Plant-soil feedbacks in declining forests: implications for species coexistence.

Plant-soil feedbacks (PSFs) play a relevant role as drivers of species abundance, coexistence, and succession in plant communities. However, the potential contribution of PSFs to community dynamics in changing forest ecosystems affected by global change drivers is still largely unexplored. We measured the direction, strength and nature (biological vs. chemical) of PSFs experienced by coexisting tree species in two types of declining Quercus suber forests of southwestern Spain (open woodland vs. closed forest) invaded by the exotic soil pathogen Phytophthora cinnamomi. To test PSFs in a realistic community context, we focused not only on individual PSFs (i.e., comparing the growth of a tree species on conspecific vs. heterospecific soil) but also calculated net-pairwise PSFs by comparing performance of coexisting tree species on their own and each other's soils. We hypothesized that the decline and death of Q. suber would alter the direction and strength of individual and net-pairwise PSFs due to the associated changes in soil nutrients and microbial communities, with implications for recruitment dynamics and species coexistence. In support of our hypothesis, we found that the decline of Q. suber translated into substantial alterations of individual and net-pairwise PSFs, which shifted from mostly neutral to significantly positive or negative, depending on the forest type. In both cases however the identified PSFs benefited other species more than Q. suber (i.e., heterospecific positive PSF in the open woodland, conspecific negative PSF in the closed forest). Our results supported PSFs driven by changes in chemical soil properties (mainly phosphorus) and arbuscular mycorrhizal fungi, but not in pathogen abundance. Overall, our study suggests that PSFs might reinforce the loss of dominance of Q. suber in declining forests invaded by P. cinnamomi by promoting the relative performance of non-declining coexisting species. More generally, our results indicate an increase in the strength of net PSFs as natural forests become disturbed by global change drivers (e.g., invasive species), suggesting an increasingly important role of PSFs in forest community dynamics in the near future.