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Plant-animal interactions are pivotal for ecosystem functioning, and usually form complex networks involving multiple species of mutualists as well as antagonists. The costs and benefits of these interactions show a strong context-dependency directly related to individual variation in partner identity and differential strength. Yet understanding the context-dependency and functional consequences of mutualistic and antagonistic interactions on individuals remains a lasting challenge. We use a network approach to characterize the individual, plant-based pollination interaction networks of the Canarian Isoplexis canariensis (Plantaginaceae) with a mixed assemblage of vertebrate mutualists (birds and lizards) and invertebrate antagonists (florivores, nectar larcenists, and predispersal seed predators). We identify and quantify interaction typologies based on the sign (mutualistic vs. antagonistic) and strength (weak vs. strong) of animal-mediated pollination and test the relationship with individual female reproductive success (FRS). In addition, we document pollinator movement patterns among individual plants to infer events of pollen transfer/receipt that define the plant mating networks and test the relationship with FRS. We identify six interaction typologies along a mutualism-antagonism gradient, with two typologies being over-represented involving both mutualists and antagonists and influencing FRS. Plants showing strong mutualistic interactions, but also (weak or strong) interactions with antagonists are relatively better connected in the mating network (i.e., with higher potential to transfer or receive pollen). Thus, mixed flower visitor assemblages with mutualists and antagonists give plants increased their importance in the mating networks, promote outcrossing and increasing both female and male fitness. Our approach helps characterize plant-animal interaction typologies, the context-specificity of diversified mutualisms, and a better forecasting of their functional consequences.

How native mass‐flowering plants affect the specialization of insects at individual and species levels and the consequences for pollination networks have received much less attention than for mass‐flowering crops or alien species and basically remain unexplored. Using existing DNA metabarcoding data on the pollen loads of 402 flower‐visiting insects, we assessed the effects of a native mass‐flowering plant of high reward quality, the shrub Rhododendron ferrugineum, on pollination networks by investigating: (a) the food niches of individual pollinators and pollinator species and (b) the structure of individual and species networks in subalpine heathland patches with extremely contrasted densities of R. ferrugineum. Relative to its high abundance in high‐density patches, the shrub was greatly underrepresented and did not dominate individual's or species' generalized networks, rather individual and species specialization increased with a decrease in R. ferrugineum density. Furthermore, individuals of the more generalist dipteran Empididae species tended to extend exclusive interactions with rare plant species in low‐density networks. The same trend was observed in the more specialist Apidea but toward rare species in high‐density networks. Our results reveal a quite paradoxical view of pollination and a functional complementarity within networks. Niche and network indices mostly based on the occurrence of links showed that individual pollinators and pollinator species and networks were highly generalized, whereas indices of link strength revealed that species and above all individuals behave as quite strict specialists. Synthesis. Our study provides insights into the status of a native mass‐flowering plant in individual's and insect species' food niches and pollination networks. It revealed that a generalist pollinator species can be highly specialized at the individual level and how rare plant species coexisting with mass‐flowering plants may nevertheless be visited. Using metabarcoding and thus the strength of plant–pollinator links, we provide new strong insights into the status of a native mass‐flowering species (MFS; compared to crops or aliens species) in individual and species pollination networks and the general specialization of generalist individuals increasing with the decrease in the MFS abundance.

Network analysis is a powerful tool to understand community-level plant–pollinator interactions. We evaluated the role of floral visitors on plant fitness through a series of pollination exclusion experiments to test the effectiveness of pollinators of an Ipomoea community in the Pacific coast of Mexico, including: (1) all flower visitors, (2) visitors that contact the reproductive organs, (3) visitors that deposit pollen on stigmas, and (4) visitors that mediate fruit and seed production. Our results show that networks built from effective pollination interactions are smaller, less connected, more specialized and modular than floral visitor networks. Modules are associated with pollinator functional groups and they provide strong support for pollination syndromes only when non-effective interactions are excluded. In contrast to other studies, the analyzed networks are not nested. Our results also show that only 59% of floral visitors were legitimate pollinators that contribute to seed production. Furthermore, only 27% of the links in visitation network resulted in seed production. Our study shows that plant-pollination networks that consider effectiveness measures of pollination in addition to floral visitation provide insightful information about the different role floral visitors play in a community, encompassing a large number of commensalistic/antagonistic interactions and the more restricted set of mutualistic relationships that underlie the evolution of convergent floral phenotypes in plants.

The disruption of mutualisms by invasive species has consequences for biodiversity loss and ecosystem function. Although invasive plant effects on the pollination of individual native species has been the subject of much study, their impacts on entire plant–pollinator communities are less understood. Community-level studies on plant invasion have mainly focused on two fronts: understanding the mechanisms that mediate their integration; and their effects on plant–pollinator network structure. Here we briefly review current knowledge and propose a more unified framework for evaluating invasive species integration and their effects on plant–pollinator communities. We further outline gaps in our understanding and propose ways to advance knowledge in this field. Specifically, modeling approaches have so far yielded important predictions regarding the outcome and drivers of invasive species effects on plant communities. However, experimental studies that test these predictions in the field are lacking. We further emphasize the need to understand the link between invasive plant effects on pollination network structure and their consequences for native plant population dynamics (population growth). Integrating demographic studies with those on pollination networks is thus key in order to achieve a more predictive understanding of pollinator-mediated effects of invasive species on the persistence of native plant biodiversity.

In natural communities, species and their interactions are often organized as nonrandom networks, showing distinct and repeated complex patterns. A prevalent, but poorly explored pattern is ecological modularity, with weakly interlinked subsets of species (modules), which, however, internally consist of strongly connected species. The importance of modularity has been discussed for a long time, but no consensus on its prevalence in ecological networks has yet been reached. Progress is hampered by inadequate methods and a lack of large datasets. We analyzed 51 pollination networks including almost 10,000 species and 20,000 links and tested for modularity by using a recently developed simulated annealing algorithm. All networks with >150 plant and pollinator species were modular, whereas networks with <50 species were never modular. Both module number and size increased with species number. Each module includes one or a few species groups with convergent trait sets that may be considered as coevolutionary units. Species played different roles with respect to modularity. However, only 15% of all species were structurally important to their network. They were either hubs (i.e., highly linked species within their own module), connectors linking different modules, or both. If these key species go extinct, modules and networks may break apart and initiate cascades of extinction. Thus, species serving as hubs and connectors should receive high conservation priorities.

1. Pollination by insects is a key ecosystem service and important to wider ecosystem function. Most species-level pollination networks studied have a generalised structure, with plants having several potential pollinators, and pollinators in turn visiting a number of different plant species. This is in apparent contrast to a plant's need for efficient conspecific pollen transfer. 2. The aim of this study was to investigate the structure of pollen transport networks at three levels of biological hierarchy: community, species and individual. We did this using hoverflies in the genus Eristalis, a key group of non-Hymenopteran pollinators. 3. We constructed pollen transport networks using DNA metabarcoding to identify pollen. We captured hoverflies in conservation grasslands in west Wales, UK, removed external pollen loads, sequenced the pollen DNA on the Illumina MiSeq platform using the standard plant barcode rbcL, and matched sequences using a pre-existing plant DNA barcode reference library. 4. We found that Eristalis hoverflies transport pollen from 65 plant taxa, more than previously appreciated. Networks were generalised at the site and species level, suggesting some degree of functional redundancy, and were more generalised in late summer compared to early summer. In contrast, pollen transport at the individual level showed some degree of specialisation. Hoverflies defined as \"single-plant visitors\" varied from 40% of those captured in early summer to 24% in late summer. Individual hoverflies became more generalised in late summer, possibly in response to an increase in floral resources. Rubus fruticosus agg. and Succisa pratensis were key plant species for hoverflies at our sites 5. Our results contribute to resolving the apparent paradox of how generalised pollinator networks can provide efficient pollination to plant species. Generalised hoverfly pollen transport networks may result from a varied range of short-term specialised feeding bouts by individual insects. The generalisation and functional redundancy of Eristalis pollen transport networks may increase the stability of the pollination service they deliver.

Farm habitat enrichment is crucial for sustainable production of pollinator-dependent crops. Correct choice of crop and non-crop plant species in habitat management support resilient pollinator fauna and effective pollination service delivery. We identify key network metrices to recognize suitable crop and non-crop plant species for farm habitat enrichment. We also highlight the importance of seasonal variation of the key plant and pollinator species that will crucially inform farm management. Crop species played a key role in maintaining plant–pollinator network integrity. In contrast to the conventional practice of focussing on non-crop plants for pollination service restoration, we find crop plants across seasons hold a key role in maintaining healthy plant–pollinator networks. Our study highlights the importance of non-bee pollinators especially, flies and butterflies in sustaining healthy plant–pollinator network. Bees were important as connector species and controlled other species in the network. Only 16.67% bees and 33.33% of plant species acted as connector species. Our study also shows that the identity of connector species in a plant–pollinator network can change drastically across seasons.

Ecological networks depict the interactions between species, mainly based on observations in the field. The information contained in such interaction matrices depends on the sampling design, and typically, compounds preferences (specialization) and abundances (activity). Null models are the primary vehicles to disentangle the effects of specialization from those of sampling and abundance, but they ignore the feedback of network structure on abundances. Hence, network structure, as exemplified here by modularity, is difficult to link to specific causes. Indeed, various processes lead to modularity and to specific interaction patterns more generally. Inferring (co)evolutionary dynamics is even more challenging, as competition and trait matching yield identical patterns of interactions. A satisfactory resolution of the underlying factors determining network structure will require substantial additional information, not only on independently assessed abundances, but also on traits, and ideally on fitness consequences as measured in experimental setups.

Although species and their interactions in unison represent biodiversity and all the ecological and evolutionary processes associated with life, biotic interactions have, contrary to species, rarely been integrated into the concepts of spatial β-diversity. Here, we examine β-diversity of ecological networks by using pollination networks sampled across the Canary Islands. We show that adjacent and distant communities are more and less similar, respectively, in their composition of plants, pollinators and interactions than expected from random distributions. We further show that replacement of species is the major driver of interaction turnover and that this contribution increases with distance. Finally, we quantify that species-specific partner compositions (here called partner fidelity) deviate from random partner use, but vary as a result of ecological and geographical variables. In particular, breakdown of partner fidelity was facilitated by increasing geographical distance, changing abundances and changing linkage levels, but was not related to the geographical distribution of the species. This highlights the importance of space when comparing communities of interacting species and may stimulate a rethinking of the spatial interpretation of interaction networks. Moreover, geographical interaction dynamics and its causes are important in our efforts to anticipate effects of large-scale changes, such as anthropogenic disturbances.

Worldwide, hoverflies (Syrphidae: Diptera) provide crucial ecosystem services such as pollination and biological pest control. Although many hoverfly species exhibit migratory behavior, the spatiotemporal facets of these movement dynamics, and their ecosystem services implications are poorly understood. In this study, we use long-term (16-year) trapping records, trajectory analysis, and intrinsic (i.e., isotope, genetic, pollen) markers to describe migration patterns of the hoverfly Episyrphus balteatus in northern China. Our work reveals how E. balteatus migrate northward during spring–summer and exhibits return (long-range) migration during autumn. The extensive genetic mixing and high genetic diversity of E. balteatus populations underscore its adaptive capacity to environmental disturbances, for example, climate change. Pollen markers and molecular gut analysis further illuminate how E. balteatus visits min. 1012 flowering plant species (39 orders) over space and time. By thus delineating E. balteatus transregional movements and pollination networks, we advance our understanding of its migration ecology and facilitate the design of targeted strategies to conserve and enhance its ecosystem services.