Competition and climate influence in the basal area increment models for Mediterranean mixed forests

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Competition plays a key role controlling tree growth in mixed forests. Contrary to monocultures, quantifying species mixing influence on tree growth suppose a challenge since the presence of two or more species requires to estimate the degree of intra- and inter-specific competition among trees. Moreover, it is well known that aridity can also influence tree growth, especially in the Mediterranean Basin. In the present context of climate change, it is essential to take into account species mixing and aridity uncertainty in the design of sustainable management guidelines for Mediterranean mixed forests. To achieve that, data from Spanish National Forest Inventory was used in this study to fit new mixed-effects basal area increment (BAI) models for 29 two-species compositions in Spain. A wide range of different competition structures (intra-specific, inter-specific, size-symmetric and size-asymmetric) and aridity conditions (in terms of the De Martonne Index) were included and tested into the BAI models. Parameter estimations were obtained for all possible species, mixtures and combinations by Maximum Likelihood (ML). Models with all the coefficients being significant (p < 0.05) were first selected. Among these models, we used Akaike Evidence Ratios for selecting the best one by species for each mixture. The best model for each species and mixture was used to analyze the competition and climatic influence on tree growth. Regarding competition influence, a common trend among mixtures was found with higher productivity in mixed than pure stands, suggesting that BAI values may increase with the increment of species diversity. Based on intra and inter-specific competition indexes, competition seemed to be the most representative biological interaction in conifer-conifer mixtures, since neutralism and facilitation may occur more frequently in conifer-broadleaved and broadleaved-broadleaved mixtures. Our findings also suggested that tree growth may be significantly limited by arid conditions, excepting for Pinus halepensis and Pinus pinea. Our rigorous modelling approach successfully uncovered not only possible mixing effect among various species but also help us to understand the effect of aridity on tree growth. Thus, models presented in this study can be used in the design and implementation of management and adaptation guidelines under future climate change scenarios.

The authors would like to thank the Spanish Ministry of Economy and Competitiveness for funding this research through Industrial PhD project [grant DI-15-07722] and the Torres Quevedo programme [grant PTQ-12-05409]. The authors are also grateful to the Ministry for Ecological Transition and the WorldClim team for sharing and providing the data used in this study.

Peer reviewed