Shrubs facilitating seedling performance in ungulate-dominated systems: biotic versus abiotic mechanisms of plant facilitation

Shrubs are important microsites that facilitate seedling performance in abiotically stressed environments. However, shrub facilitation might work differently in ecosystems with high biotic stress (e.g., ungulate-dominated environments). We wonder whether ungulates through their preferences for different shrubs affect the facilitation process of seedlings, and whether this process remains consistent across different levels of abiotic stress. We study seedling performance (survival and growth) of a Mediterranean oak (Quercus pyrenaica) in three microsites (open, preferred and non-preferred shrubs by ungulates) for two contrasting climatic environments and throughout a complete growing season (increasing abiotic stress). Overall, there was a positive effect of shrubs in seedling survival. We found a microsite effect on seedling growth but this effect differed depending on the abiotic environment, with seedlings growing better in open microsites in the less stressful site. The larger abiotic context (climate) played an important role in determining which microsite is the most favorable for seedling growth but not necessarily for the eventual survival of seedlings. Wild ungulates started to damage seedlings early in the growing season, whereas abiotic stress (desiccation) affected mostly those plants that were not previously attacked by animals, showing the hierarchical effect (earlier and longer effect) of the biotic over the abiotic mortality agents. Non-preferred shrubs worked better as nurse shrubs only under high ungulate pressure. From the abiotic perspective, preferred and non-preferred shrubs did not apparently differ in the facilitation effect. Importantly, we found a greater facilitative effect of shrubs on the survival than on the growth of seedlings.