Extending the concept of keystone species to communities and ecosystems.

Keystone species are defined as having disproportionate importance in their community. This concept has proved useful and is now often used in conservation ecology. Here, we introduce the concept of keystone communities (and ecosystems) within metacommunities (and metaecosystems). We define keystone and burden communities as communities with impacts disproportionately large (positive or negative respectively) relative to their weight in the metacommunity. We show how a simple metric, based on the effects of single-community removals, can characterise communities along a 'keystoneness' axis. We illustrate the usefulness of this approach with examples from two different theoretical models. We further distinguish environmental heterogeneity from species trait heterogeneity as determinants of keystoneness. We suggest that the concept of keystone communities/ecosystems will be highly beneficial, not only as a fundamental step towards understanding species interactions in a spatial context, but also as a tool for the management of disturbed landscapes.