Bethanne Bruninga-Socolar, Ph.D.

Managing and restoring tallgrass prairie ecosystem is an important form of pollinator conservation in the Midwestern United States. Prairie reconstruction has been found to enhance native bee diversity and abundance, but it is less clear if prairie reconstruction conserves species thought to be at-risk. We reanalyze a previously published dataset on the bee communities of reconstructed and remnant prairie in the US state of Minnesota to investigate how the abundance of at-risk species respond to local factors, such as floral diversity and prairie type (reconstructed or remnant), and landscape factors, in the form of surrounding agricultural production. We defined at-risk species in two ways. For bumble bees, we used the IUCN red list of bumble bees for North America. As other species in the bee community have not been systematically evaluated, we used an independent data set to calculate a community-level measure of rarity as a proxy for species risk. We calculated community rarity metrics using a Species Weighted Mean (SWM) approach, with species level rarity (relative abundance and site occurrence) derived from a regional dataset comprised of over 30,000 specimens from across the US state of Minnesota. We found that the declining bumble bee Bombus fervidus had higher abundances in remnant rather than reconstructed prairies. Floral richness was associated with rarer bee communities (lower SWM values) across remnant and reconstructed prairies. We show that planting and managing prairies for floral diversity promotes bee communities with rarer species, but that remnants better support some at-risk species such as Bombus fervidus.

Floral constancy of foraging bees influences plant reproduction. Constancy as observed in nature arises from at least four distinct mechanisms frequently confounded in the literature: context-independent preferences for particular plant species, preferential visitation to the same species as the previous plant visited (simple constancy), the spatial arrangement of plants, and the relative abundances of co-flowering species. To disentangle these mechanisms, we followed individual bee flight paths within patches where all flowering plants were mapped, and we used step selection models to estimate how each mechanism influences the probability of selecting any particular plant given the available plants in a multi-species community. We found that simple constancy was positive: bees preferred to visit the same species sequentially. In addition, bees preferred to travel short distances and maintain their direction of travel between plants. After accounting for distance, we found no significant effect of site-level plant relative abundances on bee foraging choices. To explore the importance of the spatial arrangement of plants for bee foraging choices, we compared our full model containing all parameters to one with spatial arrangement removed. Due to bees’ tendency to select nearby plants, combined with strong intraspecific plant clumping, spatial arrangement was responsible for about 50% of the total observed constancy. Our results suggest that floral constancy may be overestimated in studies that do not account for the spatial arrangement of plants, especially in systems with intraspecific plant clumping. Plant spatial patterns at within-site scales are important for pollinator foraging behavior and pollination success.

Prairie restorations are increasingly being used to benefit bees and other pollinators. However, the management practices that best benefit bees remain poorly understood. We surveyed the bees of the Six Mile Marsh Prairie Restoration in Minnetrista, Minnesota, and compared the effects of different management practices on the relative abundance and diversity of bees. The restoration was divided into three sections: haying, burning, and control. Bees were surveyed from 2018–2020 on four transects per section. In total, we collected or observed 2,404 bees from 60 species or morphospecies in 20 genera. Comparing the different management techniques, we found that the haying treatment had significantly higher bee diversity in the final year of sampling. Although the small scale of this study limits the conclusions we can draw, our results suggest that managing prairie restorations by haying could benefit bees.

The relative densities of co-flowering plant species can affect pollinator foraging behavior, i.e., bee choice of which plants to visit in a foraging path. Because animal-pollinated plants depend on sequential pollinator visits to conspecifics for successful reproduction, understanding the relationship between relative densities of co-flowering plants and bee visits within a foraging path is necessary to determine whether co-flowering species facilitate or compete for bee visitation. We examined the role of relative co-flowering plant species densities in determining bee visits to two focal plant species (Astragalus scaphoides and Penstemon albertinus) within foraging paths by following individual bees in natural multispecies co-flowering plant communities. We found that conspecific and heterospecific plant density influenced the proportion of visits to the focal plant species during foraging bouts, but effects of density differed among bee groups. Considering all bees, Lupinus spp. facilitated visits to A. scaphoides while Castilleja spp. competed with A. scaphoides for visits. When three bee groups, Bombus, Anthophora/Eucera, and ‘‘small bees,’’ were modeled separately, Bombus consistently visited A. scaphoides more than expected regardless of its density and P. albertinus less than expected at intermediate densities. Anthophora/Eucera exhibited a nonlinear response to A. scaphoides floral density but visited P. albertinus in direct proportion to its density. ‘‘Small bees’’ visited both plants in direct proportion to their densities. Our results highlight that bee groups should be treated separately in models of pollinator visitation behavior, and that the composition and density of the co-flowering plant community may impact reproductive success of focal plant species.

Restored habitats require long-term management to maintain biodiversity and ensure ecosystem functions. Management strategies are often developed for plant communities, including through seeding and disturbance management, but these actions are taken with a focus on plant dynamics and with little knowledge of the effects on non-plant organisms. Wild bees are often expected to respond to such management actions via their effects on local floral resource availability, but management may also affect bees by altering survival and nesting independently of plant community responses. Working in restoration plantings within a large, actively managed tallgrass prairie preserve, we separated the effects of management and landscape context on bee community abundance and richness from the effects of these covariates on bees mediated through the abundance and richness of the local flowering plant community. We found that bees responded primarily to disturbance management (via bison) and the amount of prairie and forest habitat in the landscape, indicating that across landscapes with relatively abundant flowers and nest-sites, these landscape-level resources are more important than local floral resources for structuring bee communities. In contrast, floral communities responded to restoration age and prescribed burning. Because bees respond to different factors and at a different landscape scale than local plant communities, we conclude that management designed for plants is not sufficient for pollinators. Landscape level restoration may therefore require targeted habitat design and management to successfully restore functionally important animals.

Fire and grazing are historic ecosystem drivers of tallgrass prairie and both are used for restoration management today. The effects of these drivers on animal taxa are still incompletely resolved, especially for wild bees, a growing conservation and restoration priority. Fire and grazing could affect wild bee communities through structural changes to nest site availability via changes to soil conditions, vegetative cover, and availability of plant stems. Here, we sought to determine how different bee nesting groups are affected by the combination of fire and bison grazing management strategies. We grouped bee species by nesting substrate (ground, stem/hole, large cavity) because we expect the availability of these substrates to vary with the application of prescribed fire and grazing. We collected bees in restored and remnant high-quality tallgrass prairie and analyzed whether the proportion of each nesting group within the total bee community was predicted by fire and/or grazing. Ground-nesting bees reached their greatest proportion in bee communities immediately after prescribed fire, but declined proportionally over time since the last burn. Stem-/hole-nesting bees reached their highest proportion in the bee community with infrequent fire (6-year interval) and differed in their response to fire depending on the presence/absence of bison. Sampling year affected bee nesting groups and we found that nesting groups did not change in concert (i.e. different nesting groups had different good and bad years from each other). Our results show that spatiotemporal variation of prescribed fire and bison grazing is essential for conservation of multiple bee nesting groups.

Effective monitoring is necessary to provide robust detection of bee declines. In the United States and worldwide, bowl traps have been increasingly used to monitor native bees and purportedly detect declines. However, bowl traps have a suite of flaws that make them poorly equipped to monitor bees. We outline the drawbacks of bowl traps, as well as other passive sampling methods. We emphasize that current methods do not monitor changes in bee abundance. We then propose future approaches to improve bee monitoring efforts, which include improving our understanding of the efficacy and drawbacks of current methods, novel molecular methods, nest censusing, mark-recapture, sampling of focal plant taxa, and detection of range contractions. Overall, we hope to highlight deficiencies of the current state of bee monitoring, with an aim to stimulate research into the efficacy of existing methods and promote novel methods that provide meaningful data that can detect declines without squandering limited resources.

Ecological restoration is increasingly implemented to reverse habitat loss and concomitant declines in biological diversity. Typically, restoration success is evaluated by measuring the abundance and/or diversity of a single taxon. However, for a restoration to be successful and persistent, critical ecosystem functions such as animal-mediated pollination must be maintained. In this review, we focus on three aspects of pollination within ecological restorations. First, we address the need to measure pollination directly in restored habitats. Proxies such as pollinator abundance and richness do not always accurately assess pollination function. Pollen supplementation experiments, pollen deposition studies, and pollen transport networks are more robust methods for assessing pollination function within restorations. Second, we highlight how local-scale management and landscape-level factors may influence pollination within restorations. Localscale management actions such as prescribed fire and removal of non-native species can have large impacts on pollinator communities and ultimately on pollination services. In addition, landscape context including proximity and connectivity to natural habitats may be an important factor for land managers and conservation practitioners to consider to maximize restoration success. Third, as climate change is predicted to be a primary driver of future loss in biodiversity, we discuss the potential effects climate change may have on animal-mediated pollination within restorations. An increased mechanistic understanding of how climate change affects pollination and incorporation of climate change predictions will help practitioners design stable, functioning restorations into the future.

Restoration efforts often focus on plants, but additionally require the establishment and long-term persistence of diverse groups of nontarget organisms, such as bees, for important ecosystem functions and meeting restoration goals. We investigated long-term patterns in the response of bees to habitat restoration by sampling bee communities along a 26-year chronosequence of restored tallgrass prairie in north-central Illinois, U.S.A. Specifically, we examined how bee communities changed over time since restoration in terms of (1) abundance and richness, (2) community composition, and (3) the two components of beta diversity, one-to-one species replacement, and changes in species richness. Bee abundance and raw richness increased with restoration age from the low level of the pre-restoration (agricultural) sites to the target level of the remnant prairie within the first 2–3 years after restoration, and these high levels were maintained throughout the entire restoration chronosequence. Bee community composition of the youngest restored sites differed from that of prairie remnants, but 5–7 years post-restoration the community composition of restored prairie converged with that of remnants. Landscape context, particularly nearby wooded land, was found to affect abundance, rarefied richness, and community composition. Partitioning overall beta diversity between sites into species replacement and richness effects revealed that the main driver of community change over time was the gradual accumulation of species, rather than one-to-one species replacement. At the spatial and temporal scales we studied, we conclude that prairie restoration efforts targeting plants also successfully restore bee communities.

Animal-pollinated plants depend on sequential pollinator visits to conspecifics for successful reproduction. Therefore, in co-flowering plant communities, the proportion of visits to a focal plant species in individual pollinator foraging bouts determines reproductive outcomes for that species. We investigated the factors determining bee visits to the plant Astragalus scaphoides within foraging bouts in a natural multispecies community in the northern Rocky Mountains. We found that both conspecific and heterospecific floral density influenced the proportion of visits to A. scaphoides during foraging bouts, but these effects of floral density differed among two abundant bee groups. Our field observations reject the null expectation that bees visit plant species in direct proportion to their relative floral densities. Bombus consistently visited A. scaphoides more than expected, while solitary bees of the genera Anthophora and Eucera exhibited a nonlinear response to floral density.