A forest gap is not forever: Towards an objective standard to determine when a gap is considered closed in temperate forests

Abstract

Forest gaps create environmental heterogeneity and drive forest dynamics through gap-phase regeneration. However, it is unclear when a forest gap can be considered closed, limiting a global synthesis of forest succession that integrates gap patterns and dynamics. We created twelve forest gaps (six large gaps, RD/H [ratio of gap diameter to gap border tree height] = 1.3; six small gaps, RD/H = 0.6) and monitored tree regeneration for seventeen years to establish an objective gap-closure standard for regeneration filling in a temperate secondary forest. Our results indicated that a forest gap could be considered closed or entering a gap-closure phase when the dominant gap regeneration layer (i.e., the top 10% gap-fillers) (1) reached the crown base height (8.9 m on average) of canopy trees and (2) entered a period of stability in density, richness, and evenness. Linear mixed-effects models found that these two conditions met each other in the same period (14.5 years) in the large gaps, with a mean density, richness, and evenness of 0.1 (stem m−2), 4.9 (species gap−1), and 0.9, respectively. However, if no repeated gap disturbance occurred, small gaps were closed by lateral extension rather than by height growth. Light-demanding species (e.g., Juglans mandshurica) mainly contributed to the closure of large gaps and played negligible roles in the small gaps, but intermediate or shade-tolerant species (e.g., Acer mono) in the small gaps had potential for successful gap filling following repeated gap disturbance. Our gap-closure standard links gap structure to species composition and provides a reference for large-scale research on gap patterns and dynamics, which helps to understand the ecological process during forest succession. The standard also provides a relatively clear boundary between forest gaps and closed forest stands, which is a premise to develop appropriate forest management and shorten the restoration period of degraded secondary forests.