Enough space in a warmer world? Microhabitat diversity and small-scale distribution of alpine plants on mountain summits

Aim: Global warming is predicted to shift distributions of mountain species upwards, driven by a release from climatic restrictions at their upper distribution limit and increased biotic pressure at their lower distribution limit. In alpine ecosystems, which are characterized by large microclimatic diversity and sparse vegetation cover, the relative importance of abiotic and biotic drivers for species distribution is poorly understood. To disentangle abiotic and biotic mechanisms affecting distributions of alpine species, we investigated how alpine plant species with differing elevational ranges and frequency trends over the past century differ in their microhabitat distribution, and how they respond to neighbouring vegetation. Location: A total of 11 summits (2635-3410 m a.s.l.) in SE-Switzerland. Methods: We quantified the microscale abundance of 12 species in relation to biogeographic (frequency trend, i.e., change in occurrences over the past century, and elevational range on summits) and local microhabitat characteristics (temperature, substrate type). We assessed species size traits in relation to neighbouring vegetation characteristics to investigate possible neighbour interactions. Results: Species with increasing frequency on summits over the past century were most abundant on scree and warmer slopes. Species with negative or stable frequency trends on summits were more abundant on organic soil and colder slopes. The preferred microhabitats of the latter were rarest overall, decreased with increasing elevation, and had the most competitive neighbours. Size of one high-alpine specialist, Ranunculus glacialis was negatively related to cover of neighbouring vegetation, whereas other species showed no response to neighbours. Main conclusions: Long-term frequency trends of species correlate with their microhabitat association. Species with most negative frequency trends show preferences for the rarest microhabitat conditions, where they likely experience higher competitive pressure in a warming climate. This finding emphasizes the importance of characterizing microhabitat associations and microclimatic diversity to assess present and future distributions of alpine plant species.