The strong seasonality of soil microbial community structure in declining Mediterranean pine forests depends more on soil conditions than on tree vitality

The soil microbiome plays an important role in forest functioning. However, the impact of drought-induced dieback and tree death on soil microbial biomass, community structure, and functional composition is unknown. We also lack understanding on how soil microbiota varies seasonally in such declining stands. We used Phospholipid Fatty Acids (PLFA) analysis to quantify soil microbial biomass and study its seasonal changes in three Mediterranean forests showing dieback and dominated by three pine species (Pinus halepensis Mill, Pinus pinaster Ait. and Pinus sylvestris L.). We also measured microclimatic parameters and soil physical and chemical parameters under trees with different vigor, assessed as canopy defoliation degree, and related them to seasonal changes in the soil microbial community. We found marked differences in soil microbial community structure, total biomass, and the relative abundance of major functional groups among forests. First, soil microbial biomass peaked either in the dry summer (P. halepensis) or in autumn (P. pinaster and P. sylvestris). Accordingly, the relative abundance of most functional groups, excluding Arbuscular mycorrhizal fungi, displayed substantial variation between seasons. In addition, the relative abundance of fungi and Gram-positive bacteria exhibited an opposite pattern compared to actinomycetes and Gram-negative bacteria. Second, soil physical and chemical parameters had a significant impact on within-site PLFA variation, although their influence was less important than that of seasonal variation. Third, differences between defoliated and healthy trees were minor and restricted to averaged ratios between different PLFA markers. Overall, the structure, biomass and relative abundance of major functional groups of the soil microbiome vary considerably among stand types and seasons in forests showing ongoing dieback and high mortality. However, while the seasonal dynamics show predictable patterns, which should be accounted for in future studies, the within-site variation is highly variable and mainly depends on soil physical and chemical parameters.