Key Pollinators Research Articles

Control of two-spotted spider mite, Tetranychus urticae, on strawberry by integrating with cyetpyrafen and Phytoseiulus persimilis

The two-spotted spider mite (TSSM, Tetranychus urticae Koch) is a significant agricultural pest, particularly in strawberries. Management of TSSM has traditionally relied on synthetic acaricides, but to mitigate dependency on these chemicals, the control of TSSM on strawberry is often combined with biological control measures and chemical control strategies. The predatory mite, Phytoseiulus persimilis, is a promising biological control agent, preying on all TSSM developmental stages. In this study, we examined the toxicity of six common acaricides on TSSM and P. persimilis, and cyetpyrafen was selected due to its highest relative toxicity value. Then, we examined the compatibility of cyetpyrafen with P. persimilis for TSSM management on strawberries. The results suggested that cyetpyrafen revealed no substantial differences in prey consumption or longevity when compared to the control, though minor effects on the development durations of protonymphs and deutonymphs were noted in the subsequent generation. Additionally, cyetpyrafen's toxicity on key pollinators, such as Apis mellifera and Bombus terrestris, was found to be low. Thus, an integrated strategy combining cyetpyrafen (0.83 mg/L) with P. persimilis (predator–prey ratio of 1:30) was examined under laboratory and field conditions. Laboratory trials demonstrated a reduction in mites per leaf from 32.72 to 14.50 within 3 days, correlating to a 70.23% control efficiency. This efficacy increased to 96.04% by day 9 and was sustained until the experiment concluded on day 27. Field trials similarly showed a reduction in TSSM from 53.93 to 9.63 mites/leaf by day 6, achieving an 83.64% control efficiency, and culminated in a 98.46% reduction by day 10. These findings suggested that an integrated approach utilizing cyetpyrafen in conjunction with P. persimilis can be an effective alternative for TSSM management on strawberry plants.

A multi-taxonomic survey to determine the conservation status of native pollinators

The worldwide decline of insects calls for understanding the conservation status of key insect groups at local and regional scales. Pollinators are widely recognized as especially important to native ecosystems and agriculture and their declines have been met with calls for inventory and monitoring. In New York State, we conducted a four-year survey effort to document the distribution and status of key native pollinators in four insect orders: Coleoptera, Diptera, Hymenoptera, and Lepidoptera. In this paper, we describe the process of designing and implementing the study in the hopes that other jurisdictions could follow a similar model. We combined systematic field inventories, surveys of special habitats, target species surveys, community science observations, and museum and partner data to create a comprehensive look at the distribution of each of 457 species in two time periods (2000 to present and 1999 and earlier). We used the conservation status ranking system developed by NatureServe and generated ranks and a series of distribution maps and phenology charts for each species. We describe the effort needed to carry out the study, as well as reasons for its success and areas for improvement.

Exposure to sublethal concentrations of imidacloprid, pyraclostrobin, and glyphosate harm the behavior and fat body cells of the stingless bee Scaptotrigona postica

Pesticide use in agriculture threatens non-target insects such as bees. Considering the ecological and economic relevance of native bees, such as Scaptotrigona postica, and the insufficient studies on the effects of pesticides on their behavior and physiology, improving the current knowledge on this issue is essential. Therefore, this study investigated the sublethal effects of imidacloprid, pyraclostrobin, and glyphosate on the behavior and fat body cells of S. postica. Pesticide ingestion decreased the walking distance and mean velocity of bees compared to the control and solvent control groups. The oenocytes of the control groups were spherical, with central nuclei containing decondensed chromatin, and the trophocytes presented irregular morphology, with cells varying in shape and the cytoplasm filled with vacuoles and granules. However, bees exposed to pesticides showed extensive cytoarchitectural disruption in the fat body, such as vacuolization and shape changes in oenocytes and altered nuclei morphology in trophocytes. Moreover, pesticide exposure increased the number of atypical oenocytes and altered trophocytes, except for the PYR group, which showed a lower number of atypical oenocytes. Caspase-positive labeling significantly increased in all exposed bee groups. Alternatively, TLR4 labeling was significantly decreased in the exposed groups compared to the control groups. There was a significant increase in HSP90 immunolabeling in all exposed groups compared to the control. These findings reinforce the importance of research on the sublethal effects of low pesticide concentrations on key neotropical pollinators and prove that these toxic substances can impair their detoxification and immune defense.

Land-use-associated stressors interact to reduce bumblebee health at the individual and colony level.

In agricultural landscapes, bees face a variety of stressors, including insecticides and poor-quality food. Although both stressors individually have been shown to affect bumblebee health negatively, few studies have focused on stressor interactions, a scenario expected in intensively used agricultural landscapes. Using the bumblebee Bombus terrestris, a key pollinator in agricultural landscapes, we conducted a fully factorial laboratory experiment starting at nest initiation. We assessed the effects of food quality and insecticides, alone and in interaction, on health traits at various levels, some of which have been rarely studied. Pollen with a diluted nutrient content (low quality) reduced ovary size and delayed colony development. Wing asymmetry, indicating developmental stress, was increased during insecticide exposure and interactions with poor food, whereas both stressors reduced body size. Both stressors and their interaction changed the workers' chemical profile and reduced worker interactions and the immune response. Our findings suggest that insecticides combined with nutritional stress reduce bumblebee health at the individual and colony levels, thus possibly affecting colony performance, such as development and reproduction, and the stability of plant-pollinator networks. The synergistic effects highlight the need of combining stressors in risk assessments and when studying the complex effects of anthropogenic stressors on health outcomes.

Limited introgression from non-native commercial strains and signatures of adaptation in the key pollinator Bombus terrestris.

Insect pollination is fundamental for natural ecosystems and agricultural crops. The bumblebee species Bombus terrestris has become a popular choice for commercial crop pollination worldwide due to its effectiveness and ease of mass rearing. Bumblebee colonies are mass produced for the pollination of more than 20 crops and imported into over 50 countries including countries outside their native ranges, and the risk of invasion by commercial non-native bumblebees is considered an emerging issue for global conservation and biological diversity. Here, we use genome-wide data from seven wild populations close to and far from farms using commercial colonies, as well as commercial populations, to investigate the implications of utilizing commercial bumblebee subspecies in the UK. We find evidence for generally low levels of introgression between commercial and wild bees, with higher admixture proportions in the bees occurring close to farms. We identify genomic regions putatively involved in local and global adaptation, and genes in locally adaptive regions were found to be enriched for functions related to taste receptor activity, oxidoreductase activity, fatty acid and lipid biosynthetic processes. Despite more than 30 years of bumblebee colony importation into the UK, we observe low impact on the genetic integrity of local B. terrestris populations, but we highlight that even limited introgression might negatively affect locally adapted populations.

Temperature during pupal development affects hoverfly developmental time, adult life span, and wing length.

Hoverflies (Diptera, Syrphidae) are cosmopolitan, generalist flower visitors and among the most important pollinators after bees and bumblebees. The dronefly Eristalis tenax can be found in temperate and continental climates across the globe, often synanthropically. Eristalis tenax pupae of different generations and different climate zones are thus exposed to vastly different temperatures. In many insects, the ambient temperature during the pupal stage affects development, adult size, and survival; however, the effect of developmental temperature on these traits in hoverflies is comparatively poorly understood. We here reared E. tenax pupae at different temperatures, from 10°C to 25°C, and quantified the effect on adult hoverflies. We found that pupal rearing at 17°C appeared to be optimal, with high eclosion rates, longer wings, and increased adult longevity. Rearing temperatures above or below this optimum led to decreased eclosion rates, wing size, and adult survival. Similar thermal dependence has been observed in other insects. We found that rearing temperature had no significant effect on locomotor activity, coloration or weight, despite evidence of strong sexual dimorphism for each of these traits. Our findings are important as hoverflies are key pollinators, and understanding the effects of developmental temperature could potentially be useful for horticulture.

The first ITS2 sequence data set of eDNA from honey of Malaysian giant honeybees (Apis dorsata) and stingless bees (Heterotrigona itama) reveals plant species diversity

ObjectivesPollen is a useful tool for identifying the provenance and complex ecosystems surrounding honey production in Malaysian forests. As native key pollinators in Malaysia, Apis dorsata and Heterotrigona itama forage on various plant/pollen species to collect honey. This study aims to generate a dataset that uncovers the presence of these plant/pollen species and their relative abundance in the honey of A. dorsata and H. itama. The information gathered from this study can be used to determine the geographical and botanical origin and authenticity of the honey produced by these two species.ResultsSequence data were obtained for both A. dorsata and H. itama. The raw sequence data for A. dorsata was 5 Mb, which was assembled into 5 contigs with a size of 6,098,728 bp, an N50 of 15,534, and a GC average of 57.42. Similarly, the raw sequence data for H. itama was 6.3 Mb, which was assembled into 11 contigs with a size of 7,642,048 bp, an N50 of 17,180, and a GC average of 55.38. In the honey sample of A. dorsata, we identified five different plant/pollen species, with only one of the five species exhibiting a relative abundance of less than 1%. For H. itama, we identified seven different plant/pollen species, with only three of the species exhibiting a relative abundance of less than 1%. All of the identified plant species were native to Peninsular Malaysia, especially the East Coast area of Terengganu.Data descriptionOur data offers valuable insights into honey’s geographical and botanical origin and authenticity. Metagenomic studies could help identify the plant species that honeybees forage and provide preliminary data for researchers studying the biological development of A. dorsata and H. itama. The identification of various flowers from the eDNA of honey that are known for their medicinal properties could aid in regional honey with accurate product origin labeling, which is crucial for guaranteeing product authenticity to consumers.

Fine-mesh exclusion netting reduces Drosophila suzukii (Diptera: Drosophilidae) infestation and improves organic fall-bearing raspberry yields.

The economical production of small fruits has been significantly complicated by the spotted-wing drosophila, Drosophila suzukii Matsumura, throughout its invaded rage. Fall-bearing red raspberries are especially susceptible to D. suzukii, and significant efforts to mitigate their damage are undertaken by growers. Exclusion barriers made from fine-mesh netting has emerged as an organic compliant strategy to mitigate damage from D. suzukii. Identifying less susceptible raspberry cultivars may also alleviate D. suzukii damage. Key pollination complications arise from exclusion-based tactics, but properly timing exclusion establishment may provide a remedy. In a 2-yr-old fall-bearing raspberry planting containing 3 cultivars, exclusion barriers were erected at 2 phenological timings. Spinosad- and pyrethrin-based pesticides were also applied to separate rows throughout harvest and a fourth treatment group matured without management. A subsection of raspberries was harvested and examined for marketability or damage, and D. suzukii adult populations were sampled with baited traps. An exclusion barrier applied early in fruit development was the most effective pest management strategy of those we tested. Its use decreased D. suzukii captures by 75%, decreased D. suzukii-infested fruit weights by 48%, and increased marketable yield by 63% compared to the no management control; however, it also increased pollination deficient fruit weights. Exclusion netting applied later in fruit maturation also decreased D. suzukii captures, infested fruit weights, and the weight of fruits damaged by other arthropod pests but did not significantly increase marketable yield. Our results indicate that netting exclusions may hold promise for the economical production of fall-bearing raspberries.

What makes a good pollinator? Abundant and specialised insects with long flight periods transport the most strawberry pollen

Abstract Despite the importance of insect pollination to produce marketable fruits, insect pollination management is limited by insufficient knowledge about key crop pollinator species. This lack of knowledge is due in part to (1) the extensive labour involved in collecting direct observations of pollen transport, (2) the variability of insect assemblages over space and time and (3) the possibility that pollinators may need access to wild plants as well as crop floral resources. We address these problems using strawberry in the United Kingdom as a case study. First, we compare two proxies for estimating pollinator importance: flower visits and pollen transport. Pollen‐transport data might provide a closer approximation of pollination service, but visitation data are less time‐consuming to collect. Second, we identify insect parameters that are associated with high importance as pollinators, estimated using each of the proxies above. Third, we estimated insects' use of wild plants as well as the strawberry crop. Overall, pollinator importances estimated based on easier‐to‐collect visitation data were strongly correlated with importances estimated based on pollen loads. Both frameworks suggest that bees (Apis and Bombus) and hoverflies (Eristalis) are likely to be key pollinators of strawberries, although visitation data underestimate the importance of bees. Moving beyond species identities, abundant, relatively specialised insects with long active periods are likely to provide more pollination services. Most insects visiting strawberry plants also carried pollen from wild plants, suggesting that pollinators need diverse floral resources. Identifying essential pollinators or pollinator parameters based on visitation data will reach the same general conclusions as those using pollen transport data, at least in monoculture crop systems. Managers may be able to enhance pollination service by preserving habitats surrounding crop fields to complement pollinators' diets and provide habitats for diverse life stages of wild pollinators.

Bee pollination services and the burden of biogeography.

Native bees augment pollination services in the Northern Hemisphere, especially cultivated apple crops, yet Southern Hemisphere contexts are poorly known. We observed the foraging behaviour of 69 354 invertebrate flower visitors in Australian orchards (two regions, 3 years) to assess the efficacy of pollination service (Peff). Native stingless bees and introduced honey bees were the most abundant visitors and most efficacious pollinators (Tetragonula Peff = 6.16; Apis Peff = 13.02), with Tetragonula becoming important service providers above 22°C. However, visits by tree-nesting stingless bees decreased with distance from native forest (less than 200 m) and their tropical/subtropical distribution precludes pollination service in other major Australian apple-producing regions. More broadly distributed native allodapine and halictine bees transferred the most pollen per-visit, but their low abundances reduce efficacies (Exoneura Peff = 0.03; Lasioglossum Peff = 0.06), resulting in a general dependence on honey bees. This reliance is a burden of biogeography, since key Northern Hemisphere pollinators of apple (Andrena, Apis, Bombus, Osmia) do not naturally occur in Australasia-where there is only 15% generic overlap with Central Asian bees sympatric with wild apple distributions (cf. Palaearctic 66% and Nearctic 46% generic overlaps). The historical biogeography of bees therefore drives an extreme dependence on one introduced species for apple pollination in Australia.

Location and Creation of Nest Sites for Ground-Nesting Bees in Apple Orchards.

Wild ground-nesting bees are key pollinators of apple (Malus domestica). We explored, (1) where they choose to nest, (2) what influences site selection and (3) species richness in orchards. Twenty-three orchards were studied over three years; twelve were treated with additional herbicide to increase bare ground with the remainder as untreated controls. Vegetation cover, soil type, soil compaction, nest number and location, and species were recorded. Fourteen species of ground-nesting solitary/eusocial bee were identified. Most nests were in areas free of vegetation and areas treated with additional herbicide were utilised by ground nesting bees within three years of application. Nests were also evenly distributed along the vegetation-free strips underneath the apple trees. This area was an important ground-nesting bee habitat with mean numbers of nests at peak nest activity of 873 per ha (range 44-5705), and 1153 per ha (range 0-4082) in 2018 and 2019, respectively. Increasing and maintaining areas of bare ground in apple orchards during peak nesting events could improve nesting opportunities for some species of ground-nesting bee and, combined with flowers strips, be part of a more sustainable pollinator management approach. The area under the tree row is an important contributor to the ground-nesting bee habitat and should be kept bare during peak nesting.

Robber flies and hover flies (Insecta, Diptera, Asilidae and Syrphidae) in beech forests of the central Apennines: a contribution to the inventory of insect biodiversity in Italian State Nature Reserves.

The present paper describes a sampling-event dataset on species belonging to two families of Diptera (Syrphidae and Asilidae) collected between 2012 and 2019 in two Italian beech forests located in the central Apennines. The reference dataset consists of an annotated checklist and has been published on Zenodo. Syrphidae and Asilidae are two widespread and key ecological groups, including predator, pollinator and saproxylic species. Despite their pivotal role in both natural and man-made ecosystems, these families are still poorly known in terms of local distribution and open-access sampling-event data are rare in Italy. This open-access dataset includes 2,295 specimens for a total of 21 Asilidae and 65 Syrphidae species. Information about the collection (e.g. place, date, methods applied, collector) and the identification (e.g. species name, author, taxon ID) of the species is provided. Given the current biodiversity crisis, the publication of checklists, sampling-event data and datasets on insect communities in open-access repositories is highly recommended, as it represents the opportunity to share biodiversity information amongst different stakeholders. Moreover, such data are also a valuable source of information for nature reserve managers responsible for monitoring the conservation status of protected and endangered species and habitats and for evaluating the effects of conservation actions over time.

Provisioning Australian Seed Carrot Agroecosystems with Non-Floral Habitat Provides Oviposition Sites for Crop-Pollinating Diptera

The addition of floral resources is a common intervention to support the adult life stages of key crop pollinators. Fly (Diptera) crop pollinators, however, typically do not require floral resources in their immature life stages and are likely not supported by this management intervention. Here, we deployed portable pools filled with habitat (decaying plant materials, soil, water) in seed carrot agroecosystems with the intention of providing reproduction sites for beneficial syrphid (tribe Eristalini) fly pollinators. Within 12 to 21 days after the pools were deployed, we found that the habitat pools supported the oviposition and larval development of two species of eristaline syrphid flies, Eristalis tenax (Linnaeus, 1758) and Eristalinus punctulatus (Macquart, 1847). Each habitat pool contained an average (±S.E.) of 547 ± 117 eristaline fly eggs and 50 ± 17 eristaline fly larvae. Additionally, we found significantly more eggs were laid on decaying plant stems and carrot roots compared to other locations within the pool habitat (e.g., on decaying carrot umbels, leaves, etc.). These results suggest that deploying habitat pools in agroecosystems can be a successful management intervention that rapidly facilitates fly pollinator reproduction. This method can be used to support future studies to determine if the addition of habitat resources on intensively cultivated farms increases flower visitation and crop pollination success by flies.

Signatures of Adaptation, Constraints, and Potential Redundancy in the Canonical Immune Genes of a Key Pollinator.

All organisms require an immune system to recognize, differentiate, and defend against pathogens. From an evolutionary perspective, immune systems evolve under strong selective pressures exerted by fast-evolving pathogens. However, the functional diversity of the immune system means that different immune components and their associated genes may evolve under varying forms of selection. Insect pollinators, which provide essential ecosystem services, are an important system in which to understand how selection has shaped immune gene evolution as their populations are experiencing declines with pathogens highlighted as a potential contributing factor. To improve our understanding of the genetic variation found in the immune genes of an essential pollinator, we performed whole-genome resequencing of wild-caught Bombus terrestris males. We first assessed nucleotide diversity and extended haplotype homozygosity for canonical immune genes finding the strongest signatures of positive selection acting on genes involved in pathogen recognition and antiviral defense, possibly driven by growing pathogen spread in wild populations. We also identified immune genes evolving under strong purifying selection, highlighting potential constraints on the bumblebee immune system. Lastly, we highlight the potential loss of function alleles present in the immune genes of wild-caught haploid males, suggesting that such genes are potentially less essential for development and survival and represent redundancy in the gene repertoire of the bumblebee immune system. Collectively, our analysis provides novel insights into the recent evolutionary history of the immune system of a key pollinator, highlighting targets of selection, constraints to adaptation, and potential redundancy.

GeoDanceHive: An Operational Hive for Honeybees Dances Recording

Honeybees are known for their ability to communicate about resources in their environment. They inform the other foragers by performing specific dance sequences according to the spatial characteristics of the resource. The purpose of our study is to provide a new tool for honeybees dances recording, usable in the field, in a practical and fully automated way, without condemning the harvest of honey. We designed and equipped an outdoor prototype of a production hive, later called "GeoDanceHive", allowing the continuous recording of honeybees' behavior such as dances and their analysis. The GeoDanceHive is divided into two sections, one for the colony and the other serving as a recording studio. The time record of dances can be set up from minutes to several months. To validate the encoding and sampling quality, we used an artificial feeder and visual decoding to generate maps with the vector endpoints deduced from the dance information. The use of the GeoDanceHive is designed for a wide range of users, who can meet different objectives, such as researchers or professional beekeepers. Thus, our hive is a powerful tool for honeybees studies in the field and could highly contribute to facilitating new research approaches and a better understanding landscape ecology of key pollinators.

Optimizing low-cost sampling of pollinator insects in oilseed rape fields

Insects are key pollinators to ecosystem function, but much work remains to determine the most cost-effective, reliable scheme to monitor them. Pan traps (PT) and flight interception traps (FIT) are two of the most popular insect sampling methods used. However, their relative sampling performance and cost is poorly known for agroecosystems in China. We conducted a study across 18 oilseed rape fields in smallholder farmland in Zhejiang, China using these two traps. Our results showed that a single FIT had a greater sampling efficiency (more individuals and higher species richness) than a single PT, but controlling for cost, four PTs (the cost for four PTs is close to one FIT) showed a greater sampling efficiency than FITs. PTs collected more small-bodied individuals while FITs and PTs did not significantly differ in terms of monitoring pollinator insects with large body size. When exploring whether semi-natural habitat embedded in the agricultural landscape affected these results, results from both trap types shows that semi-natural habitat had a significant positive impact on wild pollinator diversity and rarefied species richness. Future studies that examine the effects of agricultural landscape on the wild pollinator community should combine PTs with netting or other active methods for long-term wild pollinator monitoring strategies.

Forest and grassland habitats support pollinator diversity more than wildflowers and sunflower monoculture

Abstract Intensively managed agricultural landscapes often lack suitable habitats to support diverse wildlife, particularly harming pollinator communities. Besides mass flowering crops, remnant patches of natural and semi‐natural vegetation may play a key role in maintaining and conserving biodiversity. Yet, the effects of different natural habitats, including forests and grasslands, on different pollinator communities are poorly understood at the landscape scale. We examined the abundance, richness, and diversity of wild bees and hoverflies, two key pollinator groups, across a land‐use gradient spanning forest edges, grassland, wildflower strips, and sunflower monoculture. We also examined the distribution of hoverfly larvae trophic guilds and wild bee nesting traits across the above‐mentioned land‐use gradient. Finally, we evaluated the impact of landscape structure (forest, grassland, and water cover in the surrounding landscape) on pollinator community composition. Our results indicate that forest and grassland habitats supported a higher abundance and greater richness of pollinators than wildflower strips and sunflower monocultures. Furthermore, hoverflies were more sensitive to habitat and floristic homogenization than wild bees. Sunflower and wildflower habitats also hosted a lower diversity of larvae trophic guilds and wild bee nesting guilds as compared to forests and grasslands. Our study suggests that conserving and restoring forest and grassland habitats within agricultural mosaics may serve as the main ‘refuge’ for wild pollinators.

Understanding climate change effects on the potential distribution of an important pollinator species, Ceratina moerenhouti (Apidae: Ceratinini), in the Eastern Afromontane biodiversity hotspot, Kenya

Monitoring key pollinator taxa such as the genus Ceratina requires precise near real-time predictions to facilitate better surveillance. The potential habitat suitability of Ceratina moerenhouti was predicted in the Eastern Afromontane biodiversity hotspot (EABH) in Kenya using presence-only data, to identify their potential distribution and vulnerability due to climate change. Bioclimatic, edaphic, terrain, land surface temperature, and land use and land cover (LULC) variables were used as predictors. Three machine learning techniques, together with their ensemble model, were evaluated on their suitability to predict current and future (the shared socioeconomic pathways (SSPs), i.e., SSP245 and SSP585) habitat suitability. Predictors were subjected to variable selection using the variance inflation factor resulting in a few (n = 9) optimum variables. The area under the curve (AUC) and true skill statistic (TSS) were used for the accuracy assessment of the modeling outputs. The results indicated that 30% and 10% of the EABH in Murang'a and Taita Taveta counties are currently suitable for C. moerenhouti occurrence, respectively. However, future projections show a ±5% decrease in C. moerenhouti habitats in the two counties. Further, the ensemble model harnessed the algorithm differences while the random forest had the highest individual predictive power (AUC = 0.97; TSS = 0.96). Clay content, LULC, and the slope were the most relevant variables together with temperature and precipitation. Integrating multi-source data in predicting suitable habitats improves model prediction capacity. This study can be used to support the maintenance of flowering plant communities around agricultural areas to improve pollination services.

Effectiveness landscape of crop pollinator assemblages: Implications to pollination service management

There is a growing consensus that the world is facing a pollination crisis. To mitigate crop pollination deficits, some management strategies include the massive introduction of managed bee species, yet quite often they are applied blindly, as information on crop pollination effectiveness for each single pollinator species of assemblages is usually not available. Therefore, the introduction on managed species is not always the best option to improve crop yields. Here, by using the highly pollinator-dependent alfalfa crop (Medicago sativa L.) as a case study, we propose the use of the effectiveness landscape framework to identify key crop pollinator species. According to this framework, in a mutualistic interaction, each species´ effectiveness is represented by the product of a quantitative component and a qualitative one, these being measures of the outcomes of this interaction. We applied this framework for two managed and four wild bee species that visit alfalfa in fields southwest of Buenos Aires province, Argentina. We dissected the quantity components of the pollinator effectiveness landscape by estimating two quantitative subcomponents: visitation rate and flower tripping rate. Also, we estimate pod set as a qualitative component without dissecting it in subcomponents. Our results showed that the contribution of both components and the resulting pollinator effectiveness varied among pollinator species, indicating a contrasting effectiveness of different bee species on alfalfa pollination. For example, pollinator effectiveness was higher for managed than for wild bees, as consequence of their very high visitation rate, however, wild bee flower tripping rate and pod set were as high as managed ones. In fact, wild bees were more effective in promoting flower tripping than one of the managed bees (A. mellifera). This approach allowed us to assess which effectiveness components and subcomponents make pollinator species more or less effective, thus providing valuable information to identify key species to be enhanced to help in closing yield gaps. We suggest that the application of the effectiveness landscape framework would be useful to develop strategies to improve crop pollination service in pollinator-dependent crop systems.

Range‐wide floral trait variation reflects shifts in pollinator assemblages, consistent with pollinator‐mediated divergence despite generalized visitation

Floral trait evolution mediated by pollinators is important in the diversification of flowering plants, yet few studies have demonstrated the range‐wide geographic variation in both floral traits and pollinators which represents a predicted precursor for pollinator‐mediated speciation. This study explores whether geographic variation in pollinator interactions underlies the observed patterns of floral divergence both 1) among species of theCastilleja purpureacomplex (C. purpurea,C. citrinaandC. lindheimeri) and the congenerC. sessiliflora, as well as 2) withinC. sessiliflora, across its wide geographic range. We sampled floral visitors and floral traits (morphology and color) at 23 populations across a 1900 km‐wide study area in 1–3 years, with reproductive fitness (fruit set) data for 18 of these populations. A wide diversity of pollinator functional groups visited the focal species, including bees, butterflies, hawkmoths and hummingbirds, and visitor assemblages varied among species and across geography. We identified relationships between floral traits and visitation by certain pollinator groups, which often aligned with predictions based on pollination syndromes. Despite visitor assemblages being largely generalized across most populations, we found that the observed changes in floral traits were associated with shifts in the relative frequencies of key pollinator functional groups. Hence this study demonstrates that variation in pollinator assemblages across the distributions of taxa may underlie divergence in floral traits and suggests that highly specialized relationships may not be required for early stages of pollinator‐mediated floral divergence. Our extensive sampling of 23 populations over multiple years across a large geographic area highlights the value of range‐wide studies for characterizing patterns of divergence mediated by ecological interactions.