Phylogenetic constraints to soil properties determine elevational diversity gradients of forest understory vegetation

Elevational diversity gradients (EDGs) of vegetation are shaped by the evolutionary histories of plants as well as by ecological factors. However, few studies of EDGs have focused on the roles of phylogenetic constraints and the effects of complicated interactions among environmental factors. Here, we examine the direct and indirect effects of environmental factors in forming EDGs of forest understory vegetation. The study plots were selected along elevational gradients in cool-temperate and sub-alpine forests in the University of Tokyo Chichibu Forest of central Japan. Tree seedlings and herbs were identified, and environmental factors (elevation, soil temperature, soil pH, soil CN ratio, forest type, basal area, canopy openness, and slope) were measured in these plots. Structural equation modeling (SEM) including taxonomic and phylogenetic diversity was used to consider the causal relationships between environmental conditions and the diversity of understory vegetation. In addition, phylogenetic signals in habitat requirements were tested. The taxonomic and phylogenetic diversities of tree seedlings increased monotonically with elevation, and the same pattern was found for the taxonomic diversity of herbs. The SEM indicated that both the taxonomic and phylogenetic diversity of tree seedlings were most affected by soil properties, although the phylogenetic diversity of herbs was determined by light conditions. These results highlight the importance of environmental filtering by soil properties in shaping EDGs of tree seedlings. This study implies that phylogenetic constraints in the adaptation to soil properties should be considered when predicting changes in EDGs under environmental fluctuations.