Disturbances and Climate Drive Structure, Stability, and Growth in Mixed Temperate Old-growth Rainforests in the Caucasus

The Colchic rainforest of the Western Caucasus is one of the few temperate rainforests dominated by broadleaf deciduous trees. Understanding natural dynamics of broadleaf-dominated temperate rainforests is essential for their conservation and management. Here, we investigate for the first time the structure, natural disturbance, and recruitment dynamics of a mixed Colchic old-growth rainforest, dominated by Fagus orientalis and Picea orientalis . We used forest inventories and dendrochronological analysis of tree growth in five 30-m-radius plots to quantify forest structure, growth, and disturbances. For the last 400 years, the forest experienced a mixed disturbance regime dominated by frequent small gaps superimposed onto medium disturbances with about a 25-year recurrence period, with no evidences of stand-replacing disturbances. This disturbance regime favored the dominance of shade-tolerant, late successional species with slow tree canopy access through multiple growth releases. These dynamics impose low growth rates and continuous recruitment of spruce and beech, and contributed to a high heterogeneity of tree ages and sizes that result in stable forest structure, as suggested by the low stand slenderness. Spruces were the oldest (up to 427 years) and fastest growing trees in the forest, suggesting that their low presence in the forest is due to low disturbance rates that limit their recruitment. Spring climate conditions that promoted beech growth were detrimental for spruce growth, suggesting that interspecies interactions may condition the effect of climate on forest growth and development. The dynamic equilibrium state we reconstructed in this old-growth forest could likely be disrupted by anthropogenic disturbances or management.