Spatial and temporal regeneration patterns within gaps in the primary forests vs. secondary forests of Northeast China

Abstract

Forest gaps play an important role during forest succession in temperate forest ecosystems. However, the differences in spatial distribution and replacement patterns of woody plants (trees and shrubs) between primary and secondary forests remain unclear during the gap-filling processes, especially for temperate forests in Northeast China. We recorded 45,619 regenerated trees and shrubs in young gaps (<10 years), old gaps (10~20 years), and closed forest stands (i.e., filled gaps) in the primary broadleaved Korean pine (Pinus koraiensis Sieb. Rt Zucc.) forests vs. secondary forests (degraded from primary forests). The gap-filling processes along horizontal (Cartesian coordinate system) and vertical (lower layer: 0~5 m, medium layer: 5~10 m, and upper layer: >10 m) dimensions were quantified by shade tolerance groups of trees and shrubs. We found that gap age, competition between species, and pre-existing regeneration status resulted in different species replacement patterns within gaps in primary vs. secondary forests. Gap formation in both primary and secondary forests increased species richness, with 33, 38, 39, and 41 in the primary closed stands, primary forest gaps, secondary closed stands, and secondary forest gaps, respectively. However, only 35.9% of species in primary forest gaps and 34.1% in secondary forest gaps successfully reached the upper layer. Based on the importance values (IVs) of tree species across different canopy heights, light-demanding trees in the upper layer of the secondary forests were gradually replaced by intermediate and shade-tolerant trees. In the primary forests, Korean pine exhibited intermittent growth patterns at different canopy heights, while it had continuous regeneration along vertical height gradients in the secondary forests. The differences in Korean pine regeneration between the primary and secondary forests existed before gap formation and continued during the gap-filling processes. The interspecific competition among different tree species gradually decreased with increasing vertical height, and compared to the primary forests, the secondary forests showed an earlier occurrence of competition exclusion within gaps. Our findings revealed the species replacement patterns within gaps and provided a further understanding of the competition dynamics among tree species during the gap-filling processes.