Pollinator individual‐based networks reveal the specialized plant–pollinator mutualism in two biodiverse communities

Generalization of pollination systems is widely accepted by ecologists in the studies of plant–pollinator interaction networks at the community level, but the degree of generalization of pollination networks remains largely unknown at the individual pollinator level. Using potential legitimate pollinators that were constantly visiting flowers in two alpine meadow communities, we analyzed the differences in the pollination network structure between the pollinator individual level and species level. The results showed that compared to the pollinator species‐based networks, the linkage density, interaction diversity, interaction evenness, the average plant linkage level, and interaction diversity increased, but connectance, degree of nestedness, the average of pollinator linkage level, and interaction diversity decreased in the pollinator individual‐based networks, indicating that pollinator individuals had a narrower food niche than their counterpart species. Pollination networks at the pollinator individual level were more specialized at the network level (H′₂) and the plant species node level (d′) than at the pollinator species‐level networks, reducing the chance of underestimating levels of specialization in pollination systems. The results emphasize that research into pollinator individual‐based pollination networks will improve our understanding of the pollination networks at the pollinator species level and the coevolution of flowering plants and pollinators.     

The role of individual variation in flowering and pollination in the reproductive success of a crepuscular buzz-pollinated plant

Background and AimsPlant individuals within a population differ in their phenology and interactions with pollinators. However, it is still unknown how individual differences affect the reproductive success of plants that have functionally specialized pollination systems. Here, we evaluated whether plant individual specialization in phenology (temporal specialization) and in pollination (pollinator specialization) affect the reproductive success of the crepuscular-bee-pollinated plant Trembleya laniflora (Melastomataceae).MethodsWe quantified flowering activity (amplitude, duration and overlap), plant–pollinator interactions (number of flowers visited by pollinators) and reproductive success (fruit set) of T. laniflora individuals from three distinct locations in rupestrian grasslands of southeastern Brazil. We estimated the degree of individual temporal specialization in flowering phenology and of individual specialization in plant–pollinator interactions, and tested their relationship with plant reproductive success.Key Results Trembleya laniflora presented overlapping flowering, a temporal generalization and specialized pollinator interactions. Flowering overlap among individuals and populations was higher than expected by chance but did not affect the individual interactions with pollinators and nor their reproductive success. In contrast, higher individual generalization in the interactions with pollinators was related to higher individual reproductive success.ConclusionsOur findings suggest that individual generalization in plant–pollinator interaction reduces the potential costs of specialization at the species level, ensuring reproductive success. Altogether, our results highlight the complexity of specialization/generalization of plant–pollinator interactions at distinct levels of organization, from individuals to populations, to species.     

Network robustness and structure depend on the phenological characteristics of plants and pollinators

Many structural patterns have been found to be important for the stability and robustness of mutualistic plant–pollinator networks. These structural patterns are impacted by a suite of variables, including species traits, species abundances, their spatial configuration, and their phylogenetic history. Here, we consider a specific trait: phenology, or the timing of life history events. We expect that timing and duration of activity of pollinators, or of flowering in plants, could greatly affect the species'' roles within networks in which they are embedded. Using plant–pollinator networks from 33 sites in southern British Columbia, Canada, we asked (a) how phenological species traits, specifically timing of first appearance in the network and duration of activity in a network, were related to species'' roles within a network, and (b) how those traits affected network robustness to phenologically biased species loss. We found that long duration of activity increased connection within modules for both pollinators and plants and among modules for plants. We also found that date of first appearance was positively related to interaction strength asymmetry in plants but negatively related to pollinators. Networks were generally more robust to the loss of pollinators than plants, and robustness increased if the models allow new interactions to form when old ones are lost, constrained by overlapping phenology of plants and pollinators. Robustness declined with the loss of late‐flowering plants, which tended to have higher interaction strength asymmetry. In addition, robustness declined with loss of early‐flying or long‐duration pollinators. These pollinators tended to be among‐module connectors. Our results point to networks being limited by early‐flying pollinators. If plants flower earlier due to climate change, plant fitness may decline as they will depend on early emerging pollinators, unless pollinators also emerge earlier.     

Urbanization increases floral specialization of pollinators

Understanding how urbanization alters functional interactions among pollinators and plants is critically important given increasing anthropogenic land use and declines in pollinator populations. Pollinators often exhibit short‐term specialization and visit plants of the same species during one foraging trip. This facilitates plant receipt of conspecific pollen—pollen on a pollinator that is the same species as the plant on which the pollinator was foraging. Conspecific pollen receipt facilitates plant reproductive success and is thus important to plant and pollinator persistence. We investigated how urbanization affects short‐term specialization of insect pollinators by examining pollen loads on insects’ bodies and identifying the number and species of pollen grains on insects caught in urban habitat fragments and natural areas. We assessed possible drivers of differences between urban and natural areas, including frequency dependence in foraging, species richness and diversity of the plant and pollinator communities, floral abundance, and the presence of invasive plant species. Pollinators were more specialized in urban fragments than in natural areas, despite no differences in the species richness of plant communities across site types. These differences were likely driven by higher specialization of common pollinators, which were more abundant in urban sites. In addition, pollinators preferred to forage on invasive plants at urban sites and native plants at natural sites. Our findings reveal indirect effects of urbanization on pollinator fidelity to individual plant species and have implications for the maintenance of plant species diversity in small habitat fragments. Higher preference of pollinators for invasive plants at urban sites suggests that native species may receive fewer visits by pollinators. Therefore, native plant species diversity may decline in urban sites without continued augmentation of urban flora or removal of invasive species.     

Functional Diversity of Plant–Pollinator Interaction Webs Enhances the Persistence of Plant Communities

Pollination is exclusively or mainly animal mediated for 70% to 90% of angiosperm species. Thus, pollinators provide an essential ecosystem service to humankind. However, the impact of human-induced biodiversity loss on the functioning of plant–pollinator interactions has not been tested experimentally. To understand how plant communities respond to diversity changes in their pollinating fauna, we manipulated the functional diversity of both plants and pollinators under natural conditions. Increasing the functional diversity of both plants and pollinators led to the recruitment of more diverse plant communities. After two years the plant communities pollinated by the most functionally diverse pollinator assemblage contained about 50% more plant species than did plant communities pollinated by less-diverse pollinator assemblages. Moreover, the positive effect of functional diversity was explained by a complementarity between functional groups of pollinators and plants. Thus, the functional diversity of pollination networks may be critical to ecosystem sustainability.     

Contrasting effects of invasive plants in plant–pollinator networks

The structural organization of mutualism networks, typified by interspecific positive interactions, is important to maintain community diversity. However, there is little information available about the effect of introduced species on the structure of such networks. We compared uninvaded and invaded ecological communities, to examine how two species of invasive plants with large and showy flowers (Carpobrotus affine acinaciformis and Opuntia stricta) affect the structure of Mediterranean plant–pollinator networks. To attribute differences in pollination to the direct presence of the invasive species, areas were surveyed that contained similar native plant species cover, diversity and floral composition, with or without the invaders. Both invasive plant species received significantly more pollinator visits than any native species and invaders interacted strongly with pollinators. Overall, the pollinator community richness was similar in invaded and uninvaded plots, and only a few generalist pollinators visited invasive species exclusively. Invasive plants acted as pollination super generalists. The two species studied were visited by 43% and 31% of the total insect taxa in the community, respectively, suggesting they play a central role in the plant–pollinator networks. Carpobrotus and Opuntia had contrasting effects on pollinator visitation rates to native plants: Carpobrotus facilitated the visit of pollinators to native species, whereas Opuntia competed for pollinators with native species, increasing the nestedness of the plant–pollinator network. These results indicate that the introduction of a new species to a community can have important consequences for the structure of the plant–pollinator network.     

Community structure of pollination webs of Mauritian heathland habitats

Pollination webs have recently deepened our understanding of complex ecosystem functions and the susceptibility of biotic networks to anthropogenic disturbances. Extensive mutualistic networks from tropical species-rich communities, however, are extremely scarce. We present fully quantitative pollination webs of two plant–pollinator communities of natural heathland sites, one of which was in the process of being restored, on the oceanic island of Mauritius. The web interaction data cover a full flowering season from September 2003 to March 2004 and include all flowering plant and their pollinator species. Pollination webs at both sites were dominated by a few super-abundant, disproportionately well-connected species, and many rare and specialised species. The webs differed greatly in size, reflecting higher plant and pollinator species richness and abundance at the restored site. About one fifth of plant species at the smaller community received <3 visits. The main pollinators were insects from diverse taxonomic groups, while the few vertebrate pollinator species were abundant and highly linked. The difference in plant community composition between sites appeared to strongly affect the associated pollinator community and interactions with native plant species. Low visitation rate to introduced plant species suggested little indirect competition for pollinators with native plant species. Overall, our results indicated that the community structure was highly complex in comparison to temperate heathland communities. We discuss the observed differences in plant linkage and pollinator diversity and abundance between the sites with respect to habitat restoration management and its influence on pollination web structure and complexity. For habitat restoration to be successful in the long term, practitioners should aim to maintain structural diversity to support a species-rich and abundant pollinator assemblage which ensures native plant reproduction.     

Functional diversity of plant-pollinator interaction webs enhances the persistence of plant communities

Pollination is exclusively or mainly animal mediated for 70% to 90% of angiosperm species. Thus, pollinators provide an essential ecosystem service to humankind. However, the impact of human-induced biodiversity loss on the functioning of plant–pollinator interactions has not been tested experimentally. To understand how plant communities respond to diversity changes in their pollinating fauna, we manipulated the functional diversity of both plants and pollinators under natural conditions. Increasing the functional diversity of both plants and pollinators led to the recruitment of more diverse plant communities. After two years the plant communities pollinated by the most functionally diverse pollinator assemblage contained about 50% more plant species than did plant communities pollinated by less-diverse pollinator assemblages. Moreover, the positive effect of functional diversity was explained by a complementarity between functional groups of pollinators and plants. Thus, the functional diversity of pollination networks may be critical to ecosystem sustainability.     

Are native and non‐native pollinator friendly plants equally valuable for native wild bee communities?

Bees rely on floral pollen and nectar for food. Therefore, pollinator friendly plantings are often used to enrich habitats in bee conservation efforts. As part of these plantings, non‐native plants may provide valuable floral resources, but their effects on native bee communities have not been assessed in direct comparison with native pollinator friendly plantings. In this study, we performed a common garden experiment by seeding mixes of 20 native and 20 non‐native pollinator friendly plant species at separate neighboring plots at three sites in Maryland, USA, and recorded flower visitors for 2 years. A total of 3,744 bees (120 species) were collected. Bee abundance and species richness were either similar across plant types (midseason and for abundance also late season) or lower at native than at non‐native plots (early season and for richness also late season). The overall bee community composition differed significantly between native and non‐native plots, with 11 and 23 bee species being found exclusively at one plot type or the other, respectively. Additionally, some species were more abundant at native plant plots, while others were more abundant at non‐natives. Native plants hosted more specialized plant–bee visitation networks than non‐native plants. Three species out of the five most abundant bee species were more specialized when foraging on native plants than on non‐native plants. Overall, visitation networks were more specialized in the early season than in late seasons. Our findings suggest that non‐native plants can benefit native pollinators, but may alter foraging patterns, bee community assemblage, and bee–plant network structures.     

Linkage Rules for Plant–Pollinator Networks: Trait Complementarity or Exploitation Barriers?

Recent attempts to examine the biological processes responsible for the general characteristics of mutualistic networks focus on two types of explanations: nonmatching biological attributes of species that prevent the occurrence of certain interactions (“forbidden links”), arising from trait complementarity in mutualist networks (as compared to barriers to exploitation in antagonistic ones), and random interactions among individuals that are proportional to their abundances in the observed community (“neutrality hypothesis”). We explored the consequences that simple linkage rules based on the first two hypotheses (complementarity of traits versus barriers to exploitation) had on the topology of plant–pollination networks. Independent of the linkage rules used, the inclusion of a small set of traits (two to four) sufficed to account for the complex topological patterns observed in real-world networks. Optimal performance was achieved by a “mixed model” that combined rules that link plants and pollinators whose trait ranges overlap (“complementarity models”) and rules that link pollinators to flowers whose traits are below a pollinator-specific barrier value (“barrier models”). Deterrence of floral parasites (barrier model) is therefore at least as important as increasing pollination efficiency (complementarity model) in the evolutionary shaping of plant–pollinator networks.     

Bird and insect pollinators differ in specialization and potential pollination services along disturbance and resource gradients

Combined studies of the communities and interaction networks of bird and insect pollinators are rare, especially along environmental gradients. Here, we determined how disturbance by fire and variation in sugar resources shape pollinator communities and interactions between plants and their pollinating insects and birds. We recorded insect and bird visits to 21 Protea species across 21 study sites and for 2 years in Fynbos ecosystems in the Western Cape, South Africa. We recorded morphological traits of all pollinator species (41 insect and nine bird species). For each site, we obtained estimates of the time since the last fire (range: 2–25 calendar years) and the Protea nectar sugar amount per hectare (range: 74–62 000 g/ha). We tested how post-fire age and sugar amount influence the total interaction frequency, species richness and functional diversity of pollinator communities, as well as pollinator specialization (the effective number of plant partners) and potential pollination services (pollination service index) of insects and birds. We found little variation in the total interaction frequency, species richness and functional diversity of insect and bird pollinator communities, but insect species richness increased with post-fire age. Pollinator specialization and potential pollination services of insects and birds varied differently along the environmental gradients. Bird pollinators visited fewer Protea species at sites with high sugar amount, while there was no such trend for insects. Potential pollination services of insect pollinators to Protea species decreased with increasing post-fire age and resource amounts, whereas potential pollination services of birds remained constant along the environmental gradients. Despite little changes in pollinator communities, our analyses reveal that insect and bird pollinators differ in their specialization on Protea species and show distinct responses to disturbance and resource gradients. Our comparative study of bird and insect pollinators demonstrates that birds may be able to provide more stable pollination services than insects.     

Pollination networks and functional specialization: a test using Lesser Antillean plant–hummingbird assemblages

Network analysis has in recent years improved our understanding of pollination systems. However, there is very little information about how functionally specialized plants and pollinators interact directly and indirectly in pollination networks. We have developed a parameter, Functional specialization index, to quantify functional specialization in pollination networks. Using this parameter, we examined whether different sized hummingbirds visit a distinct set of flowers in five hummingbird-pollinated plant assemblages from the Lesser Antilles, obtaining a simple relationship between hummingbird body size, network parameter and ecological function. In the Lesser Antilles, functionally specialized hummingbird pollination is distinct for plant species pollinated by the largest hummingbird species, whereas the pollination niche gradually integrates with the insect pollinator community as hummingbird body size decreases. The network approach applied in this study can be used to validate functional specialization and community-level interdependence between plants and pollinators, and it is therefore useful for evaluating and predicting plant resilience to pollinator loss, presently a global concern.     

Lessons from Red Data Books: Plant Vulnerability Increases with Floral Complexity

The architectural complexity of flower structures (hereafter referred to as floral complexity) may be linked to pollination by specialized pollinators that can increase the probability of successful seed set. As plant—pollinator systems become fragile, a loss of such specialized pollinators could presumably result in an increased likelihood of pollination failure. This is an issue likely to be particularly evident in plants that are currently rare. Using a novel index describing floral complexity we explored whether this aspect of the structure of flowers could be used to predict vulnerability of plant species to extinction. To do this we defined plant vulnerability using the Red Data Book of Rare and Threatened Plants of Greece, a Mediterranean biodiversity hotspot. We also tested whether other intrinsic (e.g. life form, asexual reproduction) or extrinsic (e.g. habitat, altitude, range-restrictedness) factors could affect plant vulnerability. We found that plants with high floral complexity scores were significantly more likely to be vulnerable to extinction. Among all the floral complexity components only floral symmetry was found to have a significant effect, with radial-flower plants appearing to be less vulnerable. Life form was also a predictor of vulnerability, with woody perennial plants having significantly lower risk of extinction. Among the extrinsic factors, both habitat and maximum range were significantly associated with plant vulnerability (coastal plants and narrow-ranged plants are more likely to face higher risk). Although extrinsic and in particular anthropogenic factors determine plant extinction risk, intrinsic traits can indicate a plant’s proneness to vulnerability. This raises the potential threat of declining global pollinator diversity interacting with floral complexity to increase the vulnerability of individual plant species. There is potential scope for using plant—pollinator specializations to identify plant species particularly at risk and so target conservation efforts towards them.     

Phylogenetic restriction of plant invasion in drought‐stressed environments: Implications for insect‐pollinated plant communities in water‐limited ecosystems

BackgroundPlant–pollinator community diversity has been found to decrease under conditions of drought stress; however, research into the temporal dimensions of this phenomenon remains limited. In this study, we investigated the effect of seasonal drought on the temporal niche dynamics of entomophilous flowering plants in a water‐limited ecosystem. We hypothesized that closely related native and exotic plants would tend to share similar life history and that peak flowering events would therefore coincide with phylogenetic clustering in plant communities based on expected phenological responses of plant functional types to limitations in soil moisture availability.LocationGaliano Island, British Columbia, Canada.MethodsCombining methods from pollinator research and phylogenetic community ecology, we tested the influence of environmental filtering over plant community phenology across gradients of landscape disturbance and soil moisture. Floral resource availability and community structure were quantified by counts of flowering shoots. We constructed a robust phylogeny to analyze spatial and temporal variation in phylogenetic patterns across the landscape, testing the significance of the observed patterns against a randomly generated community phylogeny. Phylogenetic metrics were then regressed against factors of disturbance and soil moisture availability.ResultsCritical seasonal fluctuations in floral resources coincided with significant phylogenetic clustering in plant communities, with decreasing plant diversity observed under conditions of increasing drought stress. Exotic plant species in the Asteraceae became increasingly pervasive across the landscape, occupying a late season temporal niche in drought‐stressed environments.Main conclusionResults suggest that environmental filtering is the dominant assembly process structuring the temporal niche of plant communities in this water‐limited ecosystem. Based on these results, and trends seen elsewhere, the overall diversity of plant–pollinator communities may be expected to decline with the increasing drought stress predicted under future climate scenarios.     

Restored and remnant Banksia woodlands elicit different foraging behavior in avian pollinators

Pollinators and the pollination services they provide are critical for seed set and self‐sustainability of most flowering plants. Despite this, pollinators are rarely assessed in restored plant communities, where their services are largely assumed to re‐establish. Bird–pollinator richness, foraging, and interaction behavior were compared between natural and restored Banksia woodland sites in Western Australia to assess their re‐establishment in restored sites. These parameters were measured for natural communities of varying size and degree of fragmentation, and restored plant communities of high and low complexity for three years, in the summer and winter flowering of Banksia attenuata and B. menziesii, respectively. Bird visitor communities varied in composition, richness, foraging movement distances, and aggression among sites. Bird richness and abundance were lowest in fragmented remnants. Differences in the composition were associated with the size and degree of fragmentation in natural sites, but this did not differ between seasons. Restored sites and their adjacent natural sites had similar species composition, suggesting proximity supports pollinator re‐establishment. Pollinator foraging movements were influenced by the territorial behavior of different species. Using a network analysis approach, we found foraging behavior varied, with more frequent aggressive chases observed in restored sites, resulting in more movements out of the survey areas, than observed in natural sites. Aggressors were larger‐bodied Western Wattlebirds (Anthochaera chrysoptera) and New Holland Honeyeaters (Phylidonyris novaehollandiae) that dominated nectar resources, particularly in winter. Restored sites had re‐established pollination services, albeit with clear differences, as the degree of variability in the composition and behavior of bird pollinators for Banksias in the natural sites created a broad completion target against which restored sites were assessed. The abundance, diversity, and behavior of pollinator services to remnant and restored Banksia woodland sites were impacted by the size and degree of fragmentation, which in turn influenced bird–pollinator composition, and were further influenced by seasonal changes between summer and winter. Consideration of the spatial and temporal landscape context of restored sites, along with plant community diversity, is needed to ensure the maintenance of the effective movement of pollinators between natural remnant woodlands and restored sites.     

Community structure of pollinating insects and its driving factors in different habitats of Shivapuri‐Nagarjun National Park,Nepal

Insect pollinators are important means for a stable ecosystem. The habitat types play a crucial role in the community composition, abundance, diversity, and species richness of the pollinators. The present study in Shivapuri‐Nagarjun National Park explored the species richness and abundances of insect pollinators in four different habitats and different environmental variables in determining the community composition of the pollinators. Data were collected from 1,500 m to 2,700 m using color pan traps and hand sweeping methods. Non‐Metric Multidimensional Scaling (NMDS) and Redundancy Analysis (RDA) were conducted to show the association between insect pollinators and environmental variables. The results firmly demonstrated that species richness and abundances were higher (158) in Open trail compared to other habitats. The distribution of the pollinator species was more uniform in the Open trail followed by the Grassland. Similarly, a strong positive correlation between flower resources and pollinators'' abundance (R² = .63, P < .001) was found. In conclusion, the Open trail harbors rich insect pollinators in lower elevation. The community structure of the pollinators was strongly influenced by the presence of flowers in the trails.     

Co-flowering neighbors influence the diversity and identity of pollinator groups visiting plant species

The generalization–specialization continuum exhibited in pollination interactions currently receives much attention. It is well-known that the pollinator assemblage of particular species varies temporally and spatially, and therefore the ecological generalization on pollinators may be a contextual attribute. However, the factors causing such variation and its ecological and evolutionary consequences are still poorly understood. This variation can be caused by spatial or temporal variation in the pollinator community, but also by variation in the plant community. Here, we examined how the floral neighbourhood influenced the generalization on pollinators and the composition of pollinators of six plant species differing in generalization levels and main pollinators. The diversity, identity and density of floral species affected both the level of generalization on pollinators and the composition of visitors of particular plant species. Although the relationships to floral neighbourhood varied considerably among species, generalization level and visitation by uncommon pollinators generally increased with floral diversity and richness. The generalization level of the neighbourhood was negatively related to the generalization level of the focal species in two species. The number of flowers of the pollinator-sharing species and the number of flowers of the focal species had different effects on the composition of visits in different species; attributable to differences in facilitation/competition for pollinator attraction. We propose that an important ecological implication of our results is that variation in species interactions caused by the pollination context may result in increased community stability. The main evolutionary implication of our results is that selection on flower and pollinator traits may depend, to an unknown extent, on the composition of the co-flowering plant community.     

Phylogenetic Community Context Influences Pollen Delivery to <Emphasis Type="Italic">Allium cernuum</Emphasis>

Few studies have examined how the number and identity of species in the neighbouring community influences the reproductive success of particular focal species. Pollen delivery, an important component of fitness of sexual plants, is a function of not just the floral traits of any particular individual, but of features of the population and community as it depends on pollinator abundance and preferences. Many pollinators in flowering communities will prefer patches with high floral abundance or diversity yet may exhibit lower floral constancy when more flowering species are present. Thus, pollination efficiency could increase or decrease with increased species richness and this will alter the selection pressures placed upon the floral traits (such as floral colour or reward) of any member of a particular community. Moreover, recent studies have indicated that plant-pollinator networks are phylogenetically structured (pollinators visit related plant species more than expected by chance) and this may be an important factor structuring flowering plant communities. Thus, the sheer number of species within a patch may be less important than the number of closely-related species. We investigate whether species richness or phylogenetic proximity of coflowering species influences the amount and proportion of conspecific pollen delivered to nodding onion, Allium cernuum, in fragment patches of Garry Oak meadows in South Western British Columbia, Canada. We find that pollen delivery depended upon the presence of close relatives far more than on species richness or population density, indicating a central role of the community structure on pollination in flowering plant communities. Insofar as pollen delivery relates to the relative number of seeds produced by members of the community, pollination may in turn determine the community structure of the next generation.     

Distribution and pollination services of wild bees and hoverflies along an altitudinal gradient in mountain hay meadows

Extensively managed and flower‐rich mountain hay meadows, hotspots of Europe''s biodiversity, are subject to environmental and climatic gradients linked to altitude. While the shift of pollinators from bee‐ to fly‐dominated communities with increasing elevation across vegetation zones is well established, the effect of highland altitudinal gradients on the community structure of pollinators within a specific habitat is poorly understood. We assessed wild bee and hoverfly communities, and their pollination service to three plant species common in mountain hay meadows, in eighteen extensively managed yellow oat grasslands (Trisetum flavescens) with an altitudinal gradient spanning approx. 300 m. Species richness and abundance of pollinators increased with elevation, but no shift between hoverflies and wild bees (mainly bumblebees) occurred. Seedset of the woodland cranesbill (Geranium sylvaticum) increased with hoverfly abundance, and seedset of the marsh thistle (Cirsium palustre) increased with wild bee abundance. Black rampion (Phyteuma nigrum) showed no significant response. The assignment of specific pollinator communities, and their response to altitude in highlands, to different plant species underlines the importance of wild bees and hoverflies as pollinators in extensive grassland systems.     

Constructing more informative plant–pollinator networks: visitation and pollen deposition networks in a heathland plant community

Interaction networks are widely used as tools to understand plant–pollinator communities, and to examine potential threats to plant diversity and food security if the ecosystem service provided by pollinating animals declines. However, most networks to date are based on recording visits to flowers, rather than recording clearly defined effective pollination events. Here we provide the first networks that explicitly incorporate measures of pollinator effectiveness (PE) from pollen deposition on stigmas per visit, and pollinator importance (PI) as the product of PE and visit frequency. These more informative networks, here produced for a low diversity heathland habitat, reveal that plant–pollinator interactions are more specialized than shown in most previous studies. At the studied site, the specialization index was lower for the visitation network than the PE network, which was in turn lower than for the PI network. Our study shows that collecting PE data is feasible for community-level studies in low diversity communities and that including information about PE can change the structure of interaction networks. This could have important consequences for our understanding of threats to pollination systems.