Characterization of the structure, dynamics, and productivity of mixed-species stands: review and perspectives

The growth and yield of mixed-species stands has become an important topic of research since there are certain advantages of this type of forest as regards functions and services. However, the concepts and methods used to characterize mixed stands need to be understood, as well as harmonized and standardized. In this review we have compiled a set of measures, indices, and methods at stand level to characterize the structure, dynamics, and productivity of mixed stands, and we discuss the pros and cons of their application in growth and yield studies. Parameters for the characterization of mixed stand structure such as stand density, species composition, horizontal (intermingling) and vertical tree distribution pattern, tree size distribution, and age composition are described, detailing the potential as well as the constraints of these parameters for understanding resource capture, use, and efficiency in mixed stands. Furthermore, a set of stand-level parameters was evaluated to characterize the dynamics of mixed stands, e.g. height growth and space partitioning, self- and alien-thinning, and growth partitioning among trees. The deviations and changes in the behaviour of the analysed parameters in comparison with pure stand conditions due to inter-specific interactions are of particular interest. As regards stand productivity, we reviewed site productivity indices, the growth–density relationship in mixed stands as well as methods to compare productivity in mixed versus monospecific stands. Finally, we discuss the main problems associated with the methodology such as up-scaling from tree to stand level as well as the relevance of standardized measures and methods for improving forest growth and yield research in mixed stands. The main challenges are also outlined, especially the need for qualitatively sound data.