Invertebrate biodiversity continues to decline in cropland.

Modern agriculture has drastically changed global landscapes and introduced pressures on wildlife populations. Policy and management of agricultural systems has changed over the last 30 years, a period characterized not only by intensive agricultural practices but also by an increasing push towards sustainability. It is crucial that we understand the long-term consequences of agriculture on beneficial invertebrates and assess if policy and management approaches recently introduced are supporting their recovery. In this study, we use large citizen science datasets to derive trends in invertebrate occupancy in Great Britain between 1990 and 2019. We compare these trends between regions of no- (0%), low- (greater than 0-50%) and high-cropland (greater than 50%) cover, which includes arable and horticultural crops. Although we detect general declines, invertebrate groups are declining most strongly in high-cropland cover regions. This suggests that even in the light of improved policy and management over the last 30 years, the way we are managing cropland is failing to conserve and restore invertebrate communities. New policy-based drivers and incentives are required to support the resilience and sustainability of agricultural ecosystems. Post-Brexit changes in UK agricultural policy and reforms under the Environment Act offer opportunities to improve agricultural landscapes for the benefit of biodiversity and society.

The lady's 'slippery' orchid: functions of the floral trap and aphid mimicry in a hoverfly-pollinated Phragmipedium species in Brazil.

BACKGROUND AND AIMS: Trap flowers are fascinating cases of adaptation, often linked to oviposition-site mimicry systems. Some trap flowers do not imprison pollinators for a pre-determined period, but rather force them to move through a specific path, manipulating their movements in a way that culminates in pollen transfer, often as they leave through a secondary opening. METHODS: We investigated the previously unknown pollination system of the lady's slipper orchid Phragmipedium vittatum and assessed the function of micro-morphological traits of its trap flowers. KEY RESULTS: Our observations revealed that P. vittatum is pollinated by females of two hoverfly species (Syrphidae). Eggs laid by flies on or near raised black spots on the flowers indicate that the orchid mimics aphids which serve as food for their aphidophagous larvae. Dark, elevated aphid-like spots appear to attract the attention of hoverflies to a slipping zone. This region has downward projecting papillate cells and mucilage secretion that promote slipperiness, causing potential pollinators to fall into the labellum. They then follow a specific upward route towards inner aphid-like spots by holding onto upward oriented hairs that aid their grip. As hoverflies are funnelled by the lateral constriction of the labellum, they pass the stigma, depositing pollen they may be carrying. Later, they squeeze under one of the articulated anthers which places pollen smears onto their upper thorax. Then, they depart through one of the narrow lateral holes by holding onto hairs projecting from the petals. CONCLUSIONS: This study confirms the system of aphid mimicry in Phragmipedium and highlights the sophisticated micro-morphological traits used by trap flowers in pollinator attraction, trapping, guidance and release, thus promoting precise pollen transfer.

The enantiostylous floral polymorphism of Barberetta aurea (Haemodoraceae) facilitates wing pollination by syrphid flies.

BACKGROUND AND AIMS: Sexual polymorphisms of flowers have traditionally been interpreted as devices that promote cross-pollination, but they may also represent adaptations for exploiting particular pollination niches in local environments. The cross-pollination function of enantiostyly, characterized by flowers having either left- or right-deflected styles, has been uncertain in some lineages, such as the Haemodoraceae, because the positioning of stamens and styles is not always completely reciprocal among morphs. METHODS: We examined the floral biology of populations of the poorly known species Barberetta aurea (Haemodoraceae) across its native range in South Africa to establish the general features of its enanatiostylous reproductive system and the agents and mechanism of pollen transfer. RESULTS: We confirmed that B. aurea has a system of dimorphic enantiostyly. Style morph ratios varied among populations sampled, but with an overall tendency to being equal. Crossing experiments demonstrated that B. aurea is fully self-compatible, that intra- and inter-morph crosses are equally fertile and that it is wholly dependent on pollinator visits for seed production. Pollination is mainly by syrphid flies that transfer the sticky pollen via their wings, which contact the anthers and stigma precisely as they hover during approach and feeding. The majority of syrphid fly visitors feed on a film of highly concentrated nectar situated at the base of ultraviolet-absorbent 'nectar guides'. Because one of the three stamens is deflected in the same direction as the style, we predicted a high likelihood of intra-morph pollination, and this was corroborated by patterns of transfer of coloured dye particles in cage experiments involving syrphid flies. CONCLUSIONS: Barbaretta aurea exhibits dimorphic enantiostyly and, in contrast to most enantiostylous species, which are pollinated by bees, its flowers are specialized for pollination by syrphid flies. The lack of complete reciprocity of the enantiostylous arrangement of sexual organs facilitates both inter- and intra-morph pollen transfer on the wings of these flies.

A near-chromosome level genome assembly of the European hoverfly, Sphaerophoria rueppellii (Diptera: Syrphidae), provides comparative insights into insecticide resistance-related gene family evolution.

BACKGROUND: Sphaerophoria rueppellii, a European species of hoverfly, is a highly effective beneficial predator of hemipteran crop pests including aphids, thrips and coleopteran/lepidopteran larvae in integrated pest management (IPM) programmes. It is also a key pollinator of a wide variety of important agricultural crops. No genomic information is currently available for S. rueppellii. Without genomic information for such beneficial predator species, we are unable to perform comparative analyses of insecticide target-sites and genes encoding metabolic enzymes potentially responsible for insecticide resistance, between crop pests and their predators. These metabolic mechanisms include several gene families - cytochrome P450 monooxygenases (P450s), ATP binding cassette transporters (ABCs), glutathione-S-transferases (GSTs), UDP-glycosyltransferases (UGTs) and carboxyl/choline esterases (CCEs). METHODS AND FINDINGS: In this study, a high-quality near-chromosome level de novo genome assembly (as well as a mitochondrial genome assembly) for S. rueppellii has been generated using a hybrid approach with PacBio long-read and Illumina short-read data, followed by super scaffolding using Hi-C data. The final assembly achieved a scaffold N50 of 87Mb, a total genome size of 537.6Mb and a level of completeness of 96% using a set of 1,658 core insect genes present as full-length genes. The assembly was annotated with 14,249 protein-coding genes. Comparative analysis revealed gene expansions of CYP6Zx P450s, epsilon-class GSTs, dietary CCEs and multiple UGT families (UGT37/302/308/430/431). Conversely, ABCs, delta-class GSTs and non-CYP6Zx P450s showed limited expansion. Differences were seen in the distributions of resistance-associated gene families across subfamilies between S. rueppellii and some hemipteran crop pests. Additionally, S. rueppellii had larger numbers of detoxification genes than other pollinator species. CONCLUSION AND SIGNIFICANCE: This assembly is the first published genome for a predatory member of the Syrphidae family and will serve as a useful resource for further research into selectivity and potential tolerance of insecticides by beneficial predators. Furthermore, the expansion of some gene families often linked to insecticide resistance and selectivity may be an indicator of the capacity of this predator to detoxify IPM selective insecticides. These findings could be exploited by targeted insecticide screens and functional studies to increase effectiveness of IPM strategies, which aim to increase crop yields by sustainably and effectively controlling pests without impacting beneficial predator populations.

Genome-wide transcriptomic changes reveal the genetic pathways involved in insect migration.

Insects are capable of extraordinary feats of long-distance movement that have profound impacts on the function of terrestrial ecosystems. The ability to undertake these movements arose multiple times through the evolution of a suite of traits that make up the migratory syndrome, however the underlying genetic pathways involved remain poorly understood. Migratory hoverflies (Diptera: Syrphidae) are an emerging model group for studies of migration. They undertake seasonal movements in huge numbers across large parts of the globe and are important pollinators, biological control agents and decomposers. Here, we assembled a high-quality draft genome of the marmalade hoverfly (Episyrphus balteatus). We leveraged this genomic resource to undertake a genome-wide transcriptomic comparison of actively migrating Episyrphus, captured from a high mountain pass as they flew south to overwinter, with the transcriptomes of summer forms which were non-migratory. We identified 1543 genes with very strong evidence for differential expression. Interrogation of this gene set reveals a remarkable range of roles in metabolism, muscle structure and function, hormonal regulation, immunity, stress resistance, flight and feeding behaviour, longevity, reproductive diapause and sensory perception. These features of the migrant phenotype have arisen by the integration and modification of pathways such as insulin signalling for diapause and longevity, JAK/SAT for immunity, and those leading to octopamine production and fuelling to boost flight capabilities. Our results provide a powerful genomic resource for future research, and paint a comprehensive picture of global expression changes in an actively migrating insect, identifying key genomic components involved in this important life-history strategy.

Establishment and management of wildflower areas for insect pollinators in commercial orchards.

Sown wildflower areas are increasingly recommended as an agri-environmental intervention measure, but evidence for their success is limited to particular insect groups or hampered by the challenges of establishing seed mixes and maintaining flower abundance over time. We conducted a replicated experiment to establish wildflower areas to support insect pollinators in apple orchards. Over three years, and across 23 commercial UK orchards with and without sown wildflowers, we conducted 828 transect surveys across various non-crop habitats. We found that the abundance of flower-visiting solitary bees, bumblebees, honeybees, and beetles was increased in sown wildflower areas, compared with existing non-crop habitats in control orchards, from the second year following floral establishment. Abundance of hoverflies and other non-syrphid flies was increased in wildflower areas from the first year. Beyond the effect of wildflower areas, solitary bee abundance was also positively related to levels of floral cover in other local habitats within orchards, but neither local nor wider landscape-scale context affected abundance of other studied insect taxa within study orchards. There was a change in plant community composition on the sown wildflower areas between years, and in patterns of flowering within and between years, showing a succession from unsown weedy species towards a dominance of sown species over time. We discuss how the successful establishment of sown wildflower areas and delivery of benefits for different insect taxa relies on appropriate and reactive management practices as a key component of any such agri-environment scheme.

Pollination by hoverflies in the Anthropocene.

Pollinator declines, changes in land use and climate-induced shifts in phenology have the potential to seriously affect ecosystem function and food security by disrupting pollination services provided by insects. Much of the current research focuses on bees, or groups other insects together as 'non-bee pollinators', obscuring the relative contribution of this diverse group of organisms. Prominent among the 'non-bee pollinators' are the hoverflies, known to visit at least 72% of global food crops, which we estimate to be worth around US$300 billion per year, together with over 70% of animal pollinated wildflowers. In addition, hoverflies provide ecosystem functions not seen in bees, such as crop protection from pests, recycling of organic matter and long-distance pollen transfer. Migratory species, in particular, can be hugely abundant and unlike many insect pollinators, do not yet appear to be in serious decline. In this review, we contrast the roles of hoverflies and bees as pollinators, discuss the need for research and monitoring of different pollinator responses to anthropogenic change and examine emerging research into large populations of migratory hoverflies, the threats they face and how they might be used to improve sustainable agriculture.

Cypripedium subtropicum (Orchidaceae) employs aphid colony mimicry to attract hoverfly (Syrphidae) pollinators.

In Orchidaceae, pollination is mostly animal-mediated, and one-third of species have evolved a deceptive pollination mechanism without rewards. Cypripedium is a representative lineage of nonrewarding orchids restricted to temperate regions. Cypripedium subtropicum flowers are pollinated by hoverflies and have hairy tufts that visually resemble an aphid colony covered with honey dew. We recorded the behavior of hoverflies on the flowers, determined the breeding system of the species and the structure of hairy tufts, and investigated the roles of hairy tufts and floral volatiles in this specialized pollination by using pollination experiments, scanning electron microscopy, bioassays and chemical analyses. The white hairy tufts covering the sidelobes of the labellum provide edible rewards and serve as crucial visual lures for hoverflies. The flowers emit primarily (E)-ß-farnesene and a smaller amount of ß-pinene that were found to attract hoverflies. Our results suggest that C. subtropicum uses both visual mimicry of an aphid-colonized labellum with a reward and chemical mimicry of aphid alarm pheromones to attract hoverflies for pollination. This is the first described example of a rewarding mimicry system in plants, where the models are animals with their secretions and the reward is similar in nutrients to that of the model mimicked.

How do hoverflies use their righting reflex?

When taking off from a sloping surface, flies have to reorient themselves dorsoventrally and stabilize their body by actively controlling their flapping wings. We have observed that righting is achieved solely by performing a rolling manoeuvre. How flies manage to do this has not yet been elucidated. It was observed here for the first time that hoverfly reorientation is entirely achieved within 6 wingbeats (48.8 ms) at angular roll velocities of up to 10×103 deg s-1 and that the onset of their head rotation consistently follows that of their body rotation after a time lag of 16 ms. The insects' body roll was found to be triggered by the asymmetric wing stroke amplitude, as expected. The righting process starts immediately with the first wingbeat and seems unlikely to depend on visual feedback. A dynamic model for the fly's righting reflex is presented, which accounts for the head/body movements and the time lag recorded in these experiments. This model consists of a closed-loop control of the body roll, combined with a feedforward control of the head/body angle. During the righting manoeuvre, a strong coupling seems to exist between the activation of the halteres (which measure the body's angular speed) and the gaze stabilization reflex. These findings again confirm the fundamental role played by the halteres in both body and head stabilization processes.

A world review of reported myiases caused by flower flies (Diptera: Syrphidae), including the first case of human myiasis from Palpada scutellaris (Fabricius, 1805).

Rat-tailed larvae of the syrphid species Palpada scutellaris (Fabricius, 1805) are documented causing an enteric human myiasis in Costa Rica. This is the first time that the genus Palpada is recorded as a human myiasis agent. We report a 68-year-old woman with intestinal pain and bloody diarrhea with several live Palpada larvae present in the stool. Using molecular techniques (DNA barcodes) and both electronic and optical microscopy to study the external morphology, the preimaginal stages of the fly were unambiguously identified. An identification key to all syrphid genera actually known as agents of human and animal myiases is provided for larvae, puparia, and adults. Moreover, a critical world review of more than 100 references of Syrphidae as myiasis agents is also given, with emphasis on the species with rat-tailed larvae.

Network modularity influences plant reproduction in a mosaic tropical agroecosystem.

Biodiversity influences ecosystem function, but there is limited understanding of the mechanisms that support this relationship across different land use types in mosaic agroecosystems. Network approaches can help to understand how community structure influences ecosystem function across landscapes; however, in ecology, network analyses have largely focused on species-species interactions. Here, we use bipartite network analysis in a novel way: to link pollinator communities to sites in a tropical agricultural landscape. We used sentinel plants of Brassica rapa to examine how the structure of the community network influences plant reproduction. Diptera was the most common order of flower visitors at every site. Syrphidae visits were the strongest contributor to the number of fertilized pods, while visits by Syrphidae, Hymenoptera and Lepidoptera had the strongest effect on the number of seeds per pod. Sentinel pots at forest sites were visited by more unique species (i.e. species with higher d') than sites in other land uses, and dairy sites had more visitors that were common across the network. Participation coefficients, which indicate how connected a single node is across network modules, were strong predictors of ecosystem function: plant reproduction increased at sites with higher participation coefficients. Flower visitor taxa with higher participation coefficients also had the strongest effect on plant reproduction. Hymenoptera visits were the best predictor for participation coefficients but an Allograpta sp. (Diptera: Syrphidae) was the most influential flower visitor species in the landscape network. A diverse insect community contributed to plant reproduction and connection among nodes in this system. Identifying the 'keystone' flower visitor species and sites that have a strong influence on network structure is a significant step forward to inform conservation priorities and decision-making in diverse agroecosystems.

The yellow specialist: dronefly Eristalis tenax prefers different yellow colours for landing and proboscis extension.

Droneflies, imagoes of the hoverfly Eristalis tenax, are known to possess a preference for yellow flowers, i.e. they prefer to visit yellow flowers and prefer to extend the proboscis to yellow colours. In this study we disentangle these colour preferences by investigating the landing reaction and proboscis reflex with particular reference to intensity, spectral purity and dominant wavelength of colour stimuli and their UV reflection properties. In multiple-choice tests, naïve and non-trained flies prefer to land on yellow colours independent of their UV reflection characteristics, but also accept blue, white and pink colours if they absorb UV and are of sufficient brightness. Flies trained to land on colours other than yellow still prefer yellow colours to some extent. Moreover, the flies prefer bright over dark yellow colours even if trained to dark yellow ones. The flies refuse to land on dark colours of all hues. Naïve flies exhibit the proboscis reflex only to pure yellow pollen. These experiments show for the first time that landing in droneflies is triggered by a yellow colour independent of its UV reflection properties, but proboscis extension is triggered by yellow colours strongly absorbing blue and UV. The ability to discriminate colours is better than predicted by the categorical colour vision model. The colour preferences in E. tenax represent a fine-tuned ability to visit yellow flowers displaying a UV bull's-eye pattern.

Disturbance-mediated heterogeneity drives pollinator diversity in boreal managed forest ecosystems.

Intensive forest management, together with fire suppression, have decreased structural complexity and altered dynamics of boreal forests profoundly. Such management threatens forest biodiversity and can reduce the provision of ecosystem services. Although the importance of ecosystem services is widely acknowledged, conservation strategies are hindered by poor knowledge about diversity patterns of service provider species as well as on mechanisms affecting these assemblages at different spatial and temporal scales. In this study, we assessed the effect of disturbance management on forest pollinator communities. To do so, we used a large-scale ecological experiment conducted in the year 2000, where forest complexity was manipulated with different harvest regimes and prescribed fire. Results were consistent with a positive response of pollinators to increasing habitat heterogeneity driven by past disturbances. Harvested sites harbored a diverse pollinator community, and showed higher spatial and temporal turnover in species richness. Conversely, old-growth forest communities were a nested subset of harvested sites and contained half of their total diversity. Variation in community composition (ß diversity) was primarily affected by species temporal turnover. Throughout the season, ß diversity was controlled by fire and harvesting legacies, which provide environmental heterogeneity in the form of flowering and nesting resources over space and time. Conservation strategies may undervalue ecosystem services in dynamic, naturally disturbance-driven, landscapes when relying solely on undisturbed forests areas. However, maintaining natural dynamics in early successional forests, by emulating natural disturbances at harvesting, hold promise for the conservation of both biodiversity and ecosystem services in boreal forests.

Why many Batesian mimics are inaccurate: evidence from hoverfly colour patterns.

Mimicry is considered a classic example of the elaborate adaptations that natural selection can produce, yet often similarity between Batesian (harmless) mimics and their unpalatable models is far from perfect. Variation in mimetic accuracy is a puzzle, as natural selection should favour mimics that are hardest to distinguish from their models. Numerous hypotheses exist to explain the persistence of inaccurate mimics, but most have rarely or never been tested against empirical observations from wild populations. One reason for this is the difficulty in measuring pattern similarity, a key aspect of mimicry. Here, we use a recently developed method, based on the distance transform of binary images, to quantify pattern similarity both within and among species for a group of hoverflies and their hymenopteran models. This allowed us to test three key hypotheses regarding inaccurate mimicry. Firstly, we tested the prediction that selection should be more relaxed in less accurate mimics, but found that levels of phenotypic variation are similar across most hoverfly species. Secondly, we found no evidence that mimics have to compromise between accuracy to multiple model species. However, we did find that darker-coloured hoverflies are less accurate mimics, which could lead to a trade-off between mimicry and thermoregulation in temperate regions. Our results shed light on a classic problem concerning the limitations of natural selection.

The forgotten flies: the importance of non-syrphid Diptera as pollinators.

Bees, hoverflies and butterflies are taxa frequently studied as pollinators in agricultural and conservation contexts. Although there are many records of non-syrphid Diptera visiting flowers, they are generally not regarded as important pollinators. We use data from 30 pollen-transport networks and 71 pollinator-visitation networks to compare the importance of various flower-visiting taxa as pollen-vectors. We specifically compare non-syrphid Diptera and Syrphidae to determine whether neglect of the former in the literature is justified. We found no significant difference in pollen-loads between the syrphid and non-syrphid Diptera. Moreover, there was no significant difference in the level of specialization between the two groups in the pollen-transport networks, though the Syrphidae had significantly greater visitation evenness. Flower visitation data from 33 farms showed that non-syrphid Diptera made up the majority of the flower-visiting Diptera in the agricultural studies (on average 82% abundance and 73% species richness), and we estimate that non-syrphid Diptera carry 84% of total pollen carried by farmland Diptera. As important pollinators, such as bees, have suffered serious declines, it would be prudent to improve our understanding of the role of non-syrphid Diptera as pollinators.

Night life on the beach: selfing to avoid pollinator competition between two sympatric Silene species.

BACKGROUND AND AIMS: Evolution of autonomous selfing may be advantageous because it allows for reproductive assurance. In co-flowering plants competing for pollinators, the least common and/or attractive could suffer pollen limitations. Silene niceensis and S. ramosissima are taxonomically related species sharing the same habitat, although S. ramosissima is less abundant and has a more restricted distribution. They also have the same a priori nocturnal pollinator syndrome, and show an overlapping flowering phenology. The aim of this study was to investigate whether a selfing strategy in S. ramosissima allows it to avoid pollinator competition and/or interspecific pollen transfer with S. niceensis, which would thus enable both species to reach high levels of fruit and seed set. METHODS: The breeding system, petal colour, flower life span and degree of overlap between male and female phases, floral visitor abundance and visitation rates were analysed in two sympatric populations of S. niceensis and S. ramosissima in southern Spain. KEY RESULTS: Autonomous selfing in S. ramosissima produced very high fruit and seed set, which was also similar to open-pollinated plants. Silene niceensis showed minimum levels of autonomous selfing, and pollen/ovule ratios were within the range expected for the breeding system. In contrast to S. niceensis, flower life span was much shorter in S. ramosissima, and male and female organs completely overlapped in space and time. Upper surface petals of both species showed differing brightness, chroma and hue. Flowers of S. niceensis were actively visited by moths, hawkmoths and syrphids, whereas those of S. ramosissima were almost never visited. CONCLUSIONS: The findings show that different breeding strategies exist between the sympatric co-flowering S. niceensis and S. ramosissima, the former specializing in crepuscular-nocturnal pollination and the latter mainly based on autonomous selfing. These two strategies allow both species to share the restricted dune habitat in which they exist, with a high female reproductive success due to the absence of pollinator competition and/or interspecific pollen flow.

Four Forensic Entomology Case Studies: Records and Behavioral Observations on Seldom Reported Cadaver Fauna With Notes on Relevant Previous Occurrences and Ecology.

A yearlong survey of insect taxa associated with human decomposition was conducted at the Southeast Texas Applied Forensic Science (STAFS) facility located in the Center for Biological Field Studies of Sam Houston State University in Huntsville, TX. During this study, four insect-cadaver interactions were observed that represent previously poorly documented yet forensically significant interactions: Syrphidae maggots colonized a corpse in an aquatic situation; Psychodidae adults mated and oviposited on an algal film that was present on a corpse that had been recently removed from water; several Panorpidae were the first insects to feed upon a freshly placed corpse in the autumn; and a noctuid caterpillar was found chewing and ingesting dried human skin. Baseline knowledge of insect-cadaver interactions is the foundation of forensic entomology, and unique observations have the potential to expand our understanding of decomposition ecology.

Does metabolic rate and evaporative water loss reflect differences in migratory strategy in sexually dimorphic hoverflies?

A typical explanation for ecologically stable strategies that apply to only a proportion of a population, is bet hedging, where increased reproductive success offsets reduced reproductive rate. One such is partial migration, where only a proportion of a population moves seasonally to avoid inclement climatic conditions. Bet hedging may overlook unseen costs to maintain broad physiological resilience, implied by encountering a breadth of environmental conditions. We investigated the physiological correlates of partial migration by measuring standard metabolic rates, and rates of evaporative water loss, and then estimating upper and lower thermal tolerance in males and females of two hoverfly species, Episyrphus balteatus and Eristalis tenax. In central Europe, females of these species may either migrate or overwinter, whereas males may migrate south to the Mediterranean, but have not been found overwintering. Both species were sexually dimorphic; female Ep. balteatus were lighter than males, but female Er. tenax were heavier than males. While allometrically- corrected metabolic rate in both species increased with temperature, the most parsimonious models included no sex-specific differences in metabolic rate for either species. Evaporative water loss of both species also increased with temperature, but was higher for females of both species than males. Assuming that resting metabolism is congruent with the activity requirements of migration, highly consistent thermal tolerance and metabolic rate suggests that any given fly could migrate, although water loss patterns suggest that females may be less well-adapted to Mediterranean climates. We infer that partial migration probably results from the imperatives of their reproductive strategies.

Diversity of the Insect Visitors on Calluna vulgaris (Ericaceae) in Southern France Heathlands.

As part of an ongoing research project on the pollination networks in European heathlands, the objective of this study was to assess the insect visitor guild on Calluna vulgaris (L.) Hull (Ericaceae). We focused the study on a region renowned for its largely well-preserved heathlands, the Cévennes National Park, Southern France. In 2013, flower visitors were observed over 3 d per site, in four heathland sites at mont Lozère. Honeybees (Apis mellifera L.) were the main visitors (62-88% of total visitors). Besides honeybees, a high diversity of visitors was detected with 57 different species identified (42 Diptera and 15 Hymenoptera). Hoverflies (Syrphidae, Diptera) visitors were abundant and diverse, especially individuals belonging to the genera Eristalis and Episyrphus. The reported diversity of visitors was probably due to the preservation of large heathland areas at mont Lozère and to the generalist pollination system of C. vulgaris.

Disentangling multiple drivers of pollination in a landscape-scale experiment.

Animal pollination is essential for the reproductive success of many wild and crop plants. Loss and isolation of (semi-)natural habitats in agricultural landscapes can cause declines of plants and pollinators and endanger pollination services. We investigated the independent effects of these drivers on pollination of young cherry trees in a landscape-scale experiment. We included (i) isolation of study trees from other cherry trees (up to 350 m), (ii) the amount of cherry trees in the landscape, (iii) the isolation from other woody habitats (up to 200 m) and (iv) the amount of woody habitats providing nesting and floral resources for pollinators. At the local scale, we considered effects of (v) cherry flower density and (vi) heterospecific flower density. Pollinators visited flowers more often in landscapes with high amount of woody habitat and at sites with lower isolation from the next cherry tree. Fruit set was reduced by isolation from the next cherry tree and by a high local density of heterospecific flowers but did not directly depend on pollinator visitation. These results reveal the importance of considering the plant's need for conspecific pollen and its pollen competition with co-flowering species rather than focusing only on pollinators' habitat requirements and flower visitation. It proved to be important to disentangle habitat isolation from habitat loss, local from landscape-scale effects, and direct effects of pollen availability on fruit set from indirect effects via pollinator visitation to understand the delivery of an agriculturally important ecosystem service.