Pathways and drivers of canopy accession across primary temperate forests of Europe.

Canopy accession strategies reveal much about tree life histories and forest stand dynamics. However, the protracted nature of ascending to the canopy makes direct observation challenging. We use a reconstructive approach based on an extensive tree ring database to study the variability of canopy accession patterns of dominant tree species (Abies alba, Acer pseudoplatanus, Fagus sylvatica, Picea abies) in temperate mountain forests of Europe and elucidate how disturbance histories, climate, and topography affect canopy accession. All four species exhibited high variability of radial growth histories leading to canopy accession and indicated varying levels of shade tolerance. Individuals of all four species survived at least 100 years of initial suppression. Fir and particularly beech, however, survived longer periods of initial suppression, exhibited more release events, and reached the canopy later on average, with a larger share of trees accessing the canopy after initially suppressed growth. These results indicate the superior shade tolerance of beech and fir compared to spruce and maple. The two less shade-tolerant species conversely relied on faster growth rates, revealing their competitive advantage in non-suppressed conditions. Additionally, spruce from higher-elevation spruce-dominated forests survived shorter periods of initial shading and exhibited fewer releases, with a larger share of trees reaching the canopy after open canopy recruitment (i.e. in absence of suppression) and no subsequent releases compared to spruce growing in lower-elevation mixed forests. Finally, disturbance factors were identified as the primary driver of canopy accession, whereby disturbances accelerate canopy accession and consequently regulate competitive interactions. Intensifying disturbance regimes could thus promote shifts in species composition, particularly in favour of faster-growing, more light-demanding species.

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Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.