Cluster planting facilitates survival but not growth in early development of restored tropical forest

Abstract

Interspecific interactions are considered to be important structuring forces in early successional vegetation. Whereas competition seems to prevail in less severe environments, facilitation tends to increase in importance in harsh environments. Hence, facilitation is expected to play an important role in degraded tropical areas with high irradiance, heat loads, and evapotranspiration, where conditions are far from optimum for most forest species. We planned a restoration project on the southeastern Brazilian coastal plain to compare growth and survival of tree seedlings planted at two densities (isolated or aggregated) in a factorial experiment with nutrient addition. We monitored survival, height, ground level diameter, and crown projection of 4132 saplings from 19 species that ranged along a successional gradient, over an 18-month period. We used mixed-effect models to analyze the relationship between species performance and treatments, and Akaike's information criterion (AIC) to select the models. The best model showed higher survival in aggregated plantations (indicating facilitation) for non-pioneer (slower growing) species. In contrast, we found lower diameter growth in aggregated plantation (indicating competition). Fertilizer addition did not affect survival in clusters, but, surprisingly, it had a negative effect on isolated plants of both pioneer and non-pioneer species. Fertilizer addition had a positive effect on diameter and crown projection growth in aggregated plantations (reducing competition), especially for pioneer species. Thus, whether facilitation or competition was the predominant interaction depended on the effect analyzed. As establishment of non-pioneer species in disturbed sites can be challenging, restoration designs could take advantage of higher survival rates in clusters and use resource addition to ameliorate growth and decrease competition for limited resources.