Moderate pine cover maximizes 10-year survival and growth in late-successional species of contrasting functional strategies

Abstract

Monospecific pine forests are widespread due to extensive afforestation efforts and natural colonization of abandoned croplands in the Mediterranean Basin. It was originally thought that pines would facilitate the natural colonization of native late-successional resprouter species (e.g., hardwoods), but these species can be compromised if competition with pines outweighs their facilitative effect on these hardwood species. Managing the density or canopy cover of these widespread pine forests can potentially provide some “optimum” balance between facilitation and competition to maximize success in the introduction of late successional species while maintaining a tree stratum. We tested the response (survival and growth across 10 years) of six resprouter species covering a wide range of plant functional strategies, from drought-tolerant sclerophyllous shrubs and trees to malacophyllous drought-sensitive trees, across an experimental gradient of Aleppo pine canopy cover. Seedling performance varied according to the functional strategy, pine cover and time. High pine cover generally enhanced seedling survival, whereas moderate pine cover generally enhanced seedling growth, although this response was modulated by the functional strategy of the seedling species. Interactions between pines and seedlings were only detectable 2–3 years after plantation, increasing in intensity with time. The latter highlights the need of medium to long-term studies to evaluate plant-plant interactions in these water-limited environments with slow successional trajectories. Our results could be attributed to the shade tolerance of most of the introduced trees, combined with their low tolerance to the combination of high sunlight radiation and drought. We found an optimal pine cover of ca. 50% (equivalent to 300–400 trees/ha) in which both survival and growth of late successional species can be maximized, which help to select best locations for more efficient reforestation programs and set a threshold value to decide whether or not to perform tree thinning to enhance ecosystem diversity and, subsequently, resilience.