Context-dependent effects of fire and browsing on woody alien invasion in mountain ecosystems.

Anthropogenic activities have increased disturbances and alien woody invasion in mountain ecosystems worldwide. Whether disturbances promote or counteract upward movement of woody aliens is poorly understood. We assessed if the most successful woody invader of low mountains of central Argentina (Gleditsia triacanthos) might expand its elevational distribution in response to the principal disturbances of these ecosystems (fire and livestock browsing) across increasing climatic severity. We assessed seedling emergence, growth and mycorrhizal colonization on sown plots distributed in burned and unburned sites, with and without browsing at the lower and upper elevation belts (i.e. 1000 and 2400 m a.s.l.). Additionally, several abiotic variables were measured to relate their influence on the seedling establishment. Disturbances reduced seedling emergence at both elevations. Burned conditions increased seedling growth and arbuscular colonization only in the lower belt. Seedling success (total seedling biomass per plot) was not modified by disturbances at the upper elevation, but was reduced by browsing and enhanced by fire in the lower elevation. The overall reduction in seedling emergence and growth in the upper elevation despite the higher soil nutrient content places climate as the strongest regulator of G. triacanthos seedling establishment. Accordingly, climate rather than disturbances would be the main limiting factor of upward expansion of this woody alien. Our findings differ from general patterns described for mountain invasion by herbaceous species, highlighting that mountain invasibility is highly growth-form dependent, and that upper range expansion by woody aliens interacting with multiple disturbances should be assessed worldwide.