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The role of tree diversity for individual tree growth, crown architecture and branch demography
(2012)
In the light of the concurrent loss of biodiversity, biodiversity and ecosystem functioning (BEF) research attracted a great deal of attention and emerged as one of the important fields of research in ecology. Since important ecological interactions such as competition occur between individuals, the understanding of individual tree growth was considered to be fundamental for forest related BEF research. Individual tree growth is determined by the above- and belowground interactions of a tree individual with its local neighbourhood. To obtain a deeper understanding of BEF relationships, I broadened the focus from individual tree growth (usually measured as diameter or biomass increment) to the arrangement and dynamics of the above-ground modules of trees in dependence of their local neighbourhood. More precisely, the main objective of the present thesis was to analyse the impact of tree diversity on individual tree growth, crown architectural and branch demographic variables. Thereby I considered crown architectural variables as important indicators of the competition for light. In addition, crown architectural variables impacted ecosystem services such as erosion control. Furthermore, the results of the present thesis contributed to the current discussion on species coexistence theories, which may be differentiated by two opposing views: one that relies on neutral processes and one that implicates a role for meaningful differences in the ecological strategy (niche) of co-occurring species. The studied forest ecosystems were the subtropical broad-leaved evergreen forests of southeast China, which have been under high human pressure due to a long history of intensive land-use. The area is of particular interest for BEF research due to the high species richness of woody plants, including many, yet poorly studied species, and due to the rough terrain with steep slopes, which cause severe soil erosion. The present thesis combines three observational with two experimental studies, applying the local neighbourhood approach along an age gradient from tree saplings to mature trees. In the Gutianshan National Nature Reserve (GNNR), I conducted two observational studies on permanent plots which were chosen according to a space-for-time substitution design. The aim of the first study was to reveal the effects of diversity (species richness, functional diversity) together with other biotic and abiotic variables on morphological growth parameters (crown area, crown displacement and stem inclination) of target trees of four tree species (Castanea henryi, Castanopsis eyrei, Quercus serrata and Schima superba). In the second study, the same target trees together with their neighbours were used to analyse the relation between stand related functional diversity and the horizontal and vertical structure of the canopy. The third study was conducted in a young secondary broad-leaved evergreen forest. Using two target species (Castanopsis fargesii and Quercus fabri), the role of diversity, intra- vs. inter-specific competition and the mode of competition (symmetric vs. asymmetric) on the target individuals was tested by analysing five-year radial growth increments. The two other studies were carried out in an experimentally established plantation, using saplings of four tree species (C. henryi, Elaeocarpus decipiens, Q. serrata and S. superba), which were planted in monoculture, twoand four-species combinations and in three densities. The fourth study focused on mechanisms of coexistence and the role of species richness, species composition, species identity and density on sapling growth. The fifth study tested the effect of sapling density and identity on the througfall kinetic energy, which represents a measure for the erosive power of rain. It was found that functional diversity does affect crown architectural and canopy related parameters of forests in the GNNR. However, no effects of species richness on radial-growth were detected in the younger forest. Since I also did not find strong effects of species richness on saplings in the experimental plantation, diversity effects may evolve at a later age stage. The importance of the diversity effect may be related reversely to that of species identity in an age gradient of forest stands. The findings suggest that different mechanisms of coexistence operate simultaneously but that their relative importance may shift through the life stages of trees. During the sapling stage, species-specific differences in growth and architectural traits support niche theory. In older forest stands, no species-specific differences in growth parameters could be detected. However, I did find effects of functional diversity on horizontal canopy structure. I conclude that mechanisms of coexistence may not only change with forest stand age, but may also differ for distinct traits. The present thesis, being the first to apply the local neighbourhood approach with regard to crown architecture and branch demography within the BEF field of research, stresses the importance of this individual based approach. Although the observed forest systems are very complex, crown architectural and canopy structural variables were found to be affected by diversity. The finding that the degree of erosive power of rain could be elucidated by crown architectural variables, encourages further studies to reveal possible relations between biodiversity and other ecosystem functions or services, which might be mediated by crown architectural and canopy structural variables.