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Pollinator-plant interactions represent a core mutualism that underpins biodiversity in terrestrial ecosystems, and the loss of flowering plants is a major driver of pollinator declines. Bee attraction to flowers is mediated by both quantity of resources (the number of available flowers for exploration) and quality of resources (pollen nutritional value), but whether and how bees prioritize these factors is not well understood. Here, we leveraged a unique plant system to investigate the floral factors influencing bee foraging decisions. Recombinant inbred plant lines were generated by crossing the self-fertilizing Capsella rubella and the pollinator-dependent outcrosser C. grandiflora , to produce plants that varied across floral traits. Using enclosed arenas, we evaluated the foraging behavior of two solitary bee species, Osmia cornifrons and Megachile rotundata , to the isolated inflorescences from these lines. Visits from O. cornifrons were significantly positively correlated with the number of flowers, while M. rotundata visits were significantly positively associated with pollen nutrition, with a preference for plants with higher pollen protein-to-lipid content. Further experiments using artificial flowers confirmed that M. rotundata preferred flowers with higher protein:lipid ratios, while O. cornifrons visits were unaffected by nutrition. These studies demonstrate that, although both bee species collect pollen as their sole source of protein and lipids for themselves and/or their offspring, they differentially prioritize resource quantity (number of flowers) and quality (pollen nutritional content). These studies lay the groundwork for understanding how different foraging strategies evolved, and influence, plant-pollinator ecological networks., (© The Author(s) 2024. Published by Oxford University Press on behalf of National Academy of Sciences.)

Apis mellifera Linnaeus (Hymenoptera: Apis), honey bees, are the most widely used managed crop pollinators. However, their high rental cost and uncertain availability for North American orchard crops have motivated growers to explore alternative pollination options. We examined whether adding solitary, spring-flying Osmia lignaria Say (Hymenoptera: Megachilidae), blue orchard bees, as co-pollinators with A. mellifera in Washington sweet cherry and pear orchards enhances fruit set and yield compared to the use of A. mellifera alone. We added managed O. lignaria to orchard sites where A. mellifera hives were already present. Fruit set, fruit yield, and O. lignaria reproduction at O. lignaria-supplemented sites were compared to nearby, paired sites pollinated only by A. mellifera (3 paired cherry and 3 paired pear sites). For both crops, the addition of O. lignaria significantly increased fruit set but did not yield at harvest. Microscopic inspection of pollen grains from O. lignaria nest cell provisions confirmed that O. lignaria primarily visited orchard flowers. Mean retention of O. lignaria in cherry orchards was slightly higher (65%) than O. lignaria retention reported in other orchard crops (30%-60%). However, retention in pear orchards was much lower (≤20%). These results show that supplementing hives with O. lignaria in Washington spring orchard crops can increase overall pollination, but that trees fail to bear developing fruit to maturity. The strategy of using co-pollinators, O. lignaria and A. mellifera, in US orchards may act as "pollination insurance" when A. mellifera hives are in low supply or when the weather is not amenable for A. mellifera flight during the bloom period., (Published by Oxford University Press on behalf of Entomological Society of America 2024.)

Characterizing the nutritional needs of wild bee species is an essential step to better understanding bee biology and providing suitable supplemental forage for at-risk species. Here, we aim to characterize the nutritional needs of a model solitary bee species, Osmia cornifrons (Radoszkowski), by using dietary protein-to-lipid ratio (P:L ratio) as a proxy for nutritional niche and niche breadth. We first identified the mean target P:L ratio (~3.02:1) and P:L collection range (0.75-6.26:1) from pollen provisions collected across a variety of sites and time points. We then investigated the P:L tolerance range of larvae by rearing bees in vitro on a variety of diets. Multifloral and single-source pollen diets with P:L ratios within the range of surveyed provisions did not always support larval development, indicating that other dietary components such as plant secondary compounds and micronutrients must also be considered in bee nutritional experiments. Finally, we used pollen metabarcoding to identify pollen from whole larval provisions to understand how much pollen bees used from plants outside of their host plant families to meet their nutritional needs, as well as pollen from individual forager bouts, to observe if bees maintained strict floral constancy or visited multiple plant genera per foraging bout. Whole larval provision surveys revealed a surprising range of host plant pollen use, ranging from ~5% to 70% of host plant pollen per provision. Samples from individual foraging trips contained pollen from multiple genera, suggesting that bees are using some form of foraging decision making. Overall, these results suggest that O. cornifrons have a wide nutritional niche breadth, but while pollen P:L ratio tolerance is broad, a tolerable P:L ratio alone is not enough to create a quality diet for O. cornifrons , and the plant species that make up these diets must also be carefully considered., Competing Interests: The authors have no competing interests to declare., (© 2023 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Seventy five percent of the world's food crops benefit from insect pollination. Hence, there has been increased interest in how global change drivers impact this critical ecosystem service. Because standardized data on crop pollination are rarely available, we are limited in our capacity to understand the variation in pollination benefits to crop yield, as well as to anticipate changes in this service, develop predictions, and inform management actions. Here, we present CropPol, a dynamic, open, and global database on crop pollination. It contains measurements recorded from 202 crop studies, covering 3,394 field observations, 2,552 yield measurements (i.e., berry mass, number of fruits, and fruit density [kg/ha], among others), and 47,752 insect records from 48 commercial crops distributed around the globe. CropPol comprises 32 of the 87 leading global crops and commodities that are pollinator dependent. Malus domestica is the most represented crop (32 studies), followed by Brassica napus (22 studies), Vaccinium corymbosum (13 studies), and Citrullus lanatus (12 studies). The most abundant pollinator guilds recorded are honey bees (34.22% counts), bumblebees (19.19%), flies other than Syrphidae and Bombyliidae (13.18%), other wild bees (13.13%), beetles (10.97%), Syrphidae (4.87%), and Bombyliidae (0.05%). Locations comprise 34 countries distributed among Europe (76 studies), North America (60), Latin America and the Caribbean (29), Asia (20), Oceania (10), and Africa (7). Sampling spans three decades and is concentrated on 2001-2005 (21 studies), 2006-2010 (40), 2011-2015 (88), and 2016-2020 (50). This is the most comprehensive open global data set on measurements of crop flower visitors, crop pollinators and pollination to date, and we encourage researchers to add more datasets to this database in the future. This data set is released for non-commercial use only. Credits should be given to this paper (i.e., proper citation), and the products generated with this database should be shared under the same license terms (CC BY-NC-SA)., (© 2021 The Ecological Society of America.)

Metrics to assess relative adult bee body size have included both mass and morphometrics, but these metrics may not equally or reliably estimate body size for all bee species and in all situations, due to bee age, diet, and/or environment. Understanding the relationships between different metrics and possible redundancies in the information they afford is important but not always known. Body size measurements provide valuable data for interpreting research outcomes for managed solitary bees, including Osmia lignaria Say and Megachile rotundata F. (Hymenoptera: Megachilidae). Applied studies of these important and readily available U.S. crop pollinators focus on refining commercial management practices, and basic empirical studies in various scientific disciplines (from genomics to ecology) employ them as model systems to study solitary bees. To examine common metrics of body size, we measured head capsule width (HCW), intertegular distance (ITD), and fresh and dry weights of newly emerged adults of both species. Using linear and exponential models, we determined relationships between these body size metrics. For M. rotundata , linear models best described relationships between ITD and all other metrics, and between HCW and fresh and dry weights. For O. lignaria , linear models best fit relationships between all metrics except for fresh weight with both ITD and HCW, which were fitted better with exponential models. For both species, model fits were strongest when males and females were pooled. Depending on the study question, knowing that only one metric may reliably measure body size can simplify evaluations of O. lignaria and M. rotundata responses to artificial or environmental variables., Competing Interests: The authors declare that they have no competing interests.

Most of the world's crops depend on pollinators, so declines in both managed and wild bees raise concerns about food security. However, the degree to which insect pollination is actually limiting current crop production is poorly understood, as is the role of wild species (as opposed to managed honeybees) in pollinating crops, particularly in intensive production areas. We established a nationwide study to assess the extent of pollinator limitation in seven crops at 131 locations situated across major crop-producing areas of the USA. We found that five out of seven crops showed evidence of pollinator limitation. Wild bees and honeybees provided comparable amounts of pollination for most crops, even in agriculturally intensive regions. We estimated the nationwide annual production value of wild pollinators to the seven crops we studied at over $1.5 billion; the value of wild bee pollination of all pollinator-dependent crops would be much greater. Our findings show that pollinator declines could translate directly into decreased yields or production for most of the crops studied, and that wild species contribute substantially to pollination of most study crops in major crop-producing regions.

Concerns over the availability of honeybees ( Apis mellifera L.) to meet pollination demands have elicited interest in alternative pollinators to mitigate pressures on the commercial beekeeping industry. The blue orchard bee, Osmia lignaria (Say), is a commercially available native bee that can be employed as a copollinator with, or alternative pollinator to, honeybees in orchards. To date, their successful implementation in agriculture has been limited by poor recovery of bee progeny for use during the next spring. This lack of reproductive success may be tied to an inadequate diversity and abundance of alternative floral resources during the foraging period. Managed, supplementary wildflower plantings may promote O. lignaria reproduction in California almond orchards. Three wildflower plantings were installed and maintained along orchard edges to supplement bee forage. Plantings were seeded with native wildflower species that overlapped with and extended beyond almond bloom. We measured bee visitation to planted wildflowers, bee reproduction, and progeny outcomes across orchard blocks at variable distances from wildflower plantings during 2015 and 2016. Pollen provision composition was also determined to confirm O. lignaria wildflower pollen use. Osmia lignaria were frequently observed visiting wildflower plantings during, and after, almond bloom. Most O. lignaria nesting occurred at orchard edges. The greatest recovery of progeny occurred along the orchard edges having the closest proximity (80 m) to managed wildflower plantings versus edges farther away. After almond bloom, O. lignaria nesting closest to the wildflower plantings collected 72% of their pollen from Phacelia spp., which supplied 96% of the managed floral area. Phacelia spp. pollen collection declined with distance from the plantings, but still reached 17% 800 m into the orchard. This study highlights the importance of landscape context and proximity to supplementary floral resources in promoting the propagation of solitary bees as alternative managed pollinators in commercial agriculture., Competing Interests: None declared., (© 2020 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Osmia lignaria is a commercially available, native solitary bee species recognized for its propensity to forage upon and pollinate tree fruit crops such as apple, almond and cherry. This study evaluated the implementation of O. lignaria co-pollination with honey bees in central Utah commercial tart cherry orchards during 2017 and 2018 bloom. Three paired 1.2 ha sites were selected for evaluation of cherry fruit set and yield with and without managed O. lignaria releases alongside the standard honey bee hive stocking rate of 2.5 hives/ha. Osmia lignaria supplementation did not measurably increase cherry fruit set, fruit per limb cross-sectional area or fruit weight. The lack of differences in yield is likely a consequence of local saturation of pollinator services supplied by managed honey bees throughout experimental orchards, such that no additive benefit of managed O. lignaria releases were measurable. An increase in managed O. lignaria populations was achieved in 2017 but not 2018, possibly due to unknown changes to orchard management or environmental factors. While flying O. lignaria in Utah tart cherries may support sustainable in-field bee propagation, their subsequent impacts on tart cherry yield were not detected when paired with standard stocking densities of honey bees., Competing Interests: The authors declare there are no competing interests.

Current pesticide risk assessment for bees relies on a single (social) species, the western honey bee, Apis mellifera L. (Hymenoptera: Apidae). However, most of the >20,000 bee species worldwide are solitary. Differences in life history traits between solitary bees (SB) and honey bees (HB) are likely to determine differences in routes and levels of pesticide exposure. The objectives of this review are to: 1) compare SB and HB life history traits relevant for risk assessment; 2) summarize current knowledge about levels of pesticide exposure for SB and HB; 3) identify knowledge gaps and research needs; 4) evaluate whether current HB risk assessment schemes cover routes and levels of exposure of SB; and 5) identify potential SB model species for risk assessment. Most SB exposure routes seem well covered by current HB risk assessment schemes. Exceptions to this are exposure routes related to nesting substrates and nesting materials used by SB. Exposure via soil is of particular concern because most SB species nest underground. Six SB species (Hymenoptera: Megachilidae - Osmia bicornis L., O. cornifrons Radoszkowski, O. cornuta Latreille, O. lignaria Say, Megachile rotundata F., and Halictidae - Nomia melanderi Cockerell) are commercially available and could be used in risk assessment. Of these, only N. melanderi nests underground, and the rest are cavity-nesters. However, the three Osmia species collect soil to build their nests. Life history traits of cavity-nesting species make them particularly suitable for semifield and, to a lesser extent, field tests. Future studies should address basic biology, rearing methods and levels of exposure of ground-nesting SB species.

Current pesticide risk assessment practices use the honey bee, Apis mellifera L., as a surrogate to characterize the likelihood of chemical exposure of a candidate pesticide for all bee species. Bees make up a diverse insect group that provides critical pollination services to both managed and wild ecosystems. Accordingly, they display a diversity of behaviors and vary greatly in their lifestyles and phenologies, such as their timing of emergence, degree of sociality, and foraging and nesting behaviors. Some of these factors may lead to disparate or variable routes of exposure when compared to honey bees. For those that possess life histories that are distinct from A. mellifera, further risk assessments may be warranted. In January 2017, 40 bee researchers, representative of regulatory agencies, academia, and agrochemical industries, gathered to discuss the current state of science on pesticide exposure to non-Apis bees and to determine how well honey bee exposure estimates, implemented by different regulatory agencies, may be protective for non-Apis bees. Workshop participants determined that although current risk assessment procedures for honey bees are largely conservative, several routes of exposure are unique to non-Apis bees and warranted further investigation. In this forum article, we discuss these key routes of exposure relevant to non-Apis bees and identify important research gaps that can help inform future bee risk assessment decisions.

Studies of bee movement and activities across a landscape are important for developing an understanding of their behavior and their ability to withstand environmental stress. Recent research has shown that proteins, such as egg albumin, are effective for mass-marking bees. However, current protein mass-marking techniques require sacrificing individual bees during the data collection process. A nonlethal sampling method for protein mark-capture research is sorely needed, particularly for vulnerable, sensitive, or economically valuable species. This study describes a nonlethal sampling method, in which three non-Apis bee species (Bombus bifarius Cresson [Hymenoptera: Apidae], Osmia lignaria Say [Hymenoptera: Megachilidae], and Megachile rotundata Fabricius [Hymenoptera: Megachilidae]) were tested for a unique protein marker by immersing them momentarily in saline buffer and releasing them. Results showed that an egg albumin-specific enzyme-linked immunosorbent assay was 100% effective at detecting the protein on bees that were sampled nonlethally. Furthermore, this sampling method did not have an impact on bee survivorship, suggesting that immersing bees in buffer is a reliable and valid surrogate to traditional, destructive sampling methods for mark-capture bee studies.

Pollination services provided by managed bees are essential for California almond (Prunus dulcis Mill.; Rosales: Rosaceae) production. Currently, pollination needs are met by rented or owned Apis mellifera L. (Hymenoptera: Apidae; honey bee) colonies. Excessive demand on a challenged A. mellifera industry to provide strong colonies in early spring has caused sharp increases in rental prices over the past decade, inviting the consideration of alternative pollinators in addition to, or in place of, A. mellifera. Osmia lignaria Say (Hymenoptera: Megachilidae; the blue orchard bee) is an excellent pollinator of fruit and nut trees, but its pollination impacts when used in tandem with A. mellifera have yet to be evaluated in commercial almond orchards. A 2-yr study was conducted in California orchards to compare almond pollination and production using A. mellifera as sole pollinator to an alternative practice of adding O. lignaria as a co-pollinator with A. mellifera. Almond orchard managerial decisions, such as for pesticide use and irrigation intensity, vary between almond growing regions because of local climates. Therefore, both north-central and southern sites of California's San Joaquin Valley are represented. We compared bee visitation, nut set, and nut yield between orchards and between tree rows within orchards. Also, O. lignaria reproductive success was recorded to assure that these bees remained in the orchards as pollinators and to assess the ability to sustain these bees under regional orchard conditions. We demonstrated that augmenting large commercial almond orchards with O. lignaria can significantly increase nut set and sometimes nut yield in both regions evaluated.

The engineering of flowering agricultural field borders has emerged as a research and policy priority to mitigate threats to pollinators. Studies have, however, rarely addressed the potential that flowering field borders might compete with neighboring crops for pollinator visits if they both are in bloom at the same time, despite this being a concern expressed by growers. We evaluated how wildflower plantings added to orchard borders in a large (512 ha) commercial almond orchard affected honey bee and wild bee visitation to orchard borders and the crop. The study was conducted over two consecutive seasons using three large (0.48 ha) wildflower plantings paired with control orchard borders in a highly simplified agricultural landscape in California. Honey bee (Apis mellifera L.) and wild bee visitation to wildflower plots were at least an order of magnitude higher than to control plots, but increased honey bee visitation to wildflower plots did not lead to any detectable shifts in honey bee visitation to almond flowers in the neighboring orchard. Wild bees were rarely observed visiting almond flowers irrespective of border treatment, indicating a limited short-term potential for augmenting crop pollination using wild bees in highly simplified agricultural landscapes. Although further studies are warranted on bee visitation and crop yield from spatially independent orchards, this study indicates that growers can support bees with alternative forage in almond orchards without risking competition between the wildflower plantings and the crop., (© The Authors 2017. Published by Oxford University Press on behalf of Entomological Society of America. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Contracted commercial beekeeping operations provide an essential pollination service to many agricultural systems worldwide. Increased use of genetically engineered crops in agriculture has raised concerns over pollinator-mediated gene flow between transgenic and conventional agricultural varieties. This study evaluated whether contracted migratory beekeeping practices influence transgenic pollen flow among spatially isolated alfalfa fields. Twelve honey bee (Apis mellifera L.) colonies were permitted to forage on transgenic alfalfa blossoms for 1 wk in Touchet, WA. The hives were then transported 112 km to caged conventional alfalfa plots following one and two nights of isolation (8 and 32 h, respectively) from the transgenic source. Alfalfa seed harvested from the conventional plots was assessed for the presence of the transgene using a new seedling germination assay. We found that 8 h of isolation from a transgenic alfalfa source virtually eliminated the incidence of cross-pollination between the two varieties.

The blue orchard bee, Osmia lignaria (Say), is a solitary bee that is an excellent pollinator of tree fruit orchards. Due to the annual rising costs of honey bee hive rentals, many orchardists are eager to develop management tools and practices to support O. lignaria as an alternative pollinator. Establishing O. lignaria pollination as a sustainable industry requires careful consideration of both bee and orchard management. Here, we test the effect of artificial nest box distribution on in-orchard propagation of O. lignaria in Utah commercial tart cherry orchards. Two nest box distributions were compared across three paired, 1.2-ha plots. One distribution, traditionally employed by O. lignaria consultants, included a centrally located tote for mass-nesting with smaller, surrounding 'satellite' nest boxes at orchard margins. The other distribution was composed of smaller, more equally distributed nest boxes throughout the 1.2-ha plots. Significantly higher propagation of O. lignaria was observed in the latter nest box distribution, although all treatments resulted in bee return exceeding the number of bees initially released. These findings provide support for the use of O. lignaria in tart cherry orchards, and demonstrate how simple changes to bee set-up and management can influence propagation efforts., (Published by Oxford University Press on behalf of the Entomological Society of America 2017. This work is written by US Government employees and is in the public domain in the US.)