Post-fire Regeneration Traits of Understorey Shrub Species Modulate Successional Responses to High Severity Fire in Mediterranean Pine Forests

Recurrent fires can impede the spontaneous recruitment capacity of pine forests. Empirical studies have suggested that this can lead to a prolonged replacement of pine forest by shrubland, especially if shrub species are pyrophytic. Model-based studies, however, have suggested that post-fire succession of pine forest under current climatic conditions will eventually tend towards the dominance of oaks under high fire severity and recurrence. These previous modelling studies did not address the role of the various post-fire regeneration traits of the understory shrub species. Considering the dichotomy of obligate seeder vs. resprouter species, either obligate or facultative resprouter, we hypothesized that when the shrubs present are post-fire seeders, the oaks steadily occupy the forest, whereas resprouter shrub species might compete with oaks and delay or arrest post-fire succession. To test this hypothesis, we developed a dynamic, cellular automaton model for simulating post-fire successional transitions in pine forests, including shrubs, pines and oaks, and stochastic fires of regular frequency. Our results showed a strong tendency towards oak dominance as final model state and a very reduced role of fire recurrence in this final state, with low yearly acorn input delaying oak dominance. Most relevantly, and in line with our hypothesis, the trend towards oak dominance depended markedly on the two types of shrub species, being delayed by resprouter species, which extended the shrub-dominated succession stage for several centuries. Our simulation results supported the view that the type of understorey species should be a key consideration in post-fire restoration strategies aiming to enhance fire resilience.