The study of mutualistic interaction networks has led to valuable insights into ecological and evolutionary processes. However, our understanding of network structure may depend upon the temporal scale at which we sample and analyze network data. To date, we lack a comprehensive assessment of the temporal scale‐dependence of network structure across a wide range of temporal scales and geographic locations. If network structure is temporally scale‐dependent, networks constructed over different temporal scales may provide very different perspectives on the structure and composition of species interactions. Furthermore, it remains unclear how various factors – including species richness, species turnover, link rewiring and sampling effort – act in concert to shape network structure across different temporal scales. To address these issues, we used a large database of temporally‐resolved plant–pollinator networks to investigate how temporal aggregation from the scale of one day to multiple years influences network structure. In addition, we used structural equation modeling to explore the direct and indirect effects of temporal scale, species richness, species turnover, link rewiring and sampling effort on network structural properties. We find that plant–pollinator network structure is strongly temporally‐scale dependent. This general pattern arises because the temporal scale determines the degree to which temporal dynamics (i.e. phenological turnover of species and links) are included in the network, in addition to how much sampling effort is put into constructing the network. Ultimately, the temporal scale‐dependence of our plant–pollinator networks appears to be mostly driven by species richness, which increases with sampling effort, and species turnover, which increases with temporal extent. In other words, after accounting for variation in species richness, network structure is increasingly shaped by its underlying temporal dynamics. Our results suggest that considering multiple temporal scales may be necessary to fully appreciate the causes and consequences of interaction network structure.
Most studies of plant-animal mutualistic networks have come from a temporally static perspective. This approach has revealed general patterns in network structure, but limits our ability to understand the ecological and evolutionary processes that shape these networks and to predict the consequences of natural and human-driven disturbance on species interactions. We review the growing literature on temporal dynamics of plant-animal mutualistic networks including pollination, seed dispersal and ant defence mutualisms. We then discuss potential mechanisms underlying such variation in interactions, ranging from behavioural and physiological processes at the finest temporal scales to ecological and evolutionary processes at the broadest. We find that at the finest temporal scales (days, weeks, months) mutualistic interactions are highly dynamic, with considerable variation in network structure. At intermediate scales (years, decades), networks still exhibit high levels of temporal variation, but such variation appears to influence network properties only weakly. At the broadest temporal scales (many decades, centuries and beyond), continued shifts in interactions appear to reshape network structure, leading to dramatic community changes, including loss of species and function. Our review highlights the importance of considering the temporal dimension for understanding the ecology and evolution of complex webs of mutualistic interactions.
In flowers with poricidal anthers, pollen is not freely accessible and legitimate access is restricted to bees capable of vibrating the anthers. Despite the protection of pollen provided by poricidal anthers, numerous illegitimate, non‐buzz‐pollinating flower visitors rob pollen. We aimed to quantify the influence of functional groups of floral visitors and illegitimate interactions on the network structure to disentangle the flower visitor network into its mutualistic and antagonistic components. We delimited three functional groups of bees based on their pollen collection behaviour in poricidal flowers: large bees that vibrate entire flowers in a single buzzing position (flower buzzing), bees vibrating single anthers in different positions (anther buzzing) and non‐vibrating flower‐damaging or gleaning bees (non‐buzzing). Moreover, we characterized legitimate and illegitimate interactions of co‐occurring and co‐flowering plants and their flower visitors based on the stigma contact during a visit. Since we independently assessed the type of interaction with bee–plant species combinations, we were able to include the behavioural variations of each bee species across different flowers. The networks were modular, with stronger interactions within subsets of species than among the subsets. All modules included a combination of flower‐, anther‐ and non‐buzzing bees, and mutualistic and antagonistic networks were intermingled. Seven bee species shifted their roles across plant species. Specialization in the subset of interactions with pollinators was higher than the overall visitation network. Flower‐buzzing bees were more specialized than anther‐buzzing and non‐buzzing bees, which used virtually all poricidal flowers similarly. Although plants with poricidal anthers shared a specialized mechanism of pollen release, their pollinators were highly dissimilar and formed compartments of interacting species. The interaction‐level approach taken in our study confers a high specificity to the pollinator network, leading to a more complex and realistic picture of mutualistic webs versus its embedded florivory, which are otherwise confounded in pooled networks across flower visitors. A http://onlinelibrary.wiley.com/doi/10.1111/1365-2435.13204/suppinfo is available for this article.
Multi-trophic interactions maintain critical ecosystem functions. Biodiversity is declining globally, while responses of trophic interactions to biodiversity change are largely unclear. Thus, studying responses of multi-trophic interaction robustness to biodiversity change is crucial for understanding ecosystem functioning and persistence. We investigate plant-Hemiptera (antagonism) and Hemiptera-ant (mutualism) interaction networks in response to experimental manipulation of tree diversity. We show increased diversity at both higher trophic levels (Hemiptera and ants) and increased robustness through redundancy of lower level species of multi-trophic interactions when tree diversity increased. Hemiptera and ant diversity increased with tree diversity through non-additive diversity effects. Network analyses identified that tree diversity also increased the number of tree and Hemiptera species used by Hemiptera and ant species, and decreased the specialization on lower trophic level species in both mutualistic and antagonist interactions. Our results demonstrate that bottom-up effects of tree diversity ascend through trophic levels regardless of interaction type. Thus, local tree diversity is a key driver of multi-trophic community diversity and interaction robustness in forests.
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