Local topographic and edaphic factors largely predict shrub encroachment in Mediterranean drylands

Shrub encroachment influences several ecosystem services in drylands worldwide. Yet, commonly used strategies to reduce encroachment show a low medium-term success, calling for a better understanding of its causes. Previous works identified multiple drivers responsible for this phenomenon, including anthropogenic and environmental causes. However, the relative effect of climate, topography and edaphic factors on shrub encroachment is not fully understood nor has been properly quantified in Mediterranean Basin drylands. Also, understanding how these drivers lead to changes in plant communities' functional traits associated to shrub encroachment is crucial, considering traits influence ecosystem processes and associated ecosystem services. Here, we studied the understory of a Mediterranean dryland ecosystem composed of savanna-like Holm-oak woodlands, along a regional climatic gradient. We specifically assessed (i) how climatic, topographic and edaphic factors influence understory relative shrub cover (RSC) and (ii) their direct and indirect effects (via RSC) on plant functional traits. We studied the mean and diversity of 12 functional traits related to plant regeneration, establishment, and dispersal, at the community-level. We found that, under similar low-intensity land use, topographic and edaphic factors, namely slope variations and soil C:N ratio, were the most important predictors of shrub encroachment, determining communities' functional characteristics. Climate, namely summer precipitation, had a much lesser influence. Our model explained 52% of the variation in relative shrub cover. Climate had a stronger effect on a set of functional traits weakly involved in shrub encroachment, related to flowering and dispersal strategies. We show that shrub encroachment is largely predicted by topo-edaphic factors in Mediterranean drylands subject to conventional low-intensity land use. Hence, management strategies to reduce encroachment need to take these drivers into account for efficient forecasting and higher cost-effectiveness. Our results suggest that climate change might not greatly impact shrub encroachment in the Mediterranean Basin, but may affect functional structure and reduce functional diversity of plant communities, thus affecting ecosystem functioning.