Effects of tree canopy on herbaceous understorey throughout the developmental cycle of a temperate mountain primary forest

Temporal dynamics of temperate montane primary forests were described based on data from Central-European nature reserves in the 20th century as a long-term cycle involving tree regeneration, growth, maturation, and disintegration. However, the lack of chronosequences focusing on the herbaceous understory has limited our understanding of plant community dynamics within this developmental cycle. To address questions regarding whether the herbaceous understory follows the cyclic trajectory of the tree layer and if the properties of the tree layer drive the dynamics of the herbaceous understory, we investigated a 19-year chronosequence in ten plots (with five sampling periods) within an unmanaged primary beech-fir forest in the Western Carpathians. The vegetation data from the resampled plots, including herb species frequencies and dendrometric variables, covered a significant portion of the developmental cycle of a single plant community type. The results confirmed the cyclical nature of herbaceous understory development in terms of species composition and diversity trajectories, demonstrating transitions between developmental stages. Diversity was highest during the disintegration stage, with a rapid decline towards the growth stage, followed by a slow recovery as it progressed towards the optimum stage. Temporal responses of individual species, along with ordination analyses, allowed for the separation of taxa associated with the disintegration, growth, and optimum stages, as well as the most stable ones throughout the entire cycle. The proportion of therophytes slightly increased after gap-opening events during disintegration, geophytes were most represented during the growth stage, and hemicryptophytes culminated during the optimum stage. The composition of herb species was significantly affected by the above-ground tree biomass (AGTB), which explained 21.4% of the temporal variability. The sum of the diameter at breast height (Sum of DBH) described an additional 7.2%. The relationship between the frequency of the herbaceous understory and AGTB followed a "U" shape, while the relationship with Sum of DBH was negative and linear. The relationship between the herbaceous understory and the tree overstory is likely driven by competition for light, water, and nutrient resources. These findings provide new insights into the temporal relationships between the herbaceous understory and tree layer development, and they may inspire future studies and management strategies aimed at conserving or enhancing biodiversity in temperate forests.