Influence of landscape context on the abundance and diversity of bees in Mediterranean olive groves

The diversity and abundance of wild bees ensures the delivery of pollination services and the maintenance of ecosystem diversity. As previous studies carried out in Central Europe and the US have shown, bee diversity and abundance is influenced by the structure and the composition of the surrounding landscape. Comparable studies have so far not been carried out in the Mediterranean region. The present study examines the influence of Mediterranean landscape context on the diversity and abundance of wild bees. To do this, we sampled bees in 13 sites in olive groves on Lesvos Island, Greece. Bees were assigned to five categories consisting of three body size groups (small, medium and large bees), the single most abundant bee species (Lasioglossum marginatum) and all species combined. The influence of the landscape context on bee abundance and species richness was assessed at five radii (250, 500, 750, 1000 and 1250 m) from the centre of each site. We found that the abundance within bee groups was influenced differently by different landscape parameters and land covers, whereas species richness was unaffected. Generally, smaller bees' abundance was impacted by landscape parameters at smaller scales and larger bees at larger scales. The land cover that influenced bee abundance positively was olive grove, while phrygana, conifer forest, broad-leaved forest, cultivated land, rock, urban areas and sea had mostly negative or no impact. We stress the need for a holistic approach, including all land covers, when assessing the effects of landscape context on bee diversity and abundance in the Mediterranean.     

Population genetics of wild and managed pollinators: implications for crop pollination and the genetic integrity of wild bees

Loss of habitat and chemical use associated with agriculture can cause population declines of wild pollinators. Less is known about the evolutionary consequences of interactions between species used in commercial agriculture and wild pollinators. Given population declines of many wild bee species, it is crucial to understand if commercial queens become established in natural areas, if wild bees visit agricultural fields and have the potential to interact with commercial bees, and if gene flow occurs between commercial and wild bees. We drew on a long-term data set that documents commercial bumble bee (Bombus impatiens) use in New England, and we conducted genetic analyses of foraging B. impatiens from areas with varying intensities of commercial bee use. In agricultural areas with a history of commercial bee use we also sampled bees directly from commercial hives. We found significant genetic differences among foraging B. impatiens and B. impatiens sampled directly from hives (average pairwise F′_ST = 0.14), but not among samples of foraging bees from natural areas (average F′_ST among foraging bees?=?0.002). Furthermore, Bayesian analysis of population structure revealed that foraging bees caught in areas with a history of commercial bee use grouped with samples from natural areas. These results document an agricultural setting where there was no widespread introgression of alleles from commercial bumble bees to wild bumble bees, commercial bumble bees did not become established in natural areas, and wild bees were providing pollination services to crops.     

Spatio-temporal variation in species assemblages in field edges: seasonally distinct responses of solitary bees to local habitat characteristics and landscape conditions

The bees (Hymenoptera: Apiformes) are important pollinators in many ecosystems, but their diversity has declined in Europe during the past century, mainly due to habitat loss. However, some of the habitat requirements of wild bees are met in anthropogenic landscape elements, such as road sides, power-line strips and field edges. Moreover, as the bee species assemblages change throughout the season the habitat requirements of the bee fauna may change accordingly. Understanding such seasonally distinct responses of solitary bees with different phenologies may be of high value for local conservation planning. The purpose of this study was to examine if the habitat quality of field edges for solitary bees change throughout the season, and how this temporal variation relates to local habitat and landscape conditions. By sampling solitary bees in 18 field edges in southeast Norway throughout the season we found that the species richness and abundance of bees was highest in sun exposed field edges, independently of the season. However, we found phenologically distinct responses to the landscape context. Moreover, field edges situated in landscapes with a high proportion of forests and semi-natural landscape elements hosted the most phenologically diverse bee species assemblages. We conclude that in order to fulfil the habitat requirements of bee species assemblages throughout the season, one should conserve and direct habitat restoration schemes towards increasing sun exposure at field edges with a diverse flora and a high proportion of semi-natural areas in the vicinity.     

Nesting success of wood‐cavity‐nesting bees declines with increasing time since wildfire

Bees require distinct foraging and nesting resources to occur in close proximity. However, spatial and temporal patterns in the availability and quantity of these resources can be affected by disturbances like wildfire. The potential for spatial or temporal separation of foraging and nesting resources is of particular concern for solitary wood‐cavity‐nesting bees as they are central‐place, short‐distance foragers once they have established their nest. Often the importance of nesting resources for bees have been tested by sampling foraging bees as a proxy, and nesting bees have rarely been studied in a community context, particularly postdisturbance. We tested how wood‐cavity‐nesting bee species richness, nesting success, and nesting and floral resources varied across gradients of wildfire severity and time‐since‐burn. We sampled nesting bees via nesting boxes within four wildfires in southwest Montana, USA, using a space‐for‐time substitution chronosequence approach spanning 3–25 years postburn and including an unburned control. We found that bee nesting success and species richness declined with increasing time postburn, with a complete lack of successful bee nesting in unburned areas. Nesting and floral resources were highly variable across both burn severity and time‐since‐burn, yet generally did not have strong effects on nesting success. Our results together suggest that burned areas may provide important habitat for wood‐cavity‐nesting bees in this system. Given ongoing fire regime shifts as well as other threats facing wild bee communities, this work helps provide essential information necessary for the management and conservation of wood‐cavity‐nesting bees.     

Diet characterisation of solitary bees on farmland: dietary specialisation predicts rarity

Changes in agricultural practice across Europe and North America have been associated with declines in wild bee populations. Bee diet breadth has been associated with sensitivity to agricultural intensification, but much of this analysis has been conducted at the categorical level of generalist or specialist, and it is not clear to what extent the level of generalisation within generalist species is also associated with species persistence. We used pollen load analysis to quantify the pollen diets of wild solitary bees on 19 farms across southern England, UK. A total of 72 species of solitary bees were recorded, but only 31 species were abundant enough to allow for formal diet characterisation. The results broadly conformed to existing literature with the majority of species polylectic and collecting pollen from a wide range of plants. Pollen load analysis consistently identified pollens from more plant species and families from each bee species than direct observation of their foraging behaviour. After rarefaction to standardise pollen load sample sizes, diet breadth was significantly associated with frequency of occurrence, with more generalist bees present on more farms than less generalist bees. Our results show that the majority of bee species present on farmland in reasonable numbers are widely variable in their pollen choices, but that those with the broadest diet were present on the greatest number of farms. Increasing the diversity of plants included in agri-environment schemes may be necessary to provide a wider range of pollen resources in order to support a diverse bee community on farmland.     

Wild bees (Hymenoptera: Apoidea) in an urban botanical garden in Buenos Aires,Argentina

We assessed bee diversity and abundance in a botanical garden in the city of Buenos Aires during two consecutive spring–summer periods. Every 15 days, we collected by hand-netting bee specimens seen foraging on flowers during five-minute censuses at every entomophilous herbaceous plant. A total of 66 bee species (Hymenoptera: Apoidea) were recorded. Richness and abundance of native, generalist, and above-ground nesting (cavities) species were higher than that of exotic, specialist, and below-ground (soil) nesting ones, respectively. Social bees were more abundant, while the richness of solitary species was higher than that of social bees. Cleptoparasitic species were represented by a high number of species, even though only a few individuals were captured. Our results suggest that the studied area is an important bee reservoir within the city.     

Small wild bee abundance declines with distance into strawberry crops regardless of field margin habitat

The preservation of pollinator habitat on croplands in the form of hedgerows, wildflower strips, and natural and semi-natural areas can help maintain and enhance wild bee populations in agricultural landscapes. However, there have been few comparisons of the effectiveness of different types of field-margin pollinator habitat in maintaining bee diversity and pollination of the focal crops. We compared wild bee abundance, species richness and community composition between strawberry crops bordered by hedgerows, and those bordered by larger expanses of natural land (forests). Strawberry is an ideal crop in which to investigate pollinator export from field margins as the rows are covered with straw, which reduces habitat for ground-nesting bees within the crop; thus, most wild pollinators need to enter the crop from the margins. We sampled bees in six strawberry fields with hedgerow margins and six strawberry fields with forested margins of at least 200 m in length, using a paired design. We examined strawberry pollen deposition at regular intervals into the fields, and the magnitude of pollinator export from the field margins towards the centre of the crops. We found that bees as a group were no more species-rich or abundant in crops bordered by forests than in crops bordered by hedgerows, although large-bodied bees were more abundant in the former than the latter. Regardless of field-margin type, we found that small wild bee abundance declined significantly from the edge to the centre of the crop, but honey bee (Apis mellifera L.) and large-bodied bee abundance did not. Strawberry pollen deposition also did not decline with distance into the crop. Although previous work indicates that small wild bees are more effective (yield-increasing) pollinators of strawberry on a per-visit basis, their limited foraging ranges suggest they may only pollinate areas near the crop margins, given typical field sizes in our area.     

Bumble bee abundance and richness improves honey bee pollination behaviour in sweet cherry

Pollination service in agricultural crops increases significantly with pollinator diversity and wild pollinator abundance. Differences in the foraging behaviour of pollinating insects are one of the reasons why pollinator diversity and abundance enhances crop pollination. Here, we focused on the foraging behaviour of honey bees and bumble bees in sweet cherry orchards. In addition, we studied the influence of bee diversity and abundance on the foraging behaviour of honey bees and bumble bees. Honey bees were found to visit fewer flowers than bumble bees. Bumble bees also showed a higher probability of changing trees between rows than honey bees. Both visitation rate and probability of row changes of honey bees increased with bumble bee diversity and with bumble bee abundance. We also found that the probability of row changes of honey bees increased with increasing bumble bee abundance. These effects of bumble bee richness and abundance on the pollination behaviour of honey bees can improve the pollination performance of honey bees in crops that depend on cross pollination. Our results highlight the higher pollination performance of bumble bees and the facilitative effect of wild pollinators to crop pollination.     

Agricultural landscapes with organic crops support higher pollinator diversity

Pollinators are traditionally thought to perceive non-flowering crop fields as hostile landscape matrix. In this study, we show that landscapes composed of higher proportions of organic crop fields support more bee species at greater abundances in fallow strips. An increase in organic cropping in the surrounding landscape from 5% to 20% enhanced bee species richness in fallow strips by 50%, density of solitary bees by 60% and bumble bee density by 150%. Bee species richness and bumble bee density responded strongest to organic cropping in landscape sectors with 500 m radius, solitary bee density in landscape sectors with 250 m radius. The most likely source of these results is that crop and noncrop habitats are strongly connected via bee foraging at the landscape scale. It seems likely that bees depending on nesting sites in fallow strips benefited from the more abundant flower resources provided by broadleaved weeds in organic crop fields. We conclude that the incorporation of organic crop fields into conventionally managed agricultural landscapes can provide food resources needed to sustain greater pollinator species richness in noncrop habitats.     

Wild bee pollination networks in northern New England

Conserving and maintaining a diverse assemblage of wild bees is essential for a healthy and functioning ecosystem, as species are uniquely evolved to deliver specific plant–pollination requirements. Understanding the biology and ecology of bees in poorly studied regions is the first step towards conservation. Detailed surveys in New Hampshire reveal a broad diversity of 118 species of wild bees in different guilds and habitats including 17 bee species representing new state records. Network analyses reveal a complex community structure and relatively poorly connected plant–pollinator associations, thus species may be susceptible to disturbance. Phenological analyses document that at least one representative of five native bee families was present throughout the foraging season and both abundance and diversity were highest in June and July. This study provides important baseline information on bee abundance, diversity, phenology, and host plant associations necessary for future conservation efforts.     

Bee Preference for Native versus Exotic Plants in Restored Agricultural Hedgerows

Habitat restoration to promote wild pollinator populations is becoming increasingly common in agricultural lands. Yet, little is known about how wild bees, globally the most important wild pollinators, use resources in restored habitats. We compared bee use of native and exotic plants in two types of restored native plant hedgerows: mature hedgerows (>10 years from establishment) designed for natural enemy enhancement and new hedgerows (≤2 years from establishment) designed to enhance bee populations. Bees were collected from flowers using timed aerial netting and flowering plant cover was estimated by species using cover classes. At mature hedgerow sites, wild bee abundance, richness, and diversity were greater on native plants than exotic plants. At new sites, where native plants were small and had limited floral display, abundance of bees was greater on native plants than exotic plants; but, controlling for floral cover, there was no difference in bee diversity and richness between the two plant types. At both mature and new hedgerows, wild bees preferred to forage from native plants than exotic plants. Honey bees, which were from managed colonies, also preferred native plants at mature hedgerow sites but exhibited no preference at new sites. Our study shows that wild bees, and managed bees in some cases, prefer to forage on native plants in hedgerows over co‐occurring weedy, exotic plants. Semi‐quantitative ranking identified which native plants were most preferred. Hedgerow restoration with native plants may help enhance wild bee abundance and diversity, and maintain honey bee health, in agricultural areas.     

Experience-dependent plasticity in the mushroom bodies of the solitary bee Osmia lignaria (Megachilidae)

All members of the solitary bee species Osmia lignaria (the orchard bee) forage upon emergence from their natal nest cell. Conversely, in the honey bee, days-to-weeks of socially regulated behavioral development precede the onset of foraging. The social honey bee's behavioral transition to foraging is accompanied by neuroanatomical changes in the mushroom bodies, a region of the insect brain implicated in learning. If these changes were general adaptations to foraging, they should also occur in the solitary orchard bee. Using unbiased stereological methods, we estimated the volume of the major compartments of the mushroom bodies, the neuropil and Kenyon cell body region, in adult orchard bees. We compared the mushroom bodies of recently emerged bees with mature bees that had extensive foraging experience. To separate effects of general maturation from field foraging, some orchard bees were confined to a cage indoors. The mushroom body neuropil of experienced field foragers was significantly greater than that of both recently emerged and mature caged orchard bees, suggesting that, like the honey bee, this increase is driven by outdoor foraging experience. Unlike the honey bee, where increases in the ratio of neuropil to Kenyon cell region occur in the worker after emerging from the hive cell, the orchard bee emerged from the natal nest cell with a ratio that did not change with maturation and was comparable to honey-bee foragers. These results suggest that a common developmental endpoint may be reached via different development paths in social and solitary species of foraging bees.     

Fruit set of highland coffee increases with the diversity of pollinating bees

The worldwide decline of pollinators may negatively affect the fruit set of wild and cultivated plants. Here, we show that fruit set of the self-fertilizing highland coffee (Coffea arabica) is highly variable and related to bee pollination. In a comparison of 24 agroforestry systems in Indonesia, the fruit set of coffee could be predicted by the number of flower-visiting bee species, and it ranged from ca. 60% (three species) to 90% (20 species). Diversity, not abundance, explained variation in fruit set, so the collective role of a species-rich bee community was important for pollination success. Additional experiments showed that single flower visits from rare solitary species led to higher fruit set than with abundant social species. Pollinator diversity was affected by two habitat parameters indicating guild-specific nesting requirements: the diversity of social bees decreased with forest distance, whereas the diversity of solitary bees increased with light intensity of the agroforestry systems. These results give empirical evidence for a positive relationship between ecosystem functions such as pollination and biodiversity. Conservation of rainforest adjacent to adequately managed agroforestry systems could improve the yields of farmers.     

Restoration of riverine inland sand dune complexes: implications for the conservation of wild bees

1.  The evaluation of restoration measures is an important task of conservation biology. Inland sand dunes and dry, oligotrophic grasslands have become rare habitat types in large parts of Central Europe and their restoration and management is of major importance for the preservation of many endangered plant and insect species. Within such habitats, it is important to restore key ecosystem services, such as pollination networks. As wild bees are the most important pollinators in many ecosystems, they represent a suitable key group to evaluate restoration measures. Furthermore, the recent decline of many bee species and the potential ecological and economic consequences are currently topics of strong scientific interest.
2.  We studied the succession of bee communities in response to restoration measures of sand dunes and sand grasslands and compared these communities with those of old sand dune complexes.
3.  Our results show that wild bees respond rapidly to restoration measures indicated by a high species richness and abundance. The community structure of bees at restoration sites converged only slightly to those of the target sites. A higher similarity was found between bee communities at the restoration sites (sand dunes and grasslands), indicating that their close proximity was an important determinant of species overlap. Environmental factors such as the number of entomophilous plant species and moisture had a strong influence on wild bee species composition.
4.   Synthesis and applications . The restoration of inland sand dune complexes provides opportunities for colonization by a diverse wild bee community. Although it is difficult to establish a given target community, restoration measures gave rise to a high pollinator diversity and abundance, suggesting that community function can be re-established.     

The bee fauna of large parks in downtown Paris,France

In recent years, a growing number of studies have focused on the potential interest of urban green areas for supporting biodiversity. Private gardens, urban parks or green roofs may support relatively high densities of diverse wild bees. Knowledge is lacking regarding bee assemblages in Paris, the French capital, and one of the most densely populated part of France. In this context, we here provide a first assessment of the taxonomic and functional composition of bee assemblages in three urban parks in downtown Paris. Bees were sampled with a set of three coloured pan traps per park. Fifteen 24-hour sessions were performed from April to July 2011. We captured 425 specimens from 44 species representing five families and 11 genera. The assemblage was dominated by Halictidae (15 species representing 70.1% of total abundance), especially the eusocial species Lasioglossum morio that made 25.2% of total abundance. From a functional point of view, there were only two species of parasitic bees that made 1.2% of total abundance. Most non-parasitic species were polylectic and below-ground nesting. This study highlights the importance of preserving and managing urban parks and other green areas to promote the conservation of wild bees and ultimately the functionality of pollination service in urban ecosystems.     

Hedgerow presence does not enhance indicators of nest‐site habitat quality or nesting rates of ground‐nesting bees

A major challenge in habitat restoration is targeting the key aspects of a species' niche for enhancement, particularly for species that use a diverse set of habitat features. However, restoration that focuses on limited aspects of a species' niche may neglect other resources that are critical to population persistence. We evaluated the ability of native plant hedgerows, planted to increase pollen and nectar resources for wild bees in agricultural landscapes, to provide suitable nesting habitat and enhance nesting rates of ground‐nesting bees. We found that, when compared to unmanaged field edges (controls), hedgerows did not augment most indicators of nest habitat quality (bare ground, soil surface irregularity, and soil hardness), although coarser soils were associated with higher incidence and richness of nesting bees. Hedgerows did not augment nesting rates when compared to control edges. Although all the bee species we detected nesting were also found foraging on floral resources, the foraging versus nesting assemblages found within a site were highly dissimilar. These results may reflect sampling error; or, species found foraging but not nesting in hedgerows could be utilizing hedgerows as “partial habitats,” nesting outside hedgerow plantings but foraging on the floral resources they provide. We conclude that although hedgerows are known to provide critical floral resources to wild bees especially in resource‐poor intensive agricultural landscapes, simply increasing vegetative diversity and structure may not be simultaneously enhancing nesting habitat for ground‐nesting bees.     

Response of wild bees (Hymenoptera: Apoidea: Anthophila) to surrounding land cover in Wisconsin pickling cucumber

Cucumber (Cucumis sativus L.) is among the plants highly dependent on insect-mediated pollination, but little is known about its unmanaged pollinators. Both domestic and wild bee populations in central Wisconsin pickling cucumber fields were assessed using a combination of pan trapping and floral observations before and during bloom. Together with land cover analyses extending 2,000 m from field centers, the relationship of land cover components and bee abundance and diversity were examined. Over a 2-yr sample interval distributed among 18 experimental sites, 3,185 wild bees were collected representing >60 species. A positive association was found between both noncrop and herbaceous areas with bee abundance and diversity only during bloom. Response of bee abundance and diversity to land cover was strongest at larger buffers presumably because of the heterogeneous nature of the landscape and connectivity between crop and noncrop areas. These results are consistent with previous research that has found a weak response of wild bees to surrounding vegetation in moderately fragmented areas. A diverse community of wild bees is present within the fields of a commercial cucumber system, and there is evidence of floral visitation by unmanaged bees. This evidence emphasizes the importance of wild pollinators in fragmented landscapes and the need for additional research to investigate the effectiveness of individual species in pollen deposition.     

环境变化对中国野生蜜蜂多样性的影响

蜜蜂尤其是野生蜜蜂对维持生态系统功能、保证粮食安全等方面具有重要的作用。近年来, 野生蜜蜂的栖息地由于天然林减少, 而现营造的又多为纯林, 以及大面积种植单一经济林而遭到了严重破坏和片断化。已有研究表明纯林、油茶(Camellia oleifera)和橡胶树(Hevea brasiliensis)经济林中的野生蜜蜂多样性较低。现代农业中新烟碱类杀虫剂、除草剂的大规模使用, 会对蜜蜂个体发育和行为产生不利影响。城市化进程潜在影响了蜜蜂的群落, 如郊区的蜜蜂平均物种丰富度要明显高于中心商业区; 废水、废气和粉尘对蜜蜂的觅食、生长发育等都具有不利影响; CO₂等温室气体导致的气候变暖影响了传粉蜜蜂与植物之间的互利共生关系, 造成时间或功能上的不匹配。综上所述, 我国的环境变化可能已导致中国野生传粉蜜蜂多样性的下降和种群的衰退。我国虽是传粉蜜蜂种质资源大国, 但缺乏种类和分布本底以及长期而有效的监测数据, 缺乏对蜜蜂多样性和种群下降机制的研究。因此亟待开展传粉蜜蜂调查、实施长期监测项目, 为之建立研究网络。并通过积累丰富的相关数据, 开展风险预测和评估, 用于管理和缓解传粉蜜蜂下降所带来的经济及非经济影响。     

Canopy thinning,not agricultural history,determines early responses of wild bees to longleaf pine savanna restoration

Longleaf pine savannas are highly threatened, fire‐maintained ecosystems unique to the southeastern United States. Fire suppression and conversion to agriculture have strongly affected this ecosystem, altering overstory canopies, understory plant communities, and animal populations. Tree thinning to reinstate open canopies can benefit understory plant diversity, but effects on animal communities are less well understood. Moreover, agricultural land‐use legacies can have long‐lasting impacts on plant communities, but their effects on animal communities either alone or through interactions with restoration are unclear. Resolving these impacts is important due to the conservation potential of fire‐suppressed and post‐agricultural longleaf savannas. We evaluated how historical agricultural land use and canopy thinning affect the diversity and abundance of wild bees in longleaf pine savannas. We employed a replicated, large‐scale factorial block experiment in South Carolina, where canopy thinning was applied to longleaf pine savannas that were either post‐agricultural or remnant (no agricultural history). Bees were sampled using elevated bee bowls. In the second growing season after restoration, thinned plots supported a greater bee abundance and bee community richness. Additionally, restored plots had altered wild bee community composition when compared to unthinned plots, indicating that reduction of canopy cover by the thinning treatment best predicted wild bee diversity and composition. Conversely, we found little evidence for differences between sites with or without historical agricultural land use. Some abundant Lasioglossum species were the most sensitive to habitat changes. Our results highlight how restoration practices that reduce canopy cover in fire‐suppressed savannas can have rapid benefits for wild bee communities.     

Visitation by wild and managed bees (Hymenoptera: Apoidea) to eastern U.S. native plants for use in conservation programs

Addition of floral resources to agricultural field margins has been shown to increase abundance of beneficial insects in crop fields, but most plants recommended for this use are non-native annuals. Native perennial plants with different bloom periods can provide floral resources for bees throughout the growing season for use in pollinator conservation projects. To identify the most suitable plants for this use, we examined the relative attractiveness to wild and managed bees of 43 eastern U.S. native perennial plants, grown in a common garden setting. Floral characteristics were evaluated for their ability to predict bee abundance and taxa richness. Of the wild bees collected, the most common species (62%) was Bombus impatiens Cresson. Five other wild bee species were present between 3 and 6% of the total: Lasioglossum admirandum (Sandhouse), Hylaeus affinis (Smith), Agapostemon virescens (F.), Halictus ligatus Say, and Ceratina calcarata/dupla Robertson/Say. The remaining wild bee species were present at <2% of the total. Abundance of honey bees (Apis mellifera L.) was nearly identical to that of B. impatiens. All plant species were visited at least once by wild bees; 9 were highly attractive, and 20 were moderately attractive. Honey bees visited 24 of the 43 plant species at least once. Floral area was the only measured factor accounting for variation in abundance and richness of wild bees but did not explain variation in honey bee abundance. Results of this study can be used to guide selection of flowering plants to provide season-long forage for conservation of wild bees.