Understory seedlings of Quercus mongolica survive by phenological escape

Understanding understory seedling regeneration mechanisms is important for the sustainable development of temperate primary forests in the context of increasingly intense climate warming events. The poor regeneration of dominant tree species, however, is one of the biggest challenges it faces at the moment. Especially, the regeneration of the shade-intolerant Quercus mongolica seedling is difficult in primary forests, which contrasts with the extreme abundance of understory seedlings in secondary forests. The mechanism behind the interesting phenomenon is still unknown. This study used in-situ monitoring and nursery-controlled experiment to investigate the survival rate, growth performance, as well as nonstructural carbohydrate (NSC) concentrations and pools of various organ tissues of seedlings for two consecutive years, further analyze the understory light availability and simulate the foliage carbon (C) gain in the secondary and primary forest. Results suggested that seedlings in the secondary forest had greater biomass allocation aboveground, height and specific leaf area (SLA) in summer, which allowed the seedling to survive longer in the canopy closure period. High light availability and positive C gain in early spring and late autumn are key factors affecting the growth and survival of understory seedlings in the secondary forest, whereas seedlings in the primary forest had annual negative carbon gain. Through the growing season, the total NSC concentrations of seedlings gradually decreased, whereas those of seedlings in the secondary forest increased significantly in autumn, and were mainly stored in roots for winter consumption and the following year's summer shade period, which was verified by the nursery-controlled experiment that simulated autumn enhanced light availability improved seedling survival rate and NSC pools. In conclusion, our results revealed the survival trade-off strategies of Quercus mongolica seedlings and highlighted the necessity of high light availability during the spring and autumn phenological periods for shade-intolerant tree seedling recruitment.