Abstract
As they fulfil many ecological and social functions and services better than even-aged monocultures, heterogeneous pure and mixed-species stands are on the advance in Central Europe. Even so, knowledge of how different stands compare in terms of the quantity and quality of the produced wood remains limited, as forest research has been focused on pure stands in the past. Therefore, the still limited comparative studies on timber quality in mixed versus pure stands were reviewed. Further, approximately 100 studies on the morphology of mixed versus pure stands have been reviewed. As is known, the close connection between morphology and timber quality from many studies in pure stands as well as the morphological and structural properties of trees in mixed stands is used as proxies for their timber quality. The number of studies reporting a decrease or increase in timber strength and stiffness in complex stands compared with homogeneous stands was balanced. Knottiness is mostly higher in complex stands. Wood density behaves indifferently. Distortion, as indicated by eccentricity of crown, bending of stems, or irregularity of the tree-ring width, is generally higher in complex forests. This rather ambiguous pattern becomes clearer by typifying the findings depending on the species-specific morphological plasticity of the trees and the spatial conditions they are exposed to. When growing in strong lateral restriction in even-aged pure or mixed-species stands (type 1), trees follow a “keep abreast” strategy which results in high-quality timber especially in case of species with low plasticity. Trees in uneven-aged forests with vertically restricted growing space (type 2) often use a “sit-and-wait” strategy that may result in tapering stem shapes, wide and long crowns with low branch diameters, and high wood density. Distortion may be low in case of species with low morphological plasticity but increase with increasing shade tolerance and plasticity. Growth in widely spaced and heavily thinned pure and mixed stands (type 3) may let trees follow the “stabilisation” strategy. Because of their strong dominance, these trees develop tapering stem shapes, knots of big size and wide appearance along the stem axis, as well as lower wood density, especially in the case of conifers. In arrangements of types 1–3, the “transition” strategy may also emerge, which leads from the “sit-and-wait” stadium to the “keep abreast” strategy. It starts when trees strongly increase their height growth at the expense of the stem diameter growth. It results in slender stems, low knottiness, high wood density, and low distortion, with the result that the tree gets access to the upper canopy at the expense of lateral expansion of stem and crown. In fact, it is not primarily the species mixing that modifies the morphology, structure, and wood quality of the trees but the species-specific morphological plasticity and the structural heterogeneity of the stand. The latter is often higher in mixed than in pure stands and in uneven-aged than in even-aged stands. The more variable the stand structure, the wider the range of wood attributes. The discussion is focused on the relevance of the results for stand management and interdisciplinary research at the intersection of forest growth and yield science and wood science.
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Acknowledgments
We thank the Bavarian State Ministry for Nutrition, Agriculture, and Forestry for its support of project X36, “Relationship between spacing and wood quality of Douglas-fir in Bavaria”, and the German Science Foundation (Deutsche Forschungsgemeinschaft) for providing the funds for project PR 292/12-1, “Tree and stand-level growth reactions to drought in mixed versus pure forests of Norway spruce and European beech”. C. T. Bues, P. Comeau, F. Hapla, S. Hein, H. Spellmann, A. Mäkelä, A. Nepveu, R. Schneider, J. P. Schütz, T. Seifert, C. H. Ung, and A. Weiskittel supported this study by valuable suggestions, overview of literature, or supply of publications. Thanks are also due to Gerhard Schütze for supporting the statistical analysis, Ulrich Kern for the graphical artwork, and two anonymous reviewers for their constructive criticism.
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Pretzsch, H., Rais, A. Wood quality in complex forests versus even-aged monocultures: review and perspectives. Wood Sci Technol 50, 845–880 (2016). https://doi.org/10.1007/s00226-016-0827-z
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DOI: https://doi.org/10.1007/s00226-016-0827-z