Frequency and intensity of facilitation reveal opposing patterns along a stress gradient

Disentangling the different processes structuring ecological communities is a long-standing challenge. In species-rich ecosystems, most emphasis has so far been given to environmental filtering and competition processes, while facilitative interactions between species remain insufficiently studied. Here, we propose an analysis framework that not only allows for identifying pairs of facilitating and facilitated species, but also estimates the strength of facilitation and its variation along environmental gradients. Our framework combines the analysis of both co-occurrence and co-abundance patterns using a moving window approach along environmental gradients to control for potentially confounding effects of environmental filtering in the co-abundance analysis. We first validate our new approach against community assembly simulations, and exemplify its potential on a large 1,134 plant community plots dataset. Our results generally show that facilitation intensity was strongest under cold stress, whereas the proportion of facilitating and facilitated species was higher under drought stress. Moreover, the functional distance between individual facilitated species and their facilitating species significantly changed along the temperature-moisture gradient, and seemed to influence facilitation intensity, although no general positive or general negative trend was discernible among species. The main advantages of our robust framework are as follows: It enables detecting facilitating and facilitated species in species-rich systems, and it allows identifying the directionality and intensity of facilitation in species pairs as well as its variation across long environmental gradients. It thus opens numerous opportunities for incorporating functional (and phylogenetic) information in the analysis of facilitation patterns. Our case study indicated high complexity in facilitative interactions across the stress gradient and revealed new evidence that facilitation, similarly to competition, can operate between functionally similar and dissimilar species. Extending the analyses to other taxa and ecosystems will foster our understanding how complex interspecific interactions promote biodiversity.