Invasive Mutualists Erode Native Pollination Webs

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Abstract
Plant–animal mutualisms are characterized by weak or asymmetric mutual dependences between interacting species, a feature that could increase community stability. If invasive species integrate into mutualistic webs, they may alter web structure, with consequences for species persistence. However, the effect of alien mutualists on the architecture of plant–pollinator webs remains largely unexplored. We analyzed the extent of mutual dependency between interacting species, as a measure of mutualism strength, and the connectivity of 10 paired plant–pollinator webs, eight from forests of the southern Andes and two from oceanic islands, with different incidences of alien species. Highly invaded webs exhibited weaker mutualism than less-invaded webs. This potential increase in network stability was the result of a disproportionate increase in the importance and participation of alien species in the most asymmetric interactions. The integration of alien mutualists did not alter overall network connectivity, but links were transferred from generalist native species to super-generalist alien species during invasion. Therefore, connectivity among native species declined in highly invaded webs. These modifications in the structure of pollination webs, due to dominance of alien mutualists, can leave many native species subject to novel ecological and evolutionary dynamics. Plant–animal mutualisms are characterized by weak or asymmetric mutual dependences between interacting species, such that if a plant species depends strongly on an animal species, the animal typically depends weakly on the plant, and vice versa. This limited reciprocal dependency, or “mutualism strength,” might increase species persistence by buffering plant and animal species against the extinction of any of their partners. Many plant–pollinator networks include a fraction of alien species, and it is not clear how these invaders might affect the structure of pollination webs. We analyzed 10 paired plant–pollinator webs, eight from forests of the southern Andes and two from oceanic islands, with different incidences of alien species. Highly invaded webs exhibited, on average, weaker mutualistic interactions, and hence a potential increase in network stability, than less-invaded webs. This was due to a disproportionate increase in the participation of some alien species in the most asymmetric interactions and their role as central nodes in the structure of the most invaded pollination webs. The increase in alien dominance involves the usurpation of interaction links, decreasing connectivity among native mutualists. Thus, many native species that rely on native generalists for either reproduction or survivorship become highly dependent on these super-generalist alien mutualists.