The 'filtering' metaphor revisited: competition and environment jointly structure invasibility and coexistence.

'Filtering', or the reduction in species diversity that occurs because not all species can persist in all locations, is thought to unfold hierarchically, controlled by the environment at large scales and competition at small scales. However, the ecological effects of competition and the environment are not independent, and observational approaches preclude investigation into their interplay. We use a demographic approach with 30 plant species to experimentally test: (i) the effect of competition on species persistence in two soil moisture environments, and (ii) the effect of environmental conditions on mechanisms underlying competitive coexistence. We find that competitors cause differential species persistence across environments even when effects are lacking in the absence of competition, and that the traits which determine persistence depend on the competitive environment. If our study had been observational and trait-based, we would have erroneously concluded that the environment filters species with low biomass, shallow roots and small seeds. Changing environmental conditions generated idiosyncratic effects on coexistence outcomes, increasing competitive exclusion of some species while promoting coexistence of others. Our results highlight the importance of considering environmental filtering in the light of, rather than in isolation from, competition, and challenge community assembly models and approaches to projecting future species distributions.