Trade-off between competition and facilitation defines gap colonization in mountains

Recent experimental observations show that gap colonization in small-stature (e.g. grassland and dwarf shrubs) vegetation strongly depends on the abiotic conditions within them. At the same time, within-gap variation in biotic interactions such as competition and facilitation, caused by distance to the gap edge, would affect colonizer performance, but a theoretical framework to explore such patterns is missing. Here, we model how competition, facilitation and environmental conditions together determine the small-scale patterns of gap colonization along a cold gradient in mountains, by simulating colonizer survival in gaps of various sizes. Our model adds another dimension to the known effects of biotic interactions along a stress gradient by focussing on the trade-off between competition and facilitation in the within-gap environment. We show that this trade-off defines a peak in colonizer survival at a specific distance from the gap edge, which progressively shifts closer to the edge as the environment gets colder, ultimately leaving a large fraction of gaps unsuitable for colonization in facilitation-dominated systems. This is reinforced when vegetation size and temperature amelioration are manipulated simultaneously with temperature in order to simulate an elevational gradient more realistically. Interestingly, all other conditions being equal, the magnitude of the realized survival peak was always lower in large than in small gaps, making large gaps harder to colonize. The model is relevant to predict effects of non-native plant invasions and climate warming on colonization processes in mountains.