Patterns of Soil Stoichiometry Driven by Mixed Tree Species Proportions in Boreal Forest

Soil stoichiometry is essential for determining the ecological functioning of terrestrial ecosystems. Understanding the stoichiometric relationships in mixed forests could enhance our knowledge of nutrient cycling. In a mixed forest zone of Larix principis-rupprechtii (LP) and Betula Platyphylla (BP) in Hebei China, we conducted a study at six different sites with varying levels of tree species mixing. The proportion of L. principis-rupprechtii ranged from 0% to 100%, with intermediate values of 8.58%, 10.44%, 18.62%, and 38.32%. We compared soil stoichiometry, including carbon (C), nitrogen (N), and phosphorus (P), as well as chemical and physical properties across these sites. Piecewise structural equation modeling (piecewiseSEM) was used to assess the direct and indirect links between key ecosystem factors and their effects on soil stoichiometry. In mixed forests, the soil exhibited higher contents of soil organic matter (SOM), total nitrogen (TN), and total phosphorus (TP) compared to those in pure LP forests. Additionally, the soil C: N ratio in the 8.58% and 18.62% mixed forests as well as pure BP forests was significantly higher than that in LP forests. Structural equation modeling (SEM) revealed that the contents and ratios of soil C, N, and P exhibited different responses to mixed species proportions. The effect of mixed species proportions on soil nutrients was predominantly indirect, mediated primarily by variations in soil-available nutrients and, to a lesser extent, by physical properties and pH. Specifically, an increase in the proportion of LP in mixed forests had a direct negative effect on soil-available nutrients, which in turn had a positive effect on the content of SOM, TN, and TP and their respective ratios. Based on these findings, we can predict that soil nutrient limitation becomes more pronounced with increasing proportions of Larix principis-rupprechtii in the mixed forest. Our results emphasized the significance of changes in mixed species proportions on soil stoichiometry, providing valuable references for the sustainable development of forests.