Potential warm-stage microrefugia for alpine plants: Feedback between geomorphological and biological processes

During interglacial stages, microrefugia are sites that support locally favorable climates within larger areas with unfavorable warmer climates. Despite recent theoretical representations of microrefugia, an appropriate ecological characterization is still lacking, mostly for warm periods. Across mountain/alpine areas, cold-adapted plant species could adopt different strategies to manage the effects of climate warming: (A) migration toward higher elevations and summits; (B) in situ resilience of communities and species populations within microrefugia; and C) adaptation and evolution by genetic differentiation. This review aims to distinguish and characterize from an ecological perspective glacial, nival, periglacial and composite landforms and deposits that may function as potential microrefugia during interglacial warm periods.We conducted a literature screening related to the geomorphological processes and landforms associated with vegetation and plant communities in alpine/mountain environments of Europe. They include glacial deposits rock glaciers, debris-covered glaciers, composite cones and channels. In Alpine regions, geomorphologic niches that constantly maintain cold-air pooling and temperature inversions are the main candidates for microrefugia. Within such microrefugia, microhabitat diversity modulates the responses of plants to disturbances caused by geomorphologic processes and supports their aptitude for surviving under extreme conditions on unstable surfaces in isolated patches. Currently, European marginal mountain chains may be considered as examples of macrorefugia where relict boreo-alpine species persist within peculiar geomorphological niches that act as microrefugia.This review contributes to identifying potential warm-stage microrefugia areas across alpine and mountain regions and determining certain landforms that play or may play such role under global-change scenarios. The occurrence of warm-stage microrefugia within these locations may be of great importance for the modeling of future distributions of species and assessing the risk of extinction for alpine species. Microrefugia may have important implications in micro-evolutionary processes that occur across alternating climatic phases.