Regeneration Dynamics Following the Formation of Understory Gaps in a Slovakian Beech Virgin Forest

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The frequency and size of canopy gaps largely determine light transmission to lower canopy strata, controlling structuring processes in the understory. However, quantitative data from temperate virgin forests on the structure of regeneration in gaps and its dynamics over time are scarce. We studied the structure and height growth of tree regeneration by means of sapling density, shoot length growth and cumulative biomass in 17 understory gaps (29 to 931 m2 in size) in a Slovakian beech (Fagus sylvatica L.) virgin forest, and compared the gaps with the regeneration under closed-canopy conditions. Spatial differences in regeneration structure and growth rate within a gap and in the gap periphery were analyzed for their dependence on the relative intensities of direct and diffuse radiation (high vs. low). We tested the hypotheses that (i) the density and cumulative biomass of saplings are higher in gaps than in closed-canopy patches, (ii) the position in a gap influences the density and height growth of saplings, and (iii) height growth of saplings increases with gap size. Sapling density and biomass were significantly higher in understory gaps than under closed canopy. Density of saplings was positively affected by comparatively high direct, but low diffuse radiation, resulting in pronounced spatial differences. In contrast, sapling shoot length growth was positively affected by higher levels of diffuse radiation and also depended on sapling size, while direct radiation intensity was not influential. Conclusively, in this forest, regeneration likely becomes suppressed after a short period by lateral canopy expansion in small gaps (<100 m2), resulting in a heterogeneous understory structure. In larger gaps (≥100 m2) saplings may be capable even at low plant densities to fill the gap, often forming a cohort-like regeneration layer. Thus, gaps of different sizes imprint on the resulting canopy structure in different ways, enhancing spatial heterogeneity.