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.     

Effects of different types of low‐intensity management on plant‐pollinator interactions in Estonian grasslands

In the face of global pollinator decline, extensively managed grasslands play an important role in supporting stable pollinator communities. However, different types of extensive management may promote particular plant species and thus particular functional traits. As the functional traits of flowering plant species (e.g., flower size and shape) in a habitat help determine the identity and frequency of pollinator visitors, they can also influence the structures of plant−pollinator interaction networks (i.e., pollination networks). The aim of this study was to examine how the type of low‐intensity traditional management influences plant and pollinator composition, the structure of plant−pollinator interactions, and their mediation by floral and insect functional traits. Specifically, we compared mown wooded meadows to grazed alvar pastures in western Estonia. We found that both management types fostered equal diversity of plants and pollinators, and overlapping, though still distinct, plant and pollinator compositions. Wooded meadow pollination networks had significantly higher connectance and specialization, while alvar pasture networks achieved higher interaction diversity at a standardized sampling of interactions. Pollinators with small body sizes and short proboscis lengths were more specialized in their preference for particular plant species and the specialization of individual pollinators was higher in alvar pastures than in wooded meadows. All in all, the two management types promoted diverse plant and pollinator communities, which enabled the development of equally even and nested pollination networks. The same generalist plant and pollinator species were important for the pollination networks of both wooded meadows and alvar pastures; however, they were complemented by management‐specific species, which accounted for differences in network structure. Therefore, the implementation of both management types in the same landscape helps to maintain high species and interaction diversity.     

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.     

生长季放牧对高寒草甸传粉网络的影响

植物-传粉者相互作用面临人类活动的威胁。在青藏高原地区,放牧是一项主要的人类活动干扰,过度放牧导致高寒草甸植被严重退化。然而在该区域放牧如何影响植物-传粉者相互作用网络还不得而知。在青藏高原东部的高寒草甸选取了两个研究样点,每个样点包括一块禁牧样地（生长季禁牧）和放牧样地（全年放牧）。在2016年至2018年每年的生长季（7月和8月）进行了连续3年观测,共构建16个传粉网络。结果发现,在研究区域的高寒草甸生态系统中,传粉者群落的物种组成以双翅目昆虫为主。放牧后双翅目、鳞翅目以及鞘翅目传粉者的种类数减少,但膜翅目传粉者的种类数未受到放牧的影响。放牧干扰显著降低了群落中植物、传粉者以及它们之间相互作用的多样性,但对传粉网络的嵌套性和特化程度（H₂''）没有显著影响,说明网络的稳定性和恢复力没有受到放牧的影响。探讨了放牧对传粉网络的影响,发现区域放牧强度过大,降低了传粉昆虫和传粉网络的多样性。未来需要进一步深入研究高寒草甸生态系统中放牧强度对传粉网络的影响模式,以期为合理的放牧制度模式提供理论依据。     

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.     

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.     

Population dynamics of plant and pollinator communities: stability reconsidered

Plant-pollinator networks are systems of outstanding ecological and economic importance. A particularly intriguing aspect of these systems is their high diversity. However, earlier studies have concluded that the specific mechanisms of plant-pollinator interactions are destabilizing and should lead to a loss of diversity. Here we present a mechanistic model of plant and pollinator population dynamics with the ability to represent a broad spectrum of interaction structures. Using this model, we examined the influence of pollinators on the stability of a plant community and the relationship between pollinator specialization and stability. In accordance with earlier work, our results show that plant-pollinator interactions may severely destabilize plant coexistence, regardless of the degree of pollinator specialization. However, if plant niche differentiation, a classical stabilizing mechanism, is sufficiently strong to overcome the minority disadvantage with respect to pollination, interactions with pollinators may even increase the stability of a plant community. In addition to plant niche differentiation, the relationship between specialization and stability depends on a number of parameters that affect pollinator growth rates. Our results highlight the complex effects of this particular type of mutualism on community stability and call for further investigations of the mechanisms of diversity maintenance in plant-pollinator systems.     

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.     

Tolerance of pollination networks to species extinctions

Mutually beneficial interactions between flowering plants and animal pollinators represent a critical 'ecosystem service' under threat of anthropogenic extinction. We explored probable patterns of extinction in two large networks of plants and flower visitors by simulating the removal of pollinators and consequent loss of the plants that depend upon them for reproduction. For each network, we removed pollinators at random, systematically from least-linked (most specialized) to most-linked (most generalized), and systematically from most- to least-linked. Plant species diversity declined most rapidly with preferential removal of the most-linked pollinators, but declines were no worse than linear. This relative tolerance to extinction derives from redundancy in pollinators per plant and from nested topology of the networks. Tolerance in pollination networks contrasts with catastrophic declines reported from standard food webs. The discrepancy may be a result of the method used: previous studies removed species from multiple trophic levels based only on their linkage, whereas our preferential removal of pollinators reflects their greater risk of extinction relative to that of plants. In both pollination networks, the most-linked pollinators were bumble-bees and some solitary bees. These animals should receive special attention in efforts to conserve temperate pollination systems.     

Adaptive foraging allows the maintenance of biodiversity of pollination networks

Pollination systems are recognized as critical for the maintenance of biodiversity in terrestrial ecosystems. Therefore, the understanding of mechanisms that promote the integrity of those mutualistic assemblages is an important issue for the conservation of biodiversity and ecosystem function. In this study we present a new population dynamics model for plant–pollinator interactions that is based on the consumer–resource approach and incorporates a few essential features of pollination ecology. The model was used to project the temporal dynamics of three empirical pollination network, in order to analyze how adaptive foraging of pollinators (AF) shapes the outcome of community dynamics in terms of biodiversity and network robustness to species loss. We found that the incorporation of AF into the dynamics of the pollination networks increased the persistence and diversity of its constituent species, and reduced secondary extinctions of both plants and animals. These findings were best explained by the following underlying processes: 1) AF increased the amount of floral resources extracted by specialist pollinators, and 2) AF raised the visitation rates received by specialist plants. We propose that the main mechanism by which AF enhanced those processes is (trophic) niche partitioning among animals, which in turn generates (pollen vector) niche partitioning among plants. Our results suggest that pollination networks can maintain their stability and diversity by the adaptive foraging of generalist pollinators.     

Resource heterogeneity, diet shifts and intra-cohort competition: effects on size divergence in YOY fish

The pollination syndrome hypothesis has provided a major conceptual framework for how plants and pollinators interact. However, the assumption of specialization in pollination systems and the reliability of floral traits in predicting the main pollinators have been questioned recently. In addition, the relationship between ecological and evolutionary specialization in pollination interactions is still poorly understood. We used data of 62 plant species from three communities across southern Norway to test: (1) the relationships between floral traits and the identity of pollinators, (2) the association between floral traits (evolutionary specialization) and ecological generalization, and (3) the consistency of both relationships across communities. Floral traits significantly affected the identity of pollinators in the three communities in a way consistent with the predictions derived from the pollination syndrome concept. However, hover flies and butterflies visited flowers with different shapes in different communities, which we mainly attribute to among-community variation in pollinator assemblages. Interestingly, ecological generalization depended more on the community-context (i.e. the plant and pollinator assemblages in the communities) than on specific floral traits. While open yellow and white flowers were the most generalist in two communities, they were the most specialist in the alpine community. Our results warn against the use of single measures of ecological generalization to question the pollination syndrome concept, and highlight the importance of community comparisons to assess the pollination syndromes, and to understand the relationships between ecological and evolutionary specialization in plant-pollinator interactions.     

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.     

Experimental reduction of pollinator visitation modifies plant–plant interactions for pollination

The strength of interactions between plants for pollination depends on the abundance of plants and pollinators in the community. The abundance of pollinators may influence plant associations and densities at which individual fitness is maximized. Reduced pollinator visitation may therefore affect the way plant species interact for pollination. We experimentally reduced pollinator visitation to six pollinator‐dependent species (three from an alpine and three from a lowland community in Norway) to study how interactions for pollination were modified by reduced pollinator availability. We related flower visitation, pollen limitation and seed set to density of conspecifics and pollinator‐sharing heterospecifics inside 30 dome‐shaped cages partially covered with fishnet (experimental plots) and in 30 control plots. We expected to find stronger interactions between plants in experimental compared to controls plots. The experiment modified plant–plant interactions for pollination in all the six species; although for two of them neighbourhood interactions did not affect seed set. The pollen limitation and seed set data showed that reduction of pollinator visits most frequently resulted in novel and/or stronger interactions between plants in the experimental plots that did not occur in the controls. Although the responses were species‐specific, there was a tendency for increasing facilitative interactions with conspecific neighbours in experimental plots where pollinator availability was reduced. Heterospecifics only influenced pollination and fecundity in species from the alpine community and in the experimental plots, where they competed with the focal species for pollination. The patterns observed for visitation rates differed from those for fecundity, with more significant interactions between plants in the controls in both communities. This study warns against the exclusive use of visitation data to interpret plant–plant interactions for pollination, and helps to understand how plant aggregations may buffer or intensify the effects of a pollinator loss on plant fitness.     

Plant species roles in pollination networks: an experimental approach

Pollination is an important ecosystem service threatened by current pollinator declines, making flower planting schemes an important strategy to recover pollination function. However, ecologists rarely test the attractiveness of chosen plants to pollinators in the field. Here, we experimentally test whether plant species roles in pollination networks can be used to identify species with the most potential to recover plant–pollinator communities. Using published pollination networks, we calculated each plant's centrality and chose five central and five peripheral plant species for introduction into replicate experimental plots. Flower visitation by pollinators was recorded in each plot and we tested the impact of introduced central and peripheral plant species on the pollinator and resident plant communities and on network structure. We found that the introduction of central plant species attracted a higher richness and abundance of pollinators than the introduction of peripheral species, and that the introduced central plant species occupied the most important network roles. The high attractiveness of central species to pollinators, however, did not negatively affect visitation to resident plant species by pollinators. We also found that the introduction of central plant species did not affect network structure, while networks with introduced peripheral species had lower centralisation and interaction evenness than networks with introduced central species. To our knowledge, this is the first time species network roles have been tested in a field experiment. Given that most restoration projects start at the plant community, being able to identify the plants with the highest potential to restore community structure and functioning should be a key goal for ecological restoration.     

The relative importance of color signaling for plant generalization in pollination networks

Plant strategies frequently vary from opportunistic pollination to specialization to single pollinators within the same community. Unraveling the proximate mechanisms that determine the degree of plant generalization to pollinators has become a primary goal of pollination ecology. Color signaling is a potentially important mechanism because it is well established that many pollinators use color stimuli to locate food items. Until now, studies on the importance of color signaling in structuring pollination networks have not considered floral coloration as it is perceived by pollinators. Here, we use a framework recently developed for network analyses to compare the relative importance of color matching (i.e. the degree of phenotypic matching between flower coloration and pollinators’ visual system) and other variables (phylogeny, co‐abundance and spatiotemporal overlap between plants and pollinators) for plant generalization. We analyzed 25 000 visits in three temperate regions. We show that color matching in combination with spatiotemporal overlap or co‐abundance significantly influences plant generalization in one of the three regions. We suggest that intense human activities in two regions have decreased the mean level of color matching, potentially disrupting the communication between plants and pollinators. This study illuminates how the sensory ecology of pollinators contributes to structure a highly diversified pollination network.     

Plant–pollinator network assembly along the chronosequence of a glacier foreland

Forelands of retreating glaciers offer an ideal model system to study community assembly processes during primary succession. As plants colonize the area that is freed from ice they should be accompanied by their pollinators to successfully reproduce and spread. However, little is known about the assembly of plant–pollinator networks. We therefore used quantitative network analysis to study the structure of plant–pollinator interactions at seven sites representing a chronosequence from 8 to 130 years since deglaciation on the foreland of the Morteratsch glacier (southeastern Switzerland). At these sites, individual visits of plant flowers by insects were recorded throughout the flowering season. Species richness of insect‐pollinated plants and plant‐pollinating insects, together with measures of interaction diversity and evenness, increased along the chronosequence at least for the first 80 years after deglaciation. Bees were the most frequent flower visitors at the two youngest sites, whereas flies dominated in mature communities. Pollinator generalization (the number of visited plant species weighted by interaction strength), but not plant generalization, strongly increased during the primary succession. This was reflected in a pronounced decline in network level specialization (measured as Blüthgen's H₂’) and interaction strength asymmetry during the first 60 years along the chronosequence, while nestedness increased along the chronosequence. Thus, our findings contradict niche‐theoretical predictions of increasing specialization of pollination systems during succession, but are in agreement with expectations from optimal foraging theory, predicting an increase in pollinator generalization with higher plant diversity but similar flower abundance, and an increase in diet breadth at higher pollinator densities during primary succession.     

Do pollination syndromes cause modularity and predict interactions in a pollination network in tropical high‐altitude grasslands?

The concept of pollination syndromes has been widely questioned, since plant–pollinator interactions have proved to be more generalist than was previously thought. We examined whether the network of a tropical high‐altitude grassland contained groups of plants and pollinators that interact preferentially with each other. A general binary matrix was created. To assess the robustness of myophily, in all analyses we considered: 1) the whole network, 2) the network after the wasps were removed, and 3) the network after the flies were removed. For each network we evaluated whether: 1) the observed interactions were more related to syndromes than expected by chance, compared to an expected matrix; 2) there was a modular structure; 3) the modules found were more related to syndromes than expected by chance, compared to another expected matrix; 4) the syndromes were equally robust. For this analysis, the general matrix was subdivided into smaller matrices that included each pollination syndrome separately. To test the influence of the functional groups of pollinators and the phylogeny of plants, in addition to the general matrix, we also considered the first expected matrix, a quantitative functional group and a plant phylogeny matrix. The pollination syndromes determined the pattern of interactions in the network: 69% of the total interactions resulted from the functional group of pollinators predicted by the plant syndrome. The network showed greater modularity (13 modules) than expected by chance, mostly consisting of the expected functional groups of pollinators and plant syndromes. The modules were associated with pollination syndromes more than was predicted by chance. Most of the variation in interactions was explained by functional groups of pollinators or by plant syndromes. Plant phylogeny did not account for a significant amount of variation in the interactions. Our findings support the concept of pollination syndromes. However, the interactions were not equally predicted by different pollination syndromes, and the accuracy of the prediction was strongest for ornithophily and melittophily.     

Pollinator specialization increases with a decrease in a mass‐flowering plant in networks inferred from DNA metabarcoding

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Between‐year changes in community composition shape species’ roles in an Arctic plant–pollinator network

Inter‐annual turnover in community composition can affect the richness and functioning of ecological communities. If incoming and outgoing species do not interact with the same partners, ecological functions such as pollination may be disrupted. Here, we explore the extent to which turnover affects species’ roles – as defined based on their participation in different motifs positions – in a series of temporally replicated plant–pollinator networks from high‐Arctic Zackenberg, Greenland. We observed substantial turnover in the plant and pollinator assemblages, combined with significant variation in species’ roles between networks. Variation in the roles of plants and pollinators tended to increase with the amount of community turnover, although a negative interaction between turnover in the plant and pollinator assemblages complicated this trend for the roles of pollinators. This suggests that increasing turnover in the future will result in changes to the roles of plants and likely those of pollinators. These changing roles may in turn affect the functioning or stability of this pollination network.