On the structure of species-function participation in multilayer ecological networks.

Understanding how biotic interactions shape ecosystems and impact their functioning, resilience and biodiversity has been a sustained research priority in ecology. Yet, traditional assessments of ecological complexity typically focus on species-species interactions that mediate a particular function (e.g., pollination), overlooking both the synergistic effect that multiple functions might develop as well as the resulting species-function participation patterns that emerge in ecosystems that harbor multiple ecological functions. Here we propose a mathematical framework that integrates various types of biotic interactions observed between different species. Its application to recently collected data of an islet ecosystem-reporting 1537 interactions between 691 plants, animals and fungi across six different functions (pollination, herbivory, seed dispersal, decomposition, nutrient uptake, and fungal pathogenicity)-unveils a non-random, nested structure in the way plant species participate across different functions. The framework further allows us to identify a ranking of species and functions, where woody shrubs and fungal decomposition emerge as keystone actors whose removal have a larger-than-random effect on secondary extinctions. The dual insight-from species and functional perspectives-offered by the framework opens the door to a richer quantification of ecosystem complexity and to better calibrate the influence of multifunctionality on ecosystem functioning and biodiversity.