Disentangling regeneration by vertical stratification: A 17-year gap-filling process in a temperate secondary forest

Forest gaps are widely spread in the forest ecosystems and play important roles in driving forest dynamics. However, long-term regeneration during the gap-filling process was less understood, especially species compo-sition and dynamics along vertical gradients over time. We monitored tree regeneration in artificial gaps of a temperate secondary forest for 17 years and analyzed the gap-filling process both as a whole and by stratification (lower layer: 0-3 m, middle layer: 3-6 m, and upper layer: >6 m) for three shade-tolerance groups of tree species. We found that the interaction effects of gap size and shade tolerance could result in a growth difference of five to ten years among different regeneration groups, with the most significant effects in the large gaps for the light-demanding species such as Juglans mandshurica and Phellodendron amurense. Regeneration density in the large and small gaps eventually converged, but there were great differences in density dynamics. The regener-ation density in the large gaps decreased gradually over time, while it decreased sharply in the small gaps after just one growing season of gap formation and consistently remained at low levels regardless of shade tolerance. Species richness decreased smoothly during the 17-year monitoring, but the richness in the large gaps was consistently higher than that in the small gaps, with all shade-tolerance groups existing, which underlined the importance of gap size in maintaining tree species diversity. Differences in the density, richness, and composition among regeneration layers reduced over time. However, vertical regeneration patterns diverged among shade -tolerance groups during the gap-filling process. Light-demanding species were likely to have only one chance for successful regeneration because almost no new recruitments appeared in the lower or middle layer as dominant stems reached the upper layer. Intermediate and shade-tolerant species, by contrast, were well -distributed in different regeneration layers, indicating a stronger capacity for resistance and resilience during uncertain disturbance events in the long run. Our results revealed the gap-filling strategies of tree species with different shade-tolerance groups, which explained why light-demanding species could not completely occupy a forest gap and highlighted the necessity of repeated gap disturbance for intermediate and shade-tolerant species regeneration.