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.
Summary Harsh abiotic conditions – such as low temperatures that lead to spring and summer frost events in high‐elevation and high‐latitude ecosystems – can have strong negative consequences for plant growth, survival and reproduction. Despite the predicted increase in episodic frost events under continued climate change in some ecosystems, our general understanding of the factors associated with frost sensitivity of reproductive and vegetative plant structures in natural plant communities is limited. The timing of growth and reproduction may be an important strategy by which plants can avoid frost. In this study, we experimentally investigated the frost sensitivity of eight long‐lived perennial herbaceous plant species from a subalpine ecosystem in the Colorado Rocky Mountains, USA. The study taxa represent four congeneric pairs from four flowering plant families; within each pair, there is a species with early and late growth and reproductive phenology. Thus, we control for evolutionary history – and therefore additional traits shared through common ancestry – to some degree, while examining the influence of phenology on frost sensitivity. Specifically, we compared frost sensitivity of vegetative and reproductive structures for each species and asked whether frost sensitivity was similar between species within congeneric pairs or, instead, was related to phenology (i.e. differences in the timing of growth and reproduction). For most species (6 of 8), flowers were more sensitive to frost than leaves. Within most congeneric pairs (3 of 4), the leaves of species with later phenology were significantly more sensitive to frost than the leaves of species with earlier phenology. For flowers, the later flowering species were more sensitive in two of the four congeneric pairs. Synthesis. This study contributes to our general understanding of factors related to interspecific differences in plant sensitivity to episodic frost events of naturally occurring species. The increased frost sensitivity of reproductive structures compared to vegetative structures may be a widespread pattern for long‐lived perennial plants. Furthermore, we find evidence for a trade‐off between phenology and frost sensitivity, whereby species with later phenology exhibit higher frost sensitivity compared to species with earlier phenology. These results have implications for plant populations, species interactions and ecological communities.
Pollination is essential for ecosystem functioning, yet our understanding of the empirical consequences of species loss for plant–pollinator interactions remains limited. It is hypothesized that the loss of abundant and generalized (well‐connected) species from a pollination network will have a large effect on the remaining species and their interactions. However, to date, relatively few studies have experimentally removed species from their natural setting to address this hypothesis. We investigated the consequences of losing an abundant, generalist native species from a series of plant–pollinator networks by experimentally removing the flowers of Helianthella quinquenervis (Asteraceae) from half of a series of 10 paired plots (15 m diameter) within a subalpine ecosystem. We then asked how the localized loss of this species influenced patterns of pollinator visitation, floral visitor composition, and interaction network structure. The experimental removal of Helianthella flowers led to an overall decline in plot‐level pollinator visitation rates and shifts in pollinator composition. Species‐level responses to floral removal differed between the two other abundant, co‐flowering plants in our experiment: Potentilla pulcherrima received higher visitation rates, whereas Erigeron speciosus visitation rates did not change. Experimental floral removal altered the structural properties of the localized plant–pollinator networks such that they were more specialized, less nested, and less robust to further species loss. Such changes to interaction network structure were consistently driven more by species turnover than by interaction rewiring. Our findings suggest that the local loss of an abundant, well‐linked, generalist plant can bring about diverse responses within intact pollination networks, including potential competitive and facilitative effects for individual species, changes to network structure that may render them more sensitive to future change, but also numerous changes to interactions that may also suggest flexibility in response to species loss.
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