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Structural diversity promotes productivity of mixed, uneven-aged forests in southwestern Germany

Abstract

Forest diversity-productivity relationships have been intensively investigated in recent decades. However, few studies have considered the interplay between species and structural diversity in driving productivity. We analyzed these factors using data from 52 permanent plots in southwestern Germany with more than 53,000 repeated tree measurements. We used basal area increment as a proxy for productivity and hypothesized that: (1) structural diversity would increase tree and stand productivity, (2) diversity-productivity relationships would be weaker for species diversity than for structural diversity, and (3) species diversity would also indirectly impact stand productivity via changes in size structure. We measured diversity using distance-independent indices. We fitted separate linear mixed-effects models for fir, spruce and beech at the tree level, whereas at the stand level we pooled all available data. We tested our third hypothesis using structural equation modeling. Structural and species diversity acted as direct and independent drivers of stand productivity, with structural diversity being a slightly better predictor. Structural diversity, but not species diversity, had a significant, albeit asymmetric, effect on tree productivity. The functioning of structurally diverse, mixed forests is influenced by both structural and species diversity. These sources of trait diversity contribute to increased vertical stratification and crown plasticity, which in turn diminish competitive interferences and lead to more densely packed canopies per unit area. Our research highlights the positive effects of species diversity and structural diversity on forest productivity and ecosystem dynamics.

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Acknowledgments

We thank our colleagues from the Forest Research Institute of Baden-Württemberg for their hard work and dedication in managing the permanent plots used for this analysis. We thank two anonymous reviewers for constructive comments on an earlier version of the manuscript and Stephanie Pollhammer for proofreading this article. This study was funded by the Ministry of the Environment, Climate Protection and the Energy Sector Baden-Württemberg (research program KLIMOPASS). Significant changes to the original manuscript and supplementary analyses were performed while A. D. was the holder of a doctoral grant from the German Federal Environmental Foundation.

Author contribution statement

A. T. A. originally formulated the idea; A. D. analyzed the data; A. D., J. B. and A. T. A. wrote the manuscript.

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Correspondence to Adrian Dănescu.

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Communicated by Daniel Laughlin.

Our study brings quantitative evidence that both structural and species diversity regulate forest ecosystem functioning. Both diversity aspects had a significant positive impact on forest productivity and ecosystem functioning. Our results indicate that structural diversity contributes to ecosystem functioning more strongly than previously known, and that adding diversity indices in growth models tends to reduce prediction errors. We discuss mechanisms which underpin our findings and highlight the importance of considering structural diversity in future studies.

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Dănescu, A., Albrecht, A.T. & Bauhus, J. Structural diversity promotes productivity of mixed, uneven-aged forests in southwestern Germany. Oecologia 182, 319–333 (2016). https://doi.org/10.1007/s00442-016-3623-4

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Keywords

  • Basal area increment
  • Complementarity
  • Ecosystem functioning
  • Biodiversity
  • Species diversity