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Tree-tree interactions and crown complementarity: The role of functional diversity and branch traits for canopy packing
Institution:1. Institute of General Ecology and Environmental Protection, Technische Universität, Pienner Straße 7, Dresden, Tharandt 01737, Germany;2. Institute of Biology/Geobotany and Botanical Garden, Martin Luther University Halle-Wittenberg, Große Steinstr. 79/80, Halle Saale 06108, Germany;3. German Centre for Integrative Biodiversity Research (iDiv), Halle-Jena-Leipzig, Leipzig 04103, Germany;4. Institute of Ecology, Leuphana University of Lüneburg, Universitätsallee 1, Lüneburg 21335, Germany;1. German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, Leipzig, Germany;2. Institute of Biology, Leipzig University, Leipzig, Germany;3. Institute of Ecology and Evolution, Friedrich Schiller University Jena, Jena, Germany;4. EcoNetLab, Institute of Biodiversity, Friedrich Schiller University Jena, Jena, Germany;5. Institute of Biology/Geobotany and Botanical Garden, Martin Luther University Halle-Wittenberg, Halle (Saale), Germany;6. Institute of Agricultural Sciences, ETH Zürich, Zürich, Switzerland;7. UFZ—Helmholtz Centre for Environmental Research, Soil Ecology Department, Halle (Saale), Germany;8. Centre d''Ecologie Fonctionnelle et Evolutive, UMR 5175 (CNRS—Université de Montpellier—Université Paul-Valéry Montpellier—EPHE), Montpellier, France;9. Institute for Chemistry and Biology of Marine Environments [ICBM], Carl-von-Ossietzky University Oldenburg, Wilhelmshaven, Germany;10. Department of Ecology, Evolution and Behavior, University of Minnesota, St. Paul, MN, United States;11. Karlsruher Institut für Technologie (KIT), Institut für Geographie und Geoökologie, Karlsruhe, Germany;12. Institute of Computer Science, Friedrich Schiller Universität Jena, Jena, Germany;13. Radboud University, Institute for Water and Wetland Research, Animal Ecology and Physiology & Experimental Plant Ecology, Nijmegen, The Netherlands;14. Terrestrial Ecology Research Group, Technical University of Munich, School of Life Sciences Weihenstephan, Freising, Germany;15. Ecotron Européen de Montpellier, Centre National de la Recherche Scientifique (CNRS), Montferrier-sur-Lez, France;p. Field Station Fabrikschleichach, Department of Animal Ecology and Tropical Biology, Biocenter, University of Würzburg, Rauhenebrach, Germany;q. Bavarian Forest National Park, Grafenau, Germany;r. Geobotany, Faculty of Biology, University of Freiburg, Freiburg, Germany;s. Department of Renewable Resources, University of Alberta, Edmonton, AB, Canada;t. Department of Biosciences, University of Salzburg, Salzburg, Austria;u. UFZ—Helmholtz Centre for Environmental Research, Department Physiological Diversity, Leipzig, Germany;v. Institute of Landscape Ecology, University of Münster, Münster, Germany;w. Department of Geography, University of Zürich, Zürich, Switzerland;x. Department of Biology, Marquette University, Milwaukee, WI, United States;y. Forest Nature Conservation, Faculty of Forest Sciences and Forest Ecology, University of Göttingen, Göttingen, Germany;z. Department of Crop Sciences, Division of Agroecology, University of Göttingen, Göttingen, Germany;11. Centre of Biodiversity and Sustainable Land Use (CBL), University of Göttingen, Göttingen, Germany;12. Institute of Biological and Medical Imaging (IBMI), Helmholtz Zentrum München (HMGU)—German Research Center for Environmental Health, Neuherberg, Germany;13. Institute of Biodiversity, Friedrich Schiller University Jena, Jena, Germany;14. Fredericton Research and Development Centre, Agriculture and Agri-Food Canada, Fredericton, NB, Canada;15. Department of Evolutionary Biology and Environmental Studies, University of Zürich, Zürich, Switzerland;16. Asian School of the Environment, Nanyang Technological University, Singapore, Singapore;17. Institute of Plant Sciences, University of Bern, Bern, Switzerland;18. Helmholtz-Institute for Functional Marine Biodiversity at the University of Oldenburg (HIFMB), Oldenburg, Germany;1. State Key Laboratory of Vegetation and Environmental Change, Institute of Botany, the Chinese Academy of Sciences, No. 20 Nanxincun, Xiangshan, 100093 Beijing, China;2. College of life sciences, University of Chinese Academy of Sciences, 100049 Beijing, China;3. Jiangxi Key Laboratory of Plant Resources and Biodiversity, Jingdezhen University, 333400 Jingdezhen, China;1. Martin Luther University Halle-Wittenberg, Institute of Biology/Geobotany and Botanical Garden, Halle (Saale), Germany;2. German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, Leipzig, Germany;3. Department of Biology, McGill University, Montreal, QC, Canada;4. Smithsonian Tropical Research Institute, Panama City, Panama
Abstract:Previous studies have shown that tree species richness increases forest productivity by allowing for greater spatial complementarity of tree crowns (crown complementarity), which in turn results in more densely packed canopies. However, the mechanisms driving crown complementarity in tree species mixtures remain unclear. Here, we take advantage of a high-resolution, three-dimensional terrestrial laser scanning approach in the context of a large-scale biodiversity-ecosystem functioning experiment in subtropical China (BEF-China) to quantify the extent to which functional dissimilarity and divergences in branch traits between neighbouring trees affect crown complementarity at the scale of tree species pairs (i.e., two adjacent trees). Overall, we found no support that functional dissimilarity (divergence in morphological flexibility, specific leaf area and wood density) promotes crown complementarity. However, we show that the effects of functional dissimilarity (the plasticity of the outer crown structure) on crown complementarity vary in their magnitude and importance depending on branch trait divergences. Firstly, crown complementarity tended to be highest for tree species pairs that strongly differed in their functional traits, but were similar in branch density. In contrast, heterospecific pairs with a low functional trait divergence benefitted the most from a large difference in branch density compared with pairs characterised by a large functional dissimilarity. Secondly, the positive effects of increasing divergence in branching intensity (the plasticity of the inner crown structure) on crown complementarity became most important at low levels of functional dissimilarity, i.e. when species pairs were similar in their branch packing and vice versa. This suggests that species mixing allows trees to occupy canopy space more efficiently mainly due to phenotypic changes associated with crown morphology and branch plasticity. Our findings highlight the importance of considering outer and inner crown structures (e.g. branching architecture) to deepen our understanding of tree-tree interactions in mixed-species communities.
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