Facilitation and Competition on Gradients in Alpine Plant Communities

We conducted a neighbor removal experiment in natural alpine plant communities of the southwestern Alps to test for the relative importance of competitive and facilitative interactions along elevational and topographical gradients. The experimental sites were chosen to encompass most of the floristic diversity observed along gradients of elevation and topography, which are the two main ecological gradients associated with alpine plant communities in the western Alps. The effects of neighbor removal on the survival, aboveground biomass, and reproduction of five target species were tested at each of six experimental sites. Using biomass data, we calculated relative competitive index (RCI) and log response ratio (LRR) as measures of interaction strength and direction. We found highly significant shifts from strong competitive effects in low and sheltered sites to strong facilitative responses in high and exposed sites. When experimental results were integrated with gradient analyses, we found that the responses of particular alpine plant species to neighbor removal generally depended on the species' position on elevational and topographical gradients. When neighbors were removed from around target species at experimental sites that were lower in elevation than the distributional mean of the target species, biomass generally increased. When neighbors were removed from around target species at experimental sites that were higher in elevation than the distributional mean of the target species, biomass decreased. In other words, facilitation appeared to allow species from lower elevations to move up the gradient, but competition at low elevations appeared to restrict species from higher elevations from moving down the gradient. In high and exposed sites, experimental evidence for facilitation was coupled to small-scale spatial associations among species, but spatial disassociation was not coupled to experimental evidence for competition at any sites. We conclude that the distribution and abundance of many species in high-elevation communities of the western Alps appears to be enhanced by neighbors, and that species continua commonly observed along environmental gradients are the result of both negative and positive plant interactions.