in Dufour's glands. In both species, fatty acids were more prevalent in glands of field-collected females compared to any other sample type. Chemical profiles of cuticles and glands were distinct from each other, and, for O. lignaria, profiles of laboratory-maintained bees could be distinguished from those of field-collected bees. Comparison of percentiles of individual components of cuticular and glandular profiles of the same bee showed that the proportions of some cuticular components were predictive of the proportion of the same glandular components, especially for nesting females. Lastly, evidence suggested that Dufour's gland is the major source of nest-marking substances in M. rotundata, but evidence for this role in O. lignaria was less conclusive.

Influence of Nest Box Color and Release Sites on Osmia lignaria (Hymenoptera: Megachilidae) Reproductive Success in a Commercial Almond Orchard.

Intensively managed, commercial orchards offer resources for managed solitary bees within agricultural landscapes and provide a means to study bee dispersal patterns, spatial movement, nest establishment, and reproduction. In 2012, we studied the impact of 1) the color of nest boxes covaried with four nest box density treatments and 2) the number of bee release sites covaried with two nest box density treatments on the reproductive success of Osmia lignaria Say in a California almond orchard pollinated by a mixture of O. lignaria and Apis mellifera L. Nest box color influenced the number of nests, total cells, and cells with male and female brood. More nests and cells were produced in light blue nest boxes than in orange or yellow nest boxes. The covariate nest box density also had a significant effect on brood production. The number of release sites did not affect O. lignaria nesting and reproduction, but the number of cavities in nest boxes influenced reproduction. Overall, the color of nest boxes and their distribution, but not the number of release sites, can greatly affect O. lignaria nest establishment and reproductive success in a commercial almond orchard. The ability to locate nesting sites in a homogenous, large orchard landscape may also be facilitated by the higher frequency of nest boxes with low numbers of cavities, and by the ability to detect certain nest box colors that best contrast with the blooming trees.

Mason Bees (Hymenoptera: Megachilidae) Exhibit No Avoidance of Imidacloprid-Treated Soils.

1) Many wild bee species interact with soil either as a nesting substrate or material. These soil interactions create a risk of exposure to agrochemicals such as imidacloprid or other neonicotinoid pesticides that can persist in soil for months after application. At the landscape level, concentrations of imidacloprid residue in soil are limited to the immediate treatment area, and thus risks to soil-interacting bees could be low if they avoid contaminated soils. 2) We utilized Osmia lignaria (Say), a solitary cavity nesting bee which collects mud to partition and seal nests, and conducted two laboratory experiments to test whether nesting females select or avoid soils containing various levels of imidacloprid residue. For the first experiment, we assessed behavioral responses of females to treated soil utilizing a choice arena and pairing various choices of soil with imidacloprid residues ranging between 0 and 780 ppb. For the second experiment, we developed a laboratory assay to assess soil selection of actively nesting O. lignaria, by providing choices of contaminated soil between 0 and 100 ppb and 0 and 1,000 ppb to nesting females. 3) We found no evidence that O. lignaria females avoided any level of imidacloprid contamination, even at the highest residue level (1,000 ppb) in both the experiments, which may have implications for risk. The in situ nesting methodology developed in this study has future applications for research on soil or pollen preferences of cavity nesting Osmia species, and potential for breeding of O. lignaria in laboratory.

Ecology and Economics of Using Native Managed Bees for Almond Pollination.

Native managed bees can improve crop pollination, but a general framework for evaluating the associated economic costs and benefits has not been developed. We conducted a cost-benefit analysis to assess how managing blue orchard bees (Osmia lignaria Say [Hymenoptera: Megachildae]) alongside honey bees (Apis mellifera Linnaeus [Hymenoptera: Apidae]) can affect profits for almond growers in California. Specifically, we studied how adjusting three strategies can influence profits: (1) number of released O. lignaria bees, (2) density of artificial nest boxes, and (3) number of nest cavities (tubes) per box. We developed an ecological model for the effects of pollinator activity on almond yields, validated the model with published data, and then estimated changes in profits for different management strategies. Our model shows that almond yields increase with O. lignaria foraging density, even where honey bees are already in use. Our cost-benefit analysis shows that profit ranged from -US$1,800 to US$2,800/acre given different combinations of the three strategies. Adding nest boxes had the greatest effect; we predict an increase in profit between low and high nest box density strategies (2.5 and 10 boxes/acre). In fact, the number of released bees and the availability of nest tubes had relatively small effects in the high nest box density strategies. This suggests that growers could improve profits by simply adding more nest boxes with moderate number of tubes in each. Our approach can support grower decisions regarding integrated crop pollination and highlight the importance of a comprehensive ecological economic framework for assessing these decisions.

Progeny of Osmia lignaria from distinct regions differ in developmental phenology and survival under a common thermal regime.

Many insects, including some bees, have extensive subcontinental distributions that can differ in climatic conditions. Within and beyond these distributions, humans intentionally transport beneficial insects, including bees, to non-natal geographic locations. Insects also are experiencing unprecedented climatic change in their resident localities. For solitary bees, we know very little about the adaptive plasticity and geographic variation in developmental physiology that accommodates the different climates experienced within distributional ranges. Osmia lignaria Say (Hymenoptera: Megachilidae) is a widely distributed North American spring-emerging bee being developed as a managed pollinator for tree fruit crops, including almonds. We examined the development and survival of O. lignaria progeny that were descended from populations sourced from southern California, western Washington, and northern Utah, and then were reared together under an hourly and weekly temperature regime simulating those of a California almond-growing region. We found that developmental physiologies of Washington and Utah progeny were generally similar. However, California progeny developed slower, were more metabolically active, and survived better under California conditions than did populations native to regions at higher latitudes. Regardless of geographic origin, cocooned adults managed under prescribed thermal regimes emerged faster and lived longer after wintering. Progeny of parents from different regions exhibited some acclimatory plasticity in developmental phenologies to a novel climatic regime, but overall their responses reflected their geographic origins. This outcome is consistent with their developmental phenologies being largely heritable adaptations to regional climates.