Bumblebees experience landscapes at different spatial scales: possible implications for coexistence

Coexistence in bumblebee communities has largely been investigated at local spatial scales. However, local resource partitioning does not fully explain the species diversity of bumblebee communities. Theoretical studies provide new evidence that partitioning of space can promote species coexistence, when species interact with their environment at different spatial scales. If bumblebee species possess specific foraging ranges, different spatial resource utilisation patterns might operate as an additional mechanism of coexistence in bumblebee communities. We investigated the effects of the landscape-wide availability of different resources (mass flowering crops and semi-natural habitats) on the local densities of four bumblebee species at 12 spatial scales (landscape sectors with 250–3,000 m radius) to indirectly identify the spatial scales at which the bumblebees perceive their environment. The densities of all bumblebee species were enhanced in landscapes with high proportions of mass flowering crops (mainly oilseed rape). We found the strongest effects for Bombus terrestris agg. and Bombus lapidarius at large spatial scales, implying foraging distances of 3,000 and 2,750 m, respectively. The densities of Bombus pascuorum were most strongly influenced at a medium spatial scale (1,000 m), and of Bombus pratorum (with marginal significance) at a small spatial scale (250 m). The estimated foraging ranges tended to be related to body and colony sizes, indicating that larger species travel over larger distances than smaller species, presumably enabling them to build up larger colonies through a better exploitation of food resources. We conclude that coexistence in bumblebee communities could potentially be mediated by species-specific differences in the spatial resource utilisation patterns, which should be considered in conservation schemes.     

Increased bumblebee abundance along the margins of a mass flowering crop: evidence for pollinator spill‐over

Although the practice of cultivating mass flowering crops (MFCs) is seen as a means of counteracting the widespread decline of insect pollinators, no study to‐date has evaluated whether the increased pollinator abundance associated with MFCs influences pollinator visits to assemblages of adjacent native, non‐crop plant species (pollinator ‘spill‐over’). In this study we quantified bumblebee abundance along hedgerow transects in MFC (field bean) vs non‐MFC (wheat) margins. Surveys were conducted on east and west‐facing margins twice daily (morning and afternoon) for three or four days during the main MFC flowering period (June) over four years involving paired bean and wheat fields in Devon and Cornwall, southwest England. Although no single bumblebee species showed any consistent change in relative frequency, when taken across all years and bumblebee species combined, we observed twice as many bumblebees visiting flowers adjacent to the MFC. However, when we compared bumblebee activity along hedgerow transects two weeks after bean flowering, there was no difference between crops. We thus conclude that although there is evidence for pollinator spill‐over from the bean MFC to nearby semi‐natural habitat, the effect is comparatively short lived and not specific to any single bumblebee species. We also suggest that while pollinator services to native plant species might be briefly enhanced by MFC cultivation, the possible repercussions of bumblebee spill‐over from MFCs to semi‐natural habitats and the other pollinator groups they support should be evaluated before MFC cultivation is considered as a means of conserving declining bumblebee populations.     

Genetic diversity and mass resources promote colony size and forager densities of a social bee (Bombus pascuorum) in agricultural landscapes

Although habitat fragmentation and agricultural intensification are known as threads to pollinator diversity, little is known about consequences for population size and genetic diversity. Here, we combined detailed field observations, molecular approaches and GIS-based quantification of landscape structure (measured by proportions of seminatural habitats and proportions of mass flowering crops) to get new insights into driving forces of population dynamics of the bumblebee species Bombus pascuorum. Comparing 13 agriculturally dominated landscape sectors, we found the proportion of mass flowering crops to positively influence bumblebee abundance whereas the proportion of seminatural habitats was of minor importance. We used microsatellites to quantify landscape-related colony densities, inbreeding and population substructure. Detected colony densities did not correlate with landscape parameters or with local worker abundance, measured by field observations. These results indicate that increased worker abundances within landscapes are rather due to greater colony sizes than due to an increased number of nests. We found significant population substructure, measured by F(ST) and seven landscape sectors to bear significantly increased inbreeding values (F(IS)). F(IS) was strongly varying between sectors but did not correlate with landscape structure. Moreover, F(IS) had a significantly negative effect on colony size, demonstrating the importance of genetic diversity on population fitness at a landscape scale. We suggest that inbreeding levels might be related to the temporal variation of food resources and population sizes in agricultural landscapes.     

Mass‐flowering crops dilute pollinator abundance in agricultural landscapes across Europe

Mass‐flowering crops (MFCs) are increasingly cultivated and might influence pollinator communities in MFC fields and nearby semi‐natural habitats (SNHs). Across six European regions and 2 years, we assessed how landscape‐scale cover of MFCs affected pollinator densities in 408 MFC fields and adjacent SNHs. In MFC fields, densities of bumblebees, solitary bees, managed honeybees and hoverflies were negatively related to the cover of MFCs in the landscape. In SNHs, densities of bumblebees declined with increasing cover of MFCs but densities of honeybees increased. The densities of all pollinators were generally unrelated to the cover of SNHs in the landscape. Although MFC fields apparently attracted pollinators from SNHs, in landscapes with large areas of MFCs they became diluted. The resulting lower densities might negatively affect yields of pollinator‐dependent crops and the reproductive success of wild plants. An expansion of MFCs needs to be accompanied by pollinator‐supporting practices in agricultural landscapes.     

Landscape context affects the relationship between local and landscape species richness of butterflies in semi‐natural habitats

Local species richness of butterflies can be expected to benefit from both local habitat properties as well as the availability of suitable habitats and source populations in the surrounding landscape. Whether local species richness is dependent on local or landscape factors can be assessed by examining the relationship between local and landscape species richness. Here we studied how local species richness is related to landscape‐level species richness in landscapes differing in agricultural intensity. The relationship was linear for field boundaries in intensively cultivated landscapes and non‐linear in less‐intensively cultivated landscapes. In landscapes containing semi‐natural grasslands (on average 4% of overall land‐use), the relationship was non‐linear for field boundaries, but linear when considering local species richness of the grasslands themselves. These results show that local factors are more important than landscape factors in determining local species richness in landscapes which contained semi‐natural grasslands. Local species richness was limited by landscape factors in intensively cultivated landscapes. This interpretation was supported by the relationship between local species richness and landscape‐scale average mobility and generalist percentage of butterfly assemblages. We conclude that the management of field boundary habitat quality for butterflies is expected to be most effective in landscapes with semi‐natural grasslands, the species composition of which in turn is dependent on the regional occurrence of grasslands. Based on our results, managing non‐crop habitats for the conservation of habitat specialists and species with poor mobility will be most efficient in regions where patches of semi‐natural grasslands occur.     

Hoverflies (Diptera: Syrphidae) benefit from a cultivation of the bioenergy crop Silphium perfoliatum L. (Asteraceae) depending on larval feeding type,landscape composition and crop management

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Habitat linkages in conservation biological control: Lessons from the land–water interface

Terrestrial landscapes, including those with embedded agroecosystems, are a mosaic of cover types varying in size. Creating or maintaining habitats that support natural enemy populations to combat agricultural pests is the primary method of conservation biological control. Non-crop habitats can be managed in an attempt to maximize the exchange of natural enemies with adjacent agroecosystems with the expectation that they will suppress damaging pest outbreaks. Despite this goal, current habitat management relying on natural enemy spillover into crops has been unreliably effective at reducing pest abundance or increasing crop yield. Furthermore, the expansion and intensification of agriculture and changes in global climate patterns threaten the foundations of conservation biological control in future agroecosystems. However, the aquatic–terrestrial interface offers a natural boundary similar to the one between agroecosystems and their neighboring non-crop habitats that can provide useful insights to the challenges facing growers. Research of the exchanges between water and land suggests general biological and physical processes that govern the movement of organisms between disparate habitats. We propose that like aquatic insects moving from water to land, natural enemy dispersal from non-crop donor habitats into recipient crop patches on the landscape is a function of (1) the production of natural enemies in the source habitat which establishes the abundance of organisms that can disperse, (2) how and why mobile natural enemies disperse themselves into neighboring recipient habitats, and (3) the configuration of donor and recipient habitats on the landscape. We suggest that conservation biological control practitioners can focus on these main components of natural enemy production and dispersal to predict the effectiveness of conservation biological control measures and guide their adaptation to future global change.     

Effects of landscape structure and land-use intensity on similarity of plant and animal communities

Aim Species richness in itself is not always sufficient to evaluate land management strategies for nature conservation. The exchange of species between local communities may be affected by landscape structure and land‐use intensity. Thus, species turnover, and its inverse, community similarity, may be useful measures of landscape integrity from a diversity perspective. Location A European transect from France to Estonia. Methods We measured the similarity of plant, bird, wild bee, true bug, carabid beetle, hoverfly and spider communities sampled along gradients in landscape composition (e.g. total availability of semi‐natural habitat), landscape configuration (e.g. fragmentation) and land‐use intensity (e.g. pesticide loads). Results Total availability of semi‐natural habitats had little effect on community similarity, except for bird communities, which were more homogeneous in more natural landscapes. Bee communities, in contrast, were less similar in landscapes with higher percentages of semi‐natural habitats. Increased landscape fragmentation decreased similarity of true bug communities, while plant communities showed a nonlinear, U‐shaped response. More intense land use, specifically increased pesticide burden, led to a homogenization of bee, bug and spider communities within sites. In these cases, habitat fragmentation interacted with pesticide load. Hoverfly and carabid beetle community similarity was differentially affected by higher pesticide levels: for carabid beetles similarity decreased, while for hoverflies we observed a U‐shaped relationship. Main conclusions Our study demonstrates the effects of landscape composition, configuration and land‐use intensity on the similarity of communities. It indicates reduced exchange of species between communities in landscapes dominated by agricultural activities. Taxonomic groups differed in their responses to environmental drivers and using but one group as an indicator for ‘biodiversity’ as such would thus not be advisable.     

Comparison of pollinators and natural enemies: a meta‐analysis of landscape and local effects on abundance and richness in crops

To manage agroecosystems for multiple ecosystem services, we need to know whether the management of one service has positive, negative, or no effects on other services. We do not yet have data on the interactions between pollination and pest‐control services. However, we do have data on the distributions of pollinators and natural enemies in agroecosystems. Therefore, we compared these two groups of ecosystem service providers, to see if the management of farms and agricultural landscapes might have similar effects on the abundance and richness of both. In a meta‐analysis, we compared 46 studies that sampled bees, predatory beetles, parasitic wasps, and spiders in fields, orchards, or vineyards of food crops. These studies used the proximity or proportion of non‐crop or natural habitats in the landscapes surrounding these crops (a measure of landscape complexity), or the proximity or diversity of non‐crop plants in the margins of these crops (a measure of local complexity), to explain the abundance or richness of these beneficial arthropods. Compositional complexity at both landscape and local scales had positive effects on both pollinators and natural enemies, but different effects on different taxa. Effects on bees and spiders were significantly positive, but effects on parasitoids and predatory beetles (mostly Carabidae and Staphylinidae) were inconclusive. Landscape complexity had significantly stronger effects on bees than it did on predatory beetles and significantly stronger effects in non‐woody rather than in woody crops. Effects on richness were significantly stronger than effects on abundance, but possibly only for spiders. This abundance‐richness difference might be caused by differences between generalists and specialists, or between arthropods that depend on non‐crop habitats (ecotone species and dispersers) and those that do not (cultural species). We call this the ‘specialist‐generalist’ or ‘cultural difference’ mechanism. If complexity has stronger effects on richness than abundance, it might have stronger effects on the stability than the magnitude of these arthropod‐mediated ecosystem services. We conclude that some pollinators and natural enemies seem to have compatible responses to complexity, and it might be possible to manage agroecosystems for the benefit of both. However, too few studies have compared the two, and so we cannot yet conclude that there are no negative interactions between pollinators and natural enemies, and no trade‐offs between pollination and pest‐control services. Therefore, we suggest a framework for future research to bridge these gaps in our knowledge.     

Hibernation of predatory arthropods in semi-natural habitats

Non-crop habitats provide important resources for natural enemies. Many natural enemies hibernate in non-crop habitats, from which they may colonise arable fields in the spring. Spring colonisation ensures annual repopulation of the crop with natural enemies, allowing them to keep pace with the development of pest populations. The availability of non-crop habitats can, therefore, be crucial to successful conservation biological control. We quantified the density of overwintering natural enemies near organic Brussels sprout crops in five different non-crop habitats (short grassy field margin, herbaceous field margin, herbaceous field margin under a tree line, ditch and forest). Soil and litter samples of non-crop habitats were taken at two sites. One site was located in an open agricultural landscape, the other in a landscape dominated by mixed forest. Insects belonging to Staphylinidae, Araneae, Carabidae, Coccinellidae and Dermaptera were the most abundant. Mean densities of predatory arthropods were higher in the open agricultural landscape (290 predators m⁻²) than in the forested landscape (137 predators m⁻²). Herbaceous habitat types supported the highest densities of overwintering predators (up to 400 predators m⁻²), whereas densities in the forest were lowest (10 predators m⁻²). These results indicate that herbaceous non-crop habitats are important refugia for predators and may play a vital role in conservation biological control. Handling Editor: Arne Janssen.     

Coccinellid response to landscape composition and configuration

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Organic farming expansion drives natural enemy abundance but not diversity in vineyard‐dominated landscapes

Organic farming is seen as a prototype of ecological intensification potentially able to conciliate crop productivity and biodiversity conservation in agricultural landscapes. However, how natural enemies, an important functional group supporting pest control services, respond to organic farming at different scales and in different landscape contexts remain unclear. Using a hierarchical design within a vineyard‐dominated region located in southwestern France, we examine the independent effects of organic farming and semi‐natural habitats at the local and landscape scales on natural enemies. We show that the proportion of organic farming is a stronger driver of species abundance than the proportion of semi‐natural habitats and is an important facet of landscape heterogeneity shaping natural enemy assemblages. Although our study highlights a strong taxonomic group‐dependency about the effect of organic farming, organic farming benefits to dominant species while rare species occur at the same frequency in the two farming systems. Independently of farming systems, enhancing field age, reducing crop productivity, soil tillage intensity, and pesticide use are key management options to increase natural enemy biodiversity. Our study indicates that policies promoting the expansion of organic farming will benefit more to ecological intensification strategies seeking to enhance ecosystem services than to biodiversity conservation.     

Spatial and temporal variations in floral resource availability affect bumblebee communities in heathlands

Modifications of landscape structure and composition can decrease the availability of floral resources, resulting in the decline of many pollinator species, including bumblebees. These declines may have significant ecological consequences, because bumblebees pollinate a large range of plant species. Our study was carried out in heathlands, open semi-natural habitats that have decreased considerably due to human activities. We analysed how floral resources affect bumblebee communities throughout the colony lifetime at three scales: plot scale, heathland patch scale, and landscape scale. Floral density at the plot scale and spruce plantations at the landscape scale influenced bumblebee communities. The abundance of bumblebees on ericaceous species was higher when the landscape included a substantial proportion of unsuitable foraging habitat (i.e., spruce plantations). Both life history traits and colony life cycle stage influenced bumblebee responses to the availability of floral resources. Bumblebees were more affected by floral resources during the colony development phase than during the nest-foundation or mating phases. Moreover, bumblebees of species that form large colonies needed larger quantities of favourable foraging habitat, compared with small-colony bees, and their proportion decreased in habitats dominated by spruce plantations. In conclusion, the conservation of plant–bumblebee interactions will require management at a larger spatial scale than the restricted protected habitats. Moreover, at the landscape scale, both quantity of favourable foraging patches and their ecological continuity are important to conserve both small- and large- colony species.     

Floral resource pulse decreases bumble bee foraging trip duration in central Wisconsin agroecosystem

1. Resource pulses, narrow periods of high resource availability, can elicit strong behavioural responses across diverse taxa. Mass‐flowering agricultural crops are an example of a resource pulse that insect pollinators exploit. However, the underlying mechanism behind changes in pollinator behaviour associated with mass‐flowering crops is still relatively unexplored. 2. The present study quantified the behavioural response of bumble bees, an important wild pollinator, to commercial cranberry bloom, an important mass‐flowering crop in Wisconsin, U.S.A. Over a 2‐year period, foraging trip duration was measured using radio frequency identification at 14 farms situated across landscape contexts, ranging from high to low natural area (woodland amount). Using transect surveys, floral resource abundance at a landscape scale was estimated. 3. It was found that bumble bees were highly sensitive to temporal changes in landscape‐level resource abundance associated with the onset of cranberry bloom, during which they decreased foraging trip duration by 22% and increased the number of foraging trips during bloom by 24% on average relative to the period before and after bloom. This phenomenon was consistent across colonies, individual bees, and landscape contexts, despite a higher abundance of flowers in low woodland landscapes. Bumble bee colonies growing in low‐ and high‐woodland landscapes exhibited a similar performance. 4. As mass‐flowering crops are probably a factor influencing bumble bee foraging behaviour in agricultural regions, investigations should continue into how variable resource landscapes, particularly those offering resource pulses, affect wild pollinators and the pollination services they provide.     

Landscape structure influences pollinator movements and directly affects plant reproductive success

Pollinators play a key role within most terrestrial communities in maintaining plant populations, as well as pollinating many agricultural crops for seed and fruit production. The mobility of pollinating animals is significant to their importance but we know little about how landscape structure influences pollinator movements. Linear landscape features such as hedgerows and embankments are conspicuous features of agricultural landscape structure and are important artificial habitats in their own right. However, there has been some debate as to the function of these landscape elements as corridors between larger expanses of semi‐natural habitat separated by urban and agricultural habitats. Few studies have specifically studied insect flight responses to linear landscape elements. By observing bumblebee flight behaviour along hedgerows and, by creating a medium‐scale experimental array of flower patches using an artificial linear feature, we examined whether such structures can elicit an oriented flight response along them and therefore facilitate insect movement through the landscape. We found that both hedgerows and artificial linear landscape features can influence the flight directions of bumblebees (Bombus spp. Hymenoptera, Apidae), one of Europe's most important groups of pollinators. A bioassay experiment in which Salvia pratensis (Lamiaceae) was planted into landscape patches with differing numbers of connecting hedgerows showed that this directional response can have a profound effect on plant reproductive success – plants had increased pollinator activity, pollen receipt and subsequent seed set in patches with more connections. The overall hedgerow connectedness of a landscape is therefore important both to bumblebee movement and to those plants which depend on bumblebees for pollination services.     

Field margin vegetation enhances biological control and crop damage suppression from multiple pests in organic tomato fields

Ephemeral cropping systems are characterized by frequent disturbances of ecological processes, which may compromise the conservation of plant and arthropod diversity and the ecosystem services they may provide. Conservation biological control practices include habitat manipulations that provide non‐pest resources and selectively enhance natural enemies' effectiveness. This study, conducted in eight commercial fields of organically grown tomato, compared the effectiveness of sown flower strips with semi‐natural margins in regulating natural enemy abundance, biocontrol, and crop damage. During repeated visits, the abundance of different arthropod groups was recorded. Crop surveys included measurement of aphid abundance, parasitism, and leaf and fruit damage from sap‐sucking and lepidopteran pests. Semi‐natural habitats were associated with higher vegetation diversity, but natural enemies were more strongly associated with sown strips during flowering. Sap‐sucking pests were always recorded in higher abundance in flower strips, but crop damage in the plots adjacent to these strips was lower, suggesting that these strips may act as a trap‐crop. The inclusion of floral supplements enhanced the parasitism rate of aphids in the crop, and reduced the rate of increase of lepidopteran‐caused foliar damage with time. Early in the growing season, semi‐natural strips showed significantly lower levels of crop damage and aphid counts, suggesting that these habitats may be important during early crop colonization by natural enemies. These results indicate that the inclusion of flower strips enhances the conservation of arthropod functional diversity in ephemeral crops, and that diverse mechanisms are important for controlling different pests. However, the efficacy of habitat manipulation is likely to be greater when it is complemented with the conservation of diverse semi‐natural vegetation in the pre‐existing field margin.     

Differentiation of spring carabid beetle assemblages between semi‐natural habitats and adjoining winter wheat

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Mass-flowering red clover crops have positive effects on bumblebee richness and diversity after bloom

Floral resource quantity in agricultural landscapes plays a key role in the persistence of wild pollinators. An equally important, but less investigated factor is how variation in floral resource availability over time, e.g. floral resource pulses, affects pollinator abundances and diversity. Despite the potential importance of late-season resource pulses for bumblebee reproduction, few studies have evaluated the effects of late-season mass-flowering crops on bumblebee abundances and diversity during and after crop bloom. We assessed how bumblebee abundances, diversity and traits associated with species rarity were affected by cultivation of late-season mass-flowering red clover grown for seed production. Bumblebees were surveyed in red clover fields and flower-rich field borders across 20 landscapes with or without a red clover field during and after crop bloom in southern Sweden. Bumblebee worker abundances were higher in clover fields compared to flower-rich borders in the surrounding landscape. There was no relationship between presence of clover fields and the abundance of males of social bumblebees, but more male cuckoo bumblebees were found in flower-rich borders in landscapes with clover following crop bloom. Mass-flowering red clover also had a positive effect on bumblebee species richness and diversity after crop bloom. Overall, clover had positive and lasting effects on less common bumblebees thereby sustaining higher bumblebee species richness after bloom. Cultivation of red clover has the potential, in combination with the management of flower-rich habitats, to benefit less common bumblebee species in temperate agroecosystems.     

Butterfly diversity at the ecotone between agricultural and semi‐natural habitats across a climatic gradient

Aim Understanding the response of species to ecotones and habitat edges is essential to designing conservation management, especially in mosaic agricultural landscapes. This study examines how species diversity and composition change with distance from semi‐natural habitats, over ecotones into agricultural fields, and how within‐site patterns of community transition change across a climatic gradient and differ between crop types. Location A total of 19 sites in Israel where semi‐natural habitats border agricultural fields (wheat fields or olive groves) distributed along a sharp climatic gradient ranging between 100 and 800 mm mean annual rainfall. Methods  We performed butterfly surveys in 2006. We analysed species richness (α‐diversity), diversity, community nestedness and species turnover (β‐diversity) within sites and between sites (γ‐diversity). We also assessed where species of conservation concern occurred. Results In wheat sites, richness and diversity declined abruptly from ecotones to fields and remained homogenously poor throughout the fields, regardless of climate. In olive sites, despite the sharp structural boundary, richness and diversity remained high from the semi‐natural habitat to the grove margins and then declined gradually into groves. Species of conservation concern occurred across all habitats at olive sites, but none were found inside wheat fields or at their ecotones. The contrast in community structure between semi‐natural habitats and fields was affected by both climate and field type. Irrigation in arid regions did not augment species diversity. Main conclusions Our results indicate that consideration of crop type, within a climatic context, should receive high priority in biodiversity conservation in agricultural areas. In ‘hostile’ crops, such as wheat, we suggest favouring a combination of high‐intensity management and wide margins over less intensive management without margins, which may merely aid generalist butterfly species. The scarcity of butterflies in arid irrigated fields suggests a need to carefully assess the effects of irrigation and agrochemicals on species’ communities.     

Foraging habitats and foraging distances of bumblebees, Bombus spp. (Hym., Apidae), in an agricultural landscape

Abstract: In selected foraging habitats of an agricultural landscape flower visits of bumblebees and community structure of foraging bumblebees were studied, with special regard to the role of crops as super-abundant resources. Most crops represent temporal foraging habitats with high abundance of bumblebees but mainly with low diversity in the bumblebee forage community, in contrast to permanent foraging habitats such as, for example, a hedgerow. The high numbers of bumblebees in the monoculture of crop plantations consisted mainly of short-tongued bumblebee species. The role of foraging distances for the visitation rate of foraging habitats was studied by performing capture–recapture experiments with natural nests of Bombus terrestris , Bombus lapidarius and Bombus muscorum . Differences were found on the species as well as the individual level. The foraging distances of B. muscorum were more restricted to the neighbourhood of the nesting habitat than the foraging activity of B. terrestris and B. lapidarius . High percentages of B. terrestris workers were recaptured while foraging on super-abundant resources in distances up to 1750 m from the nest. Isolated patches of highly rewarding forage crops, in agricultural landscapes, are probably only accessed by bumblebee species with large mean foraging distances, such as the short-tongued B. terrestris . Species like the rare, long-tongued B. muscorum depend on a close connection between nesting and foraging habitat. A restricted foraging radius might be one important factor of bumblebee species loss and potential pollinator limitation in modern agricultural landscapes. Furthermore, long-distance flights of bumblebee pollinators have to be considered in the present discussion on gene flow from transgenic plant species on a landscape scale.