Ecological network theory hypothesizes a link between structure and stability, but this has mainly been investigated in-silico. In an experimental manipulation, we sequentially removed four generalist plants from real plant-pollinator networks and explored the effects on, and drivers of, species and interaction extinctions, network structure and interaction rewiring. Our results indicate that cumulative species and interaction extinctions increased faster with generalist plant loss than what was expected by co-extinction models, which predicted the survival or extinction of many species incorrectly. In addition, network nestedness decreased, modularity increased, and opportunistic random interactions and structural unpredictability emerged, which are all indicators of network instability and fragility.Conversely, interaction reorganization (rewiring) was high, asymmetries between network levels emerged as plants increased their centrality. From the experimental manipulations of real networks, our study shows how plant-pollinator network structure has low stability and changes towards a more fragile state when generalist plants are lost.
IntroductionInteractions are organized in complex networks, and the way these structures react to disturbance is crucial for understanding network functioning, their ability to buffer negative impacts and also for their conservation 1-4 . This is usually verified with "attack tolerance tests" that assess the functionality of a system after knocking out its important components 5 . In ecology, such tests usually consist in removing all species in one trophic level and then in assessing how many species in another level lost all interactions 6,7 . So far, in pollination networks, this has been addressed mainly theoretically with numerical simulations that show a higher rate of pollinator extinction when highly linked plants are removed 7-10 . However, these theoretical predictions were not compared to empirical data from similar manipulations, which is urgently needed to assess their reliability 11 . Manipulative experiments of plant-pollinator networks can illuminate the factors maintaining network stability and the processes of network reorganization. For instance, previous experiments removing only one generalist plant 12,13 showed that networks are quite stable to this loss, and that other species occupy the role in the network of the removed species. Conversely, when multiple invasive plants are removed, network interaction diversity and generalisation are impacted 14 , indicating that losing multiple species can strongly affect real networks. Moreover, after disturbance, network stability could depend on the amount of interaction rewiring 18 , i.e. foragers' ability to use alternative resources after depletion or disappearance of those previously used 15-17 . Rewiring and the establishment of interactions between plants and pollinators may be regulated by several ecological drivers, such as species trait matching 19,20 , flower's rewards 21,22 or species abundances 23,24 . Sim...