BACKGROUND AND AIMS: Mixed forest plantations are increasingly recognized for their role in mitigating the impacts of climate change and enhancing ecosystem resilience. Yet, there remains a significant gap in understanding the early-stage dynamics of species trait diversity and interspecies interactions, particularly in pure deciduous mixtures. This study aims to explore the timing and mechanisms by which trait diversity of deciduous species and competitive interactions influence yield, carbon allocation and space occupation in mixed forests, both above and below ground. METHODS: A forest inventory was conducted in planted monocultures, two-species and four-species mixtures of European Acer, Tilia, Carpinus and Quercus, representing a spectrum from acquisitive to conservative tree species. Effects of competition were assessed with linear mixed-effects models at the level of biomass and space acquisition, including leaf, canopy, stem and fine root traits. KEY RESULTS: Early above-ground growth effects were observed 6 years post-planting, with significant biomass accumulation after 8 years, strongly influenced by species composition. Mixtures, especially with acquisitive species, exhibited above-ground overyielding, 1.5-1.9 times higher than monocultures. Fine roots showed substantial overyielding in high-diversity stands. Biomass allocation was species specific and varied markedly by tree size and the level of diversity and between acquisitive Acer and the more conservative species. No root segregation was found. CONCLUSIONS: Our findings underscore the crucial role of species trait diversity in enhancing productivity in mixed deciduous forest plantations. Allometric changes highlight the need to differentiate between (active) acclimatizations and (passive) tree size-related changes, but illustrate major consequences of competitive interactions for the functional relationship between leaves, stem and roots. This study points towards the significant contributions of both above- and below-ground components to overall productivity of planted mixed-species forests.

Federal Research and Training Center for Forests Department of Forest Protection Seckendorff Gudent Weg 8 1131 Vienna Austria

Forest Soils and Biogeochemistry Swiss Federal Institute for Forest Snow and Landscape Research 8903 Birmensdorf Switzerland

Institute of Forest Ecology Department of Forest and Sciences University of Natural Resources and Life Sciences Gregor Mendel Straße 33 1190 Vienna Austria

Mendel University in Brno Faculty of Forestry and Wood Technology Department of Forest Protection and Wildlife Management Zemědělská 3 61300 Brno Czech Republic

Vienna Scientific Instruments Heiligenkreuzer Straße 433 2534 Alland Austria

Zobrazit více v PubMed

Ahmed IU, Smith AR, Godbold DL.. 2019. Polyculture affects biomass production of component species but not total standing biomass and soil carbon stocks in a temperate forest plantation. Annals of Forest Science 76: 91.

Ammer C. 2019. Diversity and forest productivity in a changing climate. The New Phytologist 221: 50–66. PubMed

Barbeito I, Collet C, Ningre F.. 2014. Crown responses to neighbor density and species identity in a young mixed deciduous stand. Trees 28: 1751–1765.

Bastin J-F, Finegold Y, Garcia C, et al.2019. The global tree restoration potential. Science 365: 76–79. PubMed

Bauhus J, Forrester DI, Gardiner B, Jactel H, Vallejo R, Pretzsch H.. 2017. Ecological stability of mixed-species forests. In: Pretzsch H, Forrester DI, Bauhus J, eds. Mixed-species forests: ecology and management. New York, NY: Springer Berlin Heidelberg, 337–382.

Berry E, Anfodillo T, Castorena M, Echeverría A, Olson ME.. 2024. Scaling of leaf area with biomass in trees reconsidered: constant metabolically active sapwood volume per unit leaf area with height growth. Journal of Experimental Botany 75: 3993–4004. PubMed

Blondeel H, Guillemot J, Martin‐StPaul N, et al.2024. Tree diversity reduces variability in sapling survival under drought. Journal of Ecology 112: 1164–1180.

Bloom AJ, Chapin FS, Mooney HA.. 1985. Resource limitation in plants—an economic analogy. Annual Review of Ecology, Evolution, and Systematics 16: 363–392.

Bolte A, Ammer C, Löf M, et al.2009. Adaptive forest management in central Europe: climate change impacts, strategies and integrative concept. Scandinavian Journal of Forest Research 24: 473–482.

Bongers FJ, Schmid B, Bruelheide H, et al.2021. Functional diversity effects on productivity increase with age in a forest biodiversity experiment. Nature Ecology & Evolution 5: 1594–1603. PubMed

Brandeis TJ, Newton M, Cole EC. 2001. Underplanted conifer seedling survival and growth in thinned Douglas-fir stands. Canadian Journal of Forest Research 13: 302–312.

Büttner V, Leuschner C.. 1994. Spatial and temporal patterns of fine root abundance in a mixed oak-beech forest. Forest Ecology and Management 70: 11–21.

Cadotte MW, Carscadden K, Mirotchnick N.. 2011. Beyond species: functional diversity and the maintenance of ecological processes and services. Journal of Applied Ecology 48: 1079–1087.

Caudullo G, de Rigo D.. 2016. Acer platanoides in Europe: distribution, habitat, usage and threats. In: San-Miguel-Ayanz J, de Rigo D, Caudullo G, Durrant TH, Mauri A, eds. European atlas of forest tree species. Luxembourg: Publication Office of the European Union.

Cavard X, Macdonald SE, Bergeron Y, Chen HY.. 2011. Importance of mixedwoods for biodiversity conservation: evidence for understory plants, songbirds, soil fauna, and ectomycorrhizae in northern forests. Environmental Reviews 19: 142–161.

Cope OL, Lindroth RL, Helm A, Keefover‐Ring K, Kruger EL.. 2021. Trait plasticity and trade-offs shape intra-specific variation in competitive response in a foundation tree species. New Phytologist 230: 710–719. PubMed

Davis MA, Wrage KJ, Reich PB.. 1998. Competition between tree seedlings and herbaceous vegetation: support for a theory of resource supply and demand. Journal of Ecology 86: 652–661.

Dietrich P, Ferlian O, Huang Y, Luo S, Quosh J, Eisenhauer N.. 2023. Tree diversity effects on productivity depend on mycorrhizae and life strategies in a temperate forest experiment. Ecology 104: e3896. PubMed

Domisch T, Finér L, Dawud SM, Vesterdal L, Raulund-Rasmussen K.. 2015. Does species richness affect fine root biomass and production in young forest plantations? Oecologia 177: 581–594. PubMed

Dudka VA, Malysheva EF, Malysheva VF, Zhukova EA.. 2023. Mycorrhiza of Linden (Tilia spp.) in artificial plantings in St. Petersburg. Biology Bulletin Reviews 13: S17–S38.

Eaton E, Caudullo G, de Rigo D.. 2016a. Tilia cordata, Tilia platyphyllos and other limes in Europe: distribution, habitat, usage and threats. In: San-Miguel-Ayanz J, de Rigo D, Caudullo G, Durrant TH, Mauri A, eds. European atlas of forest tree species. Luxembourg: Publication Office of the European Union, 184–185.

Eaton E, Caudullo G, Sandra Oliveira, de Rigo D.. 2016b. Quercus robur and Quercus petraea in Europe: distribution, habitat, usage and threats. In: San-Miguel-Ayanz J, de Rigo D, Caudullo G, Durrant TH, Mauri A, eds. European atlas of forest tree species. Luxembourg: Publication Office of the European Union, 160–163.

European Commission. 2022. Three billion additional trees by 2030. European Union: Directorate General for Environment - Publications Office.

Fajardo A, McIntire EJB.. 2011. Under strong niche overlap conspecifics do not compete but help each other to survive: facilitation at the intraspecific level. Journal of Ecology 99: 642–650.

FAO. 2023. Towards more resilient and diverse planted forests. Rome: FAO.

Felton A, Lindbladh M, Brunet J, Fritz O.. 2010. Replacing coniferous monocultures with mixed-species production stands: an assessment of the potential benefits for forest biodiversity in northern Europe. Forest Ecology and Management 260: 939–947.

Fichtner A, Härdtle W, Li Y, Bruelheide H, Kunz M, von Oheimb G.. 2017. From competition to facilitation: how tree species respond to neighbourhood diversity. Ecology Letters 20: 892–900. PubMed

Forrester DI. 2014. The spatial and temporal dynamics of species interactions in mixed-species forests: from pattern to process. Forest Ecology and Management 312: 282–292.

Forrester DI, Ammer C, Annighöfer PJ, et al.2018. Effects of crown architecture and stand structure on light absorption in mixed and monospecific Fagus sylvatica and Pinus sylvestris forests along a productivity and climate gradient through Europe. Journal of Ecology 106: 746–760.

Franklin O, Johansson J, Dewar RC, et al.2012. Modeling carbon allocation in trees: a search for principles. Tree Physiology 32: 648–666. PubMed

Frech A, Leuschner C, Hagemeier M, Hölscher D.. 2003. Neighbor-dependent canopy dimensions of ash, hornbeam, and lime in a species-rich mixed forest (Hainich National Park, Thuringia). Forstwissenschaftliches Centralblatt 122: 22–35.

Freschet GT, Swart EM, Cornelissen JHC.. 2015. Integrated plant phenotypic responses to contrasting above- and below-ground resources: key roles of specific leaf area and root mass fraction. The New Phytologist 206: 1247–1260. PubMed

Freschet GT, Roumet C, Comas LH, et al.2021. Root traits as drivers of plant and ecosystem functioning: current understanding, pitfalls and future research needs. The New Phytologist 232: 1123–1158. PubMed

Fruleux A, Bogeat-Triboulot M-B, Collet C, et al.2018. Aboveground overyielding in a mixed temperate forest is not explained by belowground processes. Oecologia 188: 1183–1193. PubMed

Fu L, Sun H, Sharma RP, Lei Y, Zhang H, Tang S.. 2013. Nonlinear mixed-effects crown width models for individual trees of Chinese fir (Cunninghamia lanceolata) in south-central China. Forest Ecology and Management 302: 210–220.

Glatthorn J. 2021. A spatially explicit index for tree species or trait diversity at neighborhood and stand level. Ecological Indicators 130: 108073.

Gobran GR, Clegg S, Courchesne F.. 1998. Rhizospheric processes influencing the biogeochemistry of forest ecosystems. In: Breemen N, ed. Plant-induced soil changes: processes and feedbacks. Dordrecht: Springer Netherlands, 107–120.

Goldberg DE. 1996. Competitive ability: definitions, contingency and correlated traits. Philosophical Transactions of the Royal Society of London 351: 1377–1385.

Gomarasca U, Migliavacca M, Kattge J, et al.2023. Leaf-level coordination principles propagate to the ecosystem scale. Nature Communications 14: 3948. PubMed PMC

Göransson H, Bambrick MT, Godbold DL.. 2016. Overyielding of temperate deciduous tree mixtures is maintained under throughfall reduction. Plant and Soil 408: 285–298.

Gorné LD, Díaz S, Minden V, et al.2022. The acquisitive–conservative axis of leaf trait variation emerges even in homogeneous environments. Annals of Botany 129: 709–722. PubMed PMC

Grace JB, Tilman D.. 1990. Perspectives on plant competition. San Diego: Academic Press. PubMed

Grams TEE, Andersen CP.. 2007. Competition for resources in trees: physiological versus morphological plasticity. In: Esser K, Löttge U, Beyschlag W, Murata J, eds. Progress in botany. Berlin, Heidelberg: Scholars Portal, 356–381.

Griess VC, Knoke T.. 2011. Growth performance, windthrow, and insects: meta-analyses of parameters influencing performance of mixed-species stands in boreal and northern temperate biomes. Canadian Journal of Forest Research 41: 1141–1159.

Grossman JJ, Vanhellemont M, Barsoum N, et al.2018. Synthesis and future research directions linking tree diversity to growth, survival, and damage in a global network of tree diversity experiments. Environmental and Experimental Botany 152: 68–89.

Hautier Y, Niklaus PA, Hector A.. 2009. Competition for light causes plant biodiversity loss after eutrophication. Science 324: 636–638. PubMed

Hofhansl F, Chacón-Madrigal E, Brännström A, Dieckmann U, Franklin O.. 2021. Mechanisms driving plant functional trait variation in a tropical forest. Ecology and Evolution 11: 3856–3870. PubMed PMC

Huang Y, Chen Y, Castro-Izaguirre N, et al.2018. Impacts of species richness on productivity in a large-scale subtropical forest experiment. Science 362: 80–83. PubMed

Imaji A, Seiwa K.. 2010. Carbon allocation to defense, storage, and growth in seedlings of two temperate broad-leaved tree species. Oecologia 162: 273–281. PubMed

Jacob A, Hertel D, Leuschner C.. 2013. On the significance of belowground overyielding in temperate mixed forests: separating species identity and species diversity effects. Oikos 122: 463–473.

Jacobsen AL. 2023. Growing whole-plant understanding through study of below-ground structural diversity. A commentary on ‘Do root secondary xylem functional traits differ between growth forms in Fabaceae species in a seasonally dry Neotropical environment?’. Annals of Botany 132: i–ii. PubMed PMC

Jactel H, Bauhus J, Boberg J, et al.2017. Tree diversity drives forest stand resistance to natural disturbances. Current Forestry Reports 3: 223–243.

Keenan RJ. 2015. Climate change impacts and adaptation in forest management: a review. Annals of Forest Science 72: 145–167.

Kinzinger L, Mach J, Haberstroh S, et al.2024. Interaction between beech and spruce trees in temperate forests affects water use, root water uptake pattern and canopy structure. Tree Physiology 44: tpad144. PubMed

Lak ZA, Sandén H, Mayer M, Godbold DL, Rewald B.. 2020. Plasticity of root traits under competition for a nutrient-rich patch depends on tree species and possesses a large congruency between intra- and interspecific situations. Forests 11: 528.

Laubhann D, Eckmüllner O, Sterba H.. 2010. Applicability of non-destructive substitutes for leaf area in different stands of Norway spruce (Picea abies L. Karst.) focusing on traditional forest crown measures. Forest Ecology and Management 260: 1498–1506. PubMed PMC

Lei P, Scherer-Lorenzen M, Bauhus J.. 2012a. Belowground facilitation and competition in young tree species mixtures. Forest Ecology and Management 265: 191–200.

Lei P, Scherer-Lorenzen M, Bauhus J.. 2012b. The effect of tree species diversity on fine-root production in a young temperate forest. Oecologia 169: 1105–1115. PubMed

Lenth RV. 2016. Least-squares means: the R package lsmeans. Journal of Statistical Software 69: 1–33.

Leuschner C, Ellenberg H.. 2018. Ecology of central European forests, Revised and extended version of the 6th German edition, corrected publication. Cham: Springer.

Leuschner C, Hertel D, Coners H, Büttner V.. 2001. Root competition between beech and oak: a hypothesis. Oecologia 126: 276–284. PubMed

Leuschner C, Fuchs S, Wedde P, Rüther E, Schuldt B.. 2024. A multi-criteria drought resistance assessment of temperate Acer, Carpinus, Fraxinus, Quercus, and Tilia species. Perspectives in Plant Ecology, Evolution and Systematics 62: 125777.

Liang J, Buongiorno J, Monserud RA, Kruger EL, Zhou M.. 2007. Effects of diversity of tree species and size on forest basal area growth, recruitment, and mortality. Forest Ecology and Management 243: 116–127.

Liu CLC, Kuchma O, Krutovsky KV.. 2018. Mixed-species versus monocultures in plantation forestry: development, benefits, ecosystem services and perspectives for the future. Global Ecology and Conservation 15: e00419.

Lu H, Condés S, Del Río M, et al.2018. Species and soil effects on overyielding of tree species mixtures in the Netherlands. Forest Ecology and Management 409: 105–118.

Lübbe T, Schuldt B, Leuschner C.. 2017. Acclimation of leaf water status and stem hydraulics to drought and tree neighbourhood: alternative strategies among the saplings of five temperate deciduous tree species. Tree Physiology 37: 456–468. PubMed

Luo W, Ni M, Wang Y, et al.2021. Limited evidence of vertical fine‐root segregation in a subtropical forest. New Phytologist 231: 2308–2318. PubMed

Lwila AS, Mund M, Ammer C, Glatthorn J.. 2021. Site conditions more than species identity drive fine root biomass, morphology and spatial distribution in temperate pure and mixed forests. Forest Ecology and Management 499: 119581.

Madrigal-González J, Ruiz-Benito P, Ratcliffe S, et al.2016. Complementarity effects on tree growth are contingent on tree size and climatic conditions across Europe. Scientific Reports 6: 32233. PubMed PMC

Maeght J-L, Rewald B, Pierret A.. 2013. How to study deep roots—and why it matters. Frontiers in Plant Science 4: 299. PubMed PMC

Martin-Blangy S, Meredieu C, Jactel H, Bonal D, Charru M.. 2023. Species-mixing effects on crown dimensions and canopy packing in a young pine–birch plantation are modulated by stand density and irrigation. European Journal of Forest Research 142: 197–216.

Mazal L, Fajardo A, Till-Bottraud I, Corenblit D, Fumanal B.. 2023. Kin selection, kin recognition and kin discrimination in plants revisited: a claim for considering environmental and genetic variability. Plant, Cell & Environment 46: 2007–2016. PubMed

McCormack ML, Gaines KP, Pastore M, Eissenstat DM.. 2015. Early season root production in relation to leaf production among six diverse temperate tree species. Plant and Soil 389: 121–129.

Meinen C, Hertel D, Leuschner C.. 2009. Biomass and morphology of fine roots in temperate broad-leaved forests differing in tree species diversity: is there evidence of below-ground overyielding? Oecologia 161: 99–111. PubMed PMC

Midgley JJ. 2003. Is bigger better in plants? The hydraulic costs of increasing size in trees. Trends in Ecology & Evolution 18: 5–6.

Millard P, Sommerkorn M, Grelet G-A.. 2007. Environmental change and carbon limitation in trees: a biochemical, ecophysiological and ecosystem appraisal. The New Phytologist 175: 11–28. PubMed

Niinemets U. 1998. Growth of young trees of Acer platanoides and Quercus robur along a gap-understory continuum: interrelationships between allometry, biomass partitioning, nitrogen, and shade tolerance. International Journal of Plant Sciences 159: 318–330.

Niklas KJ, Enquist BJ.. 2002. Canonical rules for plant organ biomass partitioning and annual allocation. American Journal of Botany 89: 812–819. PubMed

Niklaus PA, Baruffol M, He J-S, Ma K, Schmid B.. 2017. Can niche plasticity promote biodiversity–productivity relationships through increased complementarity? Ecology 98: 1104–1116. PubMed

Pacala SW, Rees M.. 1998. Models suggesting field experiments to test two hypotheses explaining successional diversity. The American Naturalist 152: 729–737. PubMed

Palacios-Agundez I, Onaindia M, Barraqueta P, Madariaga I.. 2015. Provisioning ecosystem services supply and demand: the role of landscape management to reinforce supply and promote synergies with other ecosystem services. Land Use Policy 47: 145–155.

Pérez-Harguindeguy N, Díaz S, Garnier E, et al.2013. New handbook for standardised measurement of plant functional traits worldwide. Australian Journal of Botany 61: 167.

Petrovska R, Brang P, Gessler A, Bugmann H, Hobi ML.. 2021. Grow slowly, persist, dominate—explaining beech dominance in a primeval forest. Ecology and Evolution 11: 10077–10089. PubMed PMC

Pierik R, Mommer L, Voesenek LA.. 2013. Molecular mechanisms of plant competition: neighbour detection and response strategies. Functional Ecology 27: 841–853.

Pigott CD. 1991. Tilia cordata Miller. The Journal of Ecology 79: 1147–1207.

Pinheiro J, Bates D.. 2023. Nlme: linear and nonlinear mixed effects models: Comprehensive R Archive Network (CRAN). https://doi.org/10.32614/CRAN.package.nlme DOI

Poorter H, Sack L.. 2012. Pitfalls and possibilities in the analysis of biomass allocation patterns in plants. Frontiers in Plant Science 3: 259. PubMed PMC

Poorter H, Niklas KJ, Reich PB, Oleksyn J, Poot P, Mommer L.. 2012. Biomass allocation to leaves, stems and roots: meta-analyses of interspecific variation and environmental control. The New Phytologist 193: 30–50. PubMed

Poorter H, Jagodzinski AM, Ruiz-Peinado R, et al.2015. How does biomass distribution change with size and differ among species? An analysis for 1200 plant species from five continents. The New Phytologist 208: 736–749. PubMed PMC

Power SA, Ashmore MR.. 1996. Nutrient relations and root mycorrhizal status of healthy and declining beech (Fagus sylvatica L.) in Southern Britain. Water, Air, and Soil Pollution 86: 317–333.

Prescott CE, Grayston SJ, Helmisaari H-S, et al.2020. Surplus carbon drives allocation and plant–soil interactions. Trends in Ecology & Evolution 35: 1110–1118. PubMed

Pretzsch H, Schütze G.. 2009. Transgressive overyielding in mixed compared with pure stands of Norway spruce and European beech in Central Europe: evidence on stand level and explanation on individual tree level. European Journal of Forest Research 128: 183–204.

Pretzsch H, Schütze G.. 2016. Effect of tree species mixing on the size structure, density, and yield of forest stands. European Journal of Forest Research 135: 1–22.

Pretzsch H, Bielak K, Block J, et al.2013. Productivity of mixed versus pure stands of oak (Quercus petraea (Matt.) Liebl. and Quercus robur L.) and European beech (Fagus sylvatica L.) along an ecological gradient. European Journal of Forest Research 132: 263–280.

Qiu S, Gao P, Pan L, et al.2023. Developing nonlinear additive tree crown width models based on decomposed competition index and tree variables. Journal of Forestry Research 34: 1407–1422.

R Core Team. 2023. R: a language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing.

Rasmussen CR, Weisbach AN, Thorup-Kristensen K, Weiner J.. 2019. Size-asymmetric root competition in deep, nutrient-poor soil. Journal of Plant Ecology 12: 78–88.

Raulier F, Bernier PY, Ung C-H, Boutin R.. 2002. Structural differences and functional similarities between two sugar maple (Acer saccharum) stands. Tree Physiology 22: 1147–1156. PubMed

Ray T, Delory BM, Beugnon R, et al.2023. Tree diversity increases productivity through enhancing structural complexity across mycorrhizal types. Science Advances 9: eadi2362. PubMed PMC

Reich PB. 2014. The world‐wide ‘fast–slow’ plant economics spectrum: a traits manifesto. Journal of Ecology 102: 275–301.

Rewald B, Leuschner C.. 2009a. Belowground competition in a broad-leaved temperate mixed forest: pattern analysis and experiments in a four-species stand. European Journal of Forest Research 128: 387–398.

Rewald B, Leuschner C.. 2009b. Does root competition asymmetry increase with water availability? Plant Ecology & Diversity 2: 255–264.

Rewald B, Meinen C, Trockenbrodt M, Ephrath JE, Rachmilevitch S.. 2012. Root taxa identification in plant mixtures – current techniques and future challenges. Plant and Soil 359: 165–182.

Rewald B, Rechenmacher A, Godbold DL.. 2014. It’s complicated: intraroot system variability of respiration and morphological traits in four deciduous tree species. Plant Physiology 166: 736–745. PubMed PMC

Ricard J-P, Messier C, Delagrange S, Beaudet M.. 2003. Do understory sapling respond to both light and below-ground competition?: a field experiment in a north-eastern American hardwood forest and a literature review. Annals of Forest Science 60: 749–756.

Rosengren U, Göransson H, Jönsson U, Stjernquist I, Thelin G, Wallander H.. 2006. Functional biodiversity aspects on the nutrient sustainability in forests-importance of root distribution. Journal of Sustainable Forestry 21: 77–100.

RStudio Team. 2023. RStudio: integrated development environment for R. Boston, MA, USA: RStudio, PBC.

Schuster MJ, Williams LJ, Stefanski A, et al.2023. Patterns of belowground overyielding and fine‐root biomass in native and exotic angiosperms and gymnosperms. Oikos 2023: e08877.

Searle EB, Chen HYH, Paquette A.. 2022. Higher tree diversity is linked to higher tree mortality. Proceedings of the National Academy of Sciences of the United States of America 119: e2013171119. PubMed PMC

Seidel D, Leuschner C, Müller A, Krause B.. 2011. Crown plasticity in mixed forests—quantifying asymmetry as a measure of competition using terrestrial laser scanning. Forest Ecology and Management 261: 2123–2132.

Shu W, Shen X, Lei P, Xiang W, Ouyang S, Yan W.. 2018. Temporal changes of fine root overyielding and foraging strategies in planted monoculture and mixed forests. BMC Ecology 18: 9. PubMed PMC

Sikkema R, Caudullo G, de Rigo D.. 2016. Carpinus betulus in Europe: distribution, habitat, usage and threats. In: San-Miguel-Ayanz J, de Rigo D, Caudullo G, Durrant TH, Mauri A, eds. European atlas of forest tree species. Luxembourg: Publication Office of the European Union, 74.

Silvertown J. 2004. Plant coexistence and the niche. Trends in Ecology & Evolution 19: 605–611.

Steinparzer M, Haluza D, Godbold DL.. 2022. Integrating tree species identity and diversity in particulate matter adsorption. Forests 13: 481.

Stotz GC, Salgado‐Luarte C, Escobedo VM, Valladares F, Gianoli E.. 2022. Phenotypic plasticity and the leaf economics spectrum: plasticity is positively associated with specific leaf area. Oikos 2022: 30. doi: https://doi.org/10.1111/oik.09342 DOI

Sun H, Wang X, Fan D.. 2020. Effects of climate, biotic factors, and phylogeny on allometric relationships: testing the metabolic scaling theory in plantations and natural forests across China. Forest Ecosystems 7: 51. doi: https://doi.org/10.1186/s40663-020-00263-y DOI

Tatsumi S. 2020. Tree diversity effects on forest productivity increase through time because of spatial partitioning. Forest Ecosystems 7: 24.

Thompson RA. 2023. A neutral theory of plant carbon allocation. Tree Physiology 44: tpad151. PubMed

Thurm EA, Pretzsch H.. 2016. Improved productivity and modified tree morphology of mixed versus pure stands of European beech (Fagus sylvatica) and Douglas-fir (Pseudotsuga menziesii) with increasing precipitation and age. Annals of Forest Science 73: 1047–1061.

Urgoiti J, Messier C, Keeton WS, Reich PB, Gravel D, Paquette A.. 2022. No complementarity no gain—Net diversity effects on tree productivity occur once complementarity emerges during early stand development. Ecology Letters 25: 851–862. PubMed

Urgoiti J, Messier C, Keeton WS, Belluau M, Paquette A.. 2023a. Functional diversity and identity influence the self‐thinning process in young forest communities. Journal of Ecology 111: 2010–2022.

Urgoiti J, Messier C, Keeton WS, Paquette A.. 2023b. Tree community overyielding during early stand development is explained by asymmetric species‐specific responses to diversity. Functional Ecology 37: 2621–2633.

Usman M, Ho-Plágaro T, Frank HER, et al.2021. Mycorrhizal symbiosis for better adaptation of trees to abiotic stress caused by climate change in temperate and boreal forests. Frontiers in Forests and Global Change 4.:742392. doi: https://doi.org/10.3389/ffgc.2021.742392 DOI

Valverde-Barrantes OJ, Smemo KA, Feinstein LM, Kershner MW, Blackwood CB.. 2015. Aggregated and complementary: symmetric proliferation, overyielding, and mass effects explain fine-root biomass in soil patches in a diverse temperate deciduous forest landscape. The New Phytologist 205: 731–742. PubMed

van de Peer T, Verheyen K, Kint V, van Cleemput E, Muys B.. 2017a. Plasticity of tree architecture through interspecific and intraspecific competition in a young experimental plantation. Forest Ecology and Management 385: 1–9.

van de Peer T, Verheyen K, Ponette Q, Setiawan NN, Muys B.. 2017b. Overyielding in young tree plantations is driven by local complementarity and selection effects related to shade tolerance. Journal of Ecology 106: 1096–1105.

Vanderwel MC, Malcolm JR, Smith SM.. 2006. An integrated model for snag and downed woody debris decay class transitions. Forest Ecology and Management 234: 48–59.

Verheyen K, Vanhellemont M, Auge H, et al.2016. Contributions of a global network of tree diversity experiments to sustainable forest plantations. Ambio 45: 29–41. PubMed PMC

Vospernik S, Heym M, Pretzsch H, et al.2023. Tree species growth response to climate in mixtures of Quercus robur/Quercus petraea and Pinus sylvestris across Europe - a dynamic, sensitive equilibrium. Forest Ecology and Management 530: 120753.

Wambsganss J, Beyer F, Freschet GT, Scherer‐Lorenzen M, Bauhus J.. 2021a. Tree species mixing reduces biomass but increases length of absorptive fine roots in European forests. Journal of Ecology 109: 2678–2691.

Wambsganss J, Freschet GT, Beyer F, et al.2021b. Tree species mixing causes a shift in fine‐root soil exploitation strategies across European forests. Functional Ecology 35: 1886–1902.

Weigelt A, Mommer L, Andraczek K, et al.2021. An integrated framework of plant form and function: the belowground perspective. The New Phytologist 232: 42–59. PubMed

Weithmann G, Schuldt B, Link RM, et al.2022. Leaf trait modification in European beech trees in response to climatic and edaphic drought. Plant Biology 24: 1272–1286. PubMed

West PW. 2015. Tree and forest measurement, 3rd edn. Cham, Heidelberg, New York [etc.]: Springer.

Wickham H. 2016. ggplot2: elegant graphics for data analysis. New York, NY: Springer Science+Business Media LLC.

Williams LJ, Paquette A, Cavender-Bares J, Messier C, Reich PB.. 2017. Spatial complementarity in tree crowns explains overyielding in species mixtures. Nature Ecology & Evolution 1: 63. PubMed

Williams LJ, Butler EE, Cavender-Bares J, et al.2021. Enhanced light interception and light use efficiency explain overyielding in young tree communities. Ecology Letters 24: 996–1006. PubMed

Wortemann R, Herbette S, Barigah TS, et al.2011. Genotypic variability and phenotypic plasticity of cavitation resistance in Fagus sylvatica L. across Europe. Tree Physiology 31: 1175–1182. PubMed

Xiang Y, Li Y, Luo X, et al.2022. Mixed plantations enhance more soil organic carbon stocks than monocultures across China: implication for optimizing afforestation/reforestation strategies. The Science of the Total Environment 821: 153449. PubMed

Zheng L-T, Chen HYH, Biswas SR, et al.2021. Diversity and identity of economics traits determine the extent of tree mixture effects on ecosystem productivity. Journal of Ecology 109: 1898–1908.