Patterns and drivers of regeneration of tree species in forests of peninsular Spain

Aim Our study aimed to identify and explore the main factors that influence tree recruitment of multiple species at a regional scale across peninsular Spain, an understanding of which is essential for predicting future forest species composition in the face of ongoing environmental change. The study focused on the dynamics of the key transition phase from saplings to adult trees. Location The forests of peninsular Spain. Methods We used the extensive network of plots sampled in two consecutive Spanish national forest inventories (> 30,000 plots) to identify the factors that determine regeneration patterns of the 10 most abundant forest species of Spain at relatively large temporal (c. 10 years) and spatial scales (across Spain): five coniferous species of Pinus (pines) and five broadleaved species of the genera Fagus and Quercus. We fitted separate generalized linear models for the pine species and the broadleaved species to assess the response of sapling abundance and ingrowth rate to the spatial variability of climate (temperature, water availability and recent warming), forest structure (tree density, understorey and overstorey canopy cover, and basal area change) and disturbances (previous forest logging, wildfires and grazing). Results Mean sapling abundance was four times higher for broadleaved species than for pines, while mean annual ingrowth was twice as high. Sapling abundance and ingrowth rate were mainly determined by stand structure, both in pines and broadleaved trees. The direct effects of disturbances and climate were comparatively smaller, and there was no detectable effect of recent warming. Main conclusions The higher values of ingrowth rate of broadleaved species can be explained by their ability to maintain a higher sapling bank due to their greater shade tolerance. This differential response of pines and broadleaved species to canopy closure suggests a probable increase in broadleaved species at the expense of pines. This transition could occur earlier in stands with faster canopy closure dynamics. Spatially explicit, mixed-species demographic models incorporating both the ingrowth and the tree mortality components are needed for predicting the composition of future forests.