Seasonal variations in the mechanisms of understory herb diversity in a temperate forest in Northeast China

Herbaceous plants contribute greatly to plant diversity and play an important role in regulating the structure and functioning of forest ecosystems. Comparing with woody species, herbs exhibit higher sensitivity to seasonal changes from spring-growth to autumn-mortality. Despite extensive research on understory herb diversity (e.g., abundance and richness) in forests, how the underlying mechanisms change across seasons remain unclear, especially in temperate forests. We explored the seasonal dynamics of herb diversity (here abundance and richness) by surveying herbs of 174 quadrats (1 m × 1 m) in a 25-ha (500 m × 500 m) temperate forest in spring, summer, and autumn of 2022, respectively. We examined the effects of overstory trees (diversity, composition, and three-dimensional (3D) structural complexity detected by LiDAR), topography (elevation, slope, and aspect), and microsite conditions (light availability, soil nutrients, and soil water content) on herb diversity in different seasons. We found a similar seasonal pattern for herb abundance and richness, i.e., both decreased from spring to summer and then to autumn, but the driving mechanisms differed remarkably across seasons. Specifically, the importance of overstory trees, topography, and microsite conditions on herb abundance vs. richness varied inversely across seasons. For herb abundance, the importance of overstory trees (primarily 3D structural complexity) increased from spring to autumn, while the importance of topography decreased. Conversely, for herb richness, the importance of overstory trees decreased from spring to autumn, while the importance of topography increased. Microsite conditions governed spatial variations of herb abundance in all seasons but only in autumn for herb richness, with importance of soil nutrients remaining relatively constant while that of soil water content and light availability changed greatly across seasons. Our results demonstrate substantial seasonal variations in driving mechanisms of herb diversity in the temperate forest. These differentiated responses of herb abundance and richness to different drivers across seasons further highlight the significance of considering seasonal variations of the mechanisms underlying understory plant diversity in forest biodiversity conservation.