Global change is affecting the distribution and population dynamics of plant species across the planet, leading to trends such as shifts in distribution toward the poles and to higher elevations. Yet, we poorly understand why individual species respond differently to warming and other environmental changes, or how the trait composition of communities responds. Here we ask two questions regarding plant species and community changes over 42 years of global change in a temperate montane forest in Québec, Canada: (1) How did the trait composition, alpha diversity, and beta diversity of understory vascular plant communities change between 1970 and 2010, a period over which the region experienced 1.5°C of warming and changes in nitrogen deposition? (2) Can traits predict shifts in species elevation and abundance over this time period? For 46 understory vascular species, we locally measured six aboveground traits, and for 36 of those (not including shrubs), we also measured five belowground traits. Collectively, they capture leading dimensions of phenotypic variation that are associated with climatic and resource niches. At the community level, the trait composition of high-elevation plots shifted, primarily for two root traits: specific root length decreased and rooting depth increased. The mean trait values of high-elevation plots shifted over time toward values initially associated with low-elevation plots. These changes led to trait homogenization across elevations. The community-level shifts in traits mirrored the taxonomic shifts reported elsewhere for this site. At the species level, two of the three traits predicting changes in species elevation and abundance were belowground traits (low mycorrhizal fraction and shallow rooting). These findings highlight the importance of root traits, which, along with leaf mass fraction, were associated with shifts in distribution and abundance over four decades. Community-level trait changes were largely similar across the elevational and temporal gradients. In contrast, traits typically associated with lower elevations at the community level did not predict differences among species in their shift in abundance or distribution, indicating a decoupling between species- and community-level responses. Overall, changes were consistent with some influence of both climate warming and increased nitrogen availability.

CEFE CNRS Univ Montpellier EPHE IRD Montpellier France

Département de Biologie Université de Sherbrooke Sherbrooke Québec Canada

Department of Agriculture Faculty of Agriculture and Forestry University of Helsinki Helsinki Finland

Department of Biology University of Waterloo Waterloo Ontario Canada

Department of Vegetation Ecology Institute of Botany Czech Academy of Sciences Brno Czech Republic

State Key Laboratory of Biocontrol School of Ecology Sun Yat sen University Guangzhou China

Zobrazit více v PubMed

Ackerly, D. D., C. A. Knight, S. B. Weiss, K. Barton, and K. P. Starmer. 2002. “Leaf Size, Specific Leaf Area and Microhabitat Distribution of Chaparral Woody Plants: Contrasting Patterns in Species Level and Community Level Analyses.” Oecologia 130: 449–457.

Amatangelo, K. L., S. E. Johnson, D. A. Rogers, and D. M. Waller. 2014. “Trait‐Environment Relationships Remain Strong despite 50 Years of Trait Compositional Change in Temperate Forests.” Ecology 95: 1780–1791.

Aubin, I., L. Boisvert‐Marsh, H. Kebli, D. McKenney, J. Pedlar, K. Lawrence, E. H. Hogg, Y. Boulanger, S. Gauthier, and C. Ste‐Marie. 2018. “Tree Vulnerability to Climate Change: Improving Exposure‐Based Assessments Using Traits as Indicators of Sensitivity.” Ecosphere 9: e02108.

Aubin, I., A. D. Munson, F. Cardou, P. J. Burton, N. Isabel, J. H. Pedlar, A. Paquette, et al. 2016. “Traits to Stay, Traits to Move: A Review of Functional Traits to Assess Sensitivity and Adaptive Capacity of Temperate and Boreal Trees to Climate Change.” Environmental Reviews 23: 1–23.

Barton, K. 2024. “MuMIn: Multi‐Model Inference.” https://cran.r-project.org/web/packages/MuMIn/index.html.

Bates, D., M. Maechler, B. Bolker, and S. Walker. 2015. “Fitting Linear Mixed‐Effects Models Using lme4.” Journal of Statistical Software 67: 1–48.

Becker‐Scarpitta, A., S. Vissault, and M. Vellend. 2019. “Four Decades of Plant Community Change along a Continental Gradient of Warming.” Global Change Biology 25: 1629–1641.

Bergmann, J., A. Weigelt, F. van der Plas, D. C. Laughlin, T. W. Kuyper, N. Guerrero‐Ramirez, O. J. Valverde‐Barrantes, et al. 2020. “The Fungal Collaboration Gradient Dominates the Root Economics Space in Plants.” Science Advances 6: 1–10.

Blondeel, H., M. P. Perring, L. Depauw, E. De Lombaerde, D. Landuyt, P. De Frenne, and K. Verheyen. 2020. “Light and Warming Drive Forest Understorey Community Development in Different Environments.” Global Change Biology 26: 1681–1696.

Bobbink, R. 2004. “Plant Species Richness and the Exceedance of Empirical Nitrogen Critical Loads: An Inventory.” Report Landscape Ecology. Utrecht University/RIVM, Bilthoven, The Netherlands. https://www.researchgate.net/publication/46651318_Plant_Species_Richness_and_the_Exceedance_of_Empirical_Nitrogen_Critical_Loads_An_Inventory.

Bobbink, R., K. Hicks, J. Galloway, T. Spranger, R. Alkemade, M. Ashmore, M. Bustamante, et al. 2010. “Global Assessment of Nitrogen Deposition Effects on Terrestrial Plant Diversity: A Synthesis.” Ecological Applications 20: 30–59.

Brown, J. K., and J. C. Zinnert. 2021. “Trait‐Based Investigation Reveals Patterns of Community Response to Nutrient Enrichment in Coastal Mesic Grassland.” Diversity (Basel) 13: 1–15.

Carteron, A., V. Parasquive, F. Blanchard, X. Guilbeault‐Mayers, B. L. Turner, M. Vellend, and E. Laliberté. 2020. “Soil Abiotic and Biotic Properties Constrain the Establishment of a Dominant Temperate Tree into Boreal Forests.” Journal of Ecology 108: 931–944.

Caruso, C. M., H. Maherali, and R. A. Martin. 2020. “A Meta‐Analysis of Natural Selection on Plant Functional Traits.” International Journal of Plant Sciences 181: 44–55.

Chen, I. C., J. K. Hill, R. Ohlemüller, D. B. Roy, and C. D. Thomas. 2011. “Rapid Range Shifts of Species of Climate Warming.” Science 1979(333): 1024–1026.

Clark, J. S. 2016. “Why Species Tell More about Traits than Traits about Species: Predictive Analysis.” Ecology 97: 1979–1993.

Comas, L. H., T. J. Bouma, and D. M. Eissenstat. 2002. “Linking Root Traits to Potential Growth Rate in Six Temperate Tree Species.” Oecologia 132: 34–43.

Crimmins, S. M., S. Z. Dobrowski, J. A. Greenberg, J. T. Abatzoglou, and A. R. Mynsberge. 2011. “Changes in Climatic Water Balance Drive Downhill Shifts in Plant Species' Optimum Elevations.” Science 1979(331): 324–327.

Danneyrolles, V., S. Dupuis, G. Fortin, M. Leroyer, A. de Römer, R. Terrail, M. Vellend, et al. 2019. “Stronger Influence of Anthropogenic Disturbance than Climate Change on Century‐Scale Compositional Changes in Northern Forests.” Nature Communications 10: 1265.

Diaz, S., J. Kattge, J. H. Cornelissen, I. J. Wright, S. Lavorel, S. Dray, B. Reu, et al. 2016. “The Global Spectrum of Plant Form and Function.” Nature 14: 1–17.

Diaz, S., S. Lavorel, F. De Bello, F. Quétier, K. Grigulis, and M. Robson. 2007. “Incorporating Plant Functional Diversity Effects in Ecosystem Assessments.” Proceedings of the National Academy of Sciences of the United States of America 104: 20684–20689.

Diekmann, M., and U. Falkengren‐Grerup. 2002. “Prediction of Species Response to Atmospheric Nitrogen Deposition by Means of Ecological Measures and Life History Traits.” Journal of Ecology 90: 108–120.

Dornelas, M., A. E. Magurran, S. T. Buckland, A. Chao, R. L. Chazdon, R. K. Colwell, T. Curtis, et al. 2013. “Quantifying Temporal Change in Biodiversity: Challenges and Opportunities.” Proceedings of the Royal Society B: Biological Sciences 280: 20121931.

Dullinger, S., A. Gattringer, W. Thuiller, D. Moser, N. E. Zimmermann, A. Guisan, W. Willner, et al. 2012. “Extinction Debt of High‐Mountain Plants under Twenty‐First‐Century Climate Change.” Nature Climate Change 2: 619–622.

Environment and Climate Change Canada. 2023. “Wet Deposition Maps.” https://open.canada.ca/data/en/dataset/e8896575‐1fb8‐4e53‐8acd‐8579c3c055c2.

Fan, Y., G. Miguez‐Macho, E. G. Jobbágy, R. B. Jackson, and C. Otero‐Casal. 2017. “Hydrologic Regulation of Plant Rooting Depth.” Proceedings of the National Academy of Sciences of the United States of America 114: 10572–10577.

Feeley, K. J., M. R. Silman, M. B. Bush, W. Farfan, K. G. Cabrera, Y. Malhi, P. Meir, N. S. Revilla, M. N. R. Quisiyupanqui, and S. Saatchi. 2011. “Upslope Migration of Andean Trees.” Journal of Biogeography 38: 783–791.

Fischer, A. G. 1960. “Latitudinal Variations in Organic Diversity.” Evolution (N Y) 14: 64–81.

Fonseca, C. R., J. M. Overton, B. Collins, and M. Westoby. 2000. “Shifts in Trait‐Combinations along Rainfall and Phosphorus Gradients.” Journal of Ecology 88: 964–977.

Freschet, G. T., P. J. Bellingham, P. O. B. Lyver, K. I. Bonner, and D. A. Wardle. 2013. “Plasticity in Above‐ and Belowground Resource Acquisition Traits in Response to Single and Multiple Environmental Factors in Three Tree Species.” Ecology and Evolution 3: 1065–1078.

Freschet, G. T., C. Roumet, L. H. Comas, M. Weemstra, A. G. Bengough, B. Rewald, R. D. Bardgett, et al. 2021. “Root Traits as Drivers of Plant and Ecosystem Functioning: Current Understanding, Pitfalls and Future Research Needs.” New Phytologist 232: 1123–1158.

Freschet, G. T., E. M. Swart, and J. H. C. Cornelissen. 2015. “Integrated Plant Phenotypic Responses to Contrasting Above‐ and Below‐Ground Resources: Key Roles of Specific Leaf Area and Root Mass Fraction.” New Phytologist 206: 1247–1260.

Freschet, G. T., O. J. Valverde‐Barrantes, C. M. Tucker, J. M. Craine, M. L. McCormack, C. Violle, F. Fort, et al. 2017. “Climate, Soil and Plant Functional Types as Drivers of Global Fine‐Root Trait Variation.” Journal of Ecology 105: 1182–1196.

Freschet, G. T., C. Violle, M. Y. Bourget, M. Scherer‐Lorenzen, and F. Fort. 2018. “Allocation, Morphology, Physiology, Architecture: The Multiple Facets of Plant Above‐ and Below‐Ground Responses to Resource Stress.” New Phytologist 219: 1338–1352.

Fukami, T., and D. A. Wardle. 2005. “Long‐Term Ecological Dynamics: Reciprocal Insights from Natural and Anthropogenic Gradients.” Proceedings of the Royal Society B: Biological Sciences 272: 2105–2115.

Gallagher, R. V., L. Hughes, and M. R. Leishman. 2013. “Species Loss and Gain in Communities under Future Climate Change: Consequences for Functional Diversity.” Ecography 36: 531–540.

Garnier, E., M.‐L. Navas, and K. Grigulis. 2016. Plant Functional Diversity: Organism Traits, Community Structure, and Ecosystem Properties. Oxford, UK: Oxford University Press.

Geber, M. A., and L. R. Griffen. 2003. “Inheritance and Natural Selection on Functional Traits.” International Journal of Plant Sciences 164: S21–S42.

Givnish, T. J. 1987. “Comparative Studies of Leaf Form: Assessing the Relative Roles of Selective Pressures and Phylogenetic Constraints.” New Phytologist 106: 131–160.

Govaert, S., P. Vangansbeke, H. Blondeel, K. Steppe, K. Verheyen, and P. De Frenne. 2021. “Rapid Thermophilization of Understorey Plant Communities in a 9 Year‐Long Temperate Forest Experiment.” Journal of Ecology 109: 2434–2447.

Granier, C., B. Bessagnet, T. Bond, A. D'Angiola, H. Denier van der Gon, G. J. Frost, A. Heil, et al. 2011. “Evolution of Anthropogenic and Biomass Burning Emissions of Air Pollutants at Global and Regional Scales during the 1980–2010 Period.” Climate Change 109: 163–190.

Gutschick, V. P., and H. BassiriRad. 2003. “Extreme Events as Shaping Physiology, Ecology, and Evolution of Plants: Toward a Unified Definition & Evaluation of Their Consequences.” New Phytologist 160: 21–42.

Hall, G. 1998. “Inventaire floristique du Mont Mégantic.” Rapport préparé pour le Ministère de l'Environnement et de la Faune du Québec, Parc de conservation du Mont Mégantic.

Hallbäcken, L., and L. Q. Zhang. 1998. “Effects of Experimental Acidification, Nitrogen Addition and Liming on Ground Vegetation in a Mature Stand of Norway Spruce (Picea abies (L.) Karst.) in SE Sweden.” Forest Ecology and Management 108: 201–213.

Harrison, S. P., E. S. Gornish, and S. Copeland. 2015. “Climate‐Driven Diversity Loss in a Grassland Community.” Proceedings of the National Academy of Sciences of the United States of America 112: 8672–8677.

Hember, R. A. 2018. “Spatially and Temporally Continuous Estimates of Annual Total Nitrogen Deposition over North America, 1860–2013.” Data Brief 17: 134–140.

Holdaway, R. J., S. J. Richardson, I. A. Dickie, D. A. Peltzer, and D. A. Coomes. 2011. “Species‐ and Community‐Level Patterns in Fine Root Traits along a 120,000‐Year Soil Chronosequence in Temperate Rain Forest.” Journal of Ecology 99: 954–963.

Johnson, E. A., and K. Miyanishi. 2008. “Testing the Assumptions of Chronosequences in Succession.” Ecology Letters 11: 419–431.

Katabuchi, M., S. J. Wright, N. G. Swenson, K. J. Feeley, R. Condit, S. P. Hubbell, and S. J. Davies. 2017. “Contrasting Outcomes of Species‐ and Community‐Level Analyses of the Temporal Consistency of Functional Composition.” Ecology 98: 2273–2280.

Kattge, J., G. Bönisch, S. Díaz, S. Lavorel, I. C. Prentice, P. Leadley, S. Tautenhahn, et al. 2020. “TRY Plant Trait Database – Enhanced Coverage and Open Access.” Global Change Biology 26: 119–188.

Kelly, A. E., and M. L. Goulden. 2008. “Rapid Shifts in Plant Distribution with Recent Climate Change.” Proceedings of the National Academy of Sciences of the United States of America 105: 11823–11826.

Kerkhoff, A. J., and B. J. Enquist. 2009. “Multiplicative by Nature: Why Logarithmic Transformation Is Necessary in Allometry.” Journal of Theoretical Biology 257: 519–521.

Kermavnar, J., L. Kutnar, and A. Marinšek. 2021. “Disentangling the Ecological Determinants of Species and Functional Trait Diversity in Herb‐Layer Plant Communities in European Temperate Forests.” Forests 12: 552.

Kramer‐Walter, K. R., P. J. Bellingham, T. R. Millar, R. D. Smissen, S. J. Richardson, and D. C. Laughlin. 2016. “Root Traits Are Multidimensional: Specific Root Length Is Independent from Root Tissue Density and the Plant Economic Spectrum.” Journal of Ecology 104: 1299–1310.

Kramer‐Walter, K. R., and D. C. Laughlin. 2017. “Root Nutrient Concentration and Biomass Allocation Are More Plastic than Morphological Traits in Response to Nutrient Limitation.” Plant Soil 416: 539–550.

Laliberté, E., P. Legendre, and B. Shipley. 2014. “FD: Measuring Functional Diversity from Multiple Traits, and Other Tools for Functional Ecology.” https://cran.r-project.org/web/packages/FD/FD.pdf.

Lamarque, J. F., F. Dentener, J. McConnell, C. U. Ro, M. Shaw, R. Vet, D. Bergmann, et al. 2013. “Multi‐Model Mean Nitrogen and Sulfur Deposition from the Atmospheric Chemistry and Climate Model Intercomparison Project (ACCMIP): Evaluation of Historical and Projected Future Changes.” Atmospheric Chemistry and Physics 13: 7997–8018.

Laughlin, D. C., and J. Messier. 2015. “Fitness of Multidimensional Phenotypes in Dynamic Adaptive Landscapes.” Trends in Ecology & Evolution 30: 487–496.

Laughlin, D. C., L. Mommer, F. M. Sabatini, H. Bruelheide, T. W. Kuyper, M. L. McCormack, J. Bergmann, et al. 2021. “Root Traits Explain Plant Species Distributions along Climatic Gradients Yet Challenge the Nature of Ecological Trade‐Offs.” Nature Ecology and Evolution 5: 1123–1134.

Laughlin, D. C., R. T. Strahan, P. B. Adler, and M. M. Moore. 2018. “Survival Rates Indicate that Correlations between Community‐Weighted Mean Traits and Environments Can be Unreliable Estimates of the Adaptive Value of Traits.” Ecology Letters 21: 411–421.

Lavorel, S., and E. Garnier. 2002. “Predicting Changes in Community Composition and Ecosystem Functioning from Plant Traits: Revisiting the Holy Grail.” Functional Ecology 16: 545–556.

Lenoir, J., J. C. Gégout, P. A. Marquet, P. de Ruffray, and H. Brisse. 2008. “A Significant Upward Shift in Plant Species Optimum Elevation during the 20th Century.” Science 320: 1768–1771.

Likens, G. E., D. C. Buso, E. S. Bernhardt, and E. Rosi. 2021. “A Century of Change: Reconstructing the Biogeochemical History of Hubbard Brook.” Hydrological Processes 35: e14256.

Ma, Z., D. Guo, X. Xu, M. Lu, R. D. Bardgett, D. M. Eissenstat, M. L. McCormack, and L. O. Hedin. 2018. “Evolutionary History Resolves Global Organization of Root Functional Traits.” Nature 555: 94–97.

Maes, S. L., M. P. Perring, L. Depauw, M. Bernhardt‐Römermann, H. Blondeel, G. Brūmelis, J. Brunet, et al. 2020. “Plant Functional Trait Response to Environmental Drivers across European Temperate Forest Understorey Communities.” Plant Biology 22: 410–424.

Marcotte, G., and M. Grandtner. 1974. Étude écologique de la végétation forestière du Mont Mégantic. MSc thesis, Université Laval.

McCormack, L. M., I. A. Dickie, D. M. Eissenstat, T. J. Fahey, C. W. Fernandez, D. Guo, H. S. Helmisaari, et al. 2015. “Redefining Fine Roots Improves Understanding of Below‐Ground Contributions to Terrestrial Biosphere Processes.” New Phytologist 207: 505–518.

McCormack, M. L., T. S. Adams, E. A. H. Smithwick, and D. M. Eissenstat. 2012. “Predicting Fine Root Lifespan from Plant Functional Traits in Temperate Trees.” New Phytologist 195: 823–831.

McKenney, D. W., J. H. Pedlar, P. Papadopol, and M. F. Hutchinson. 2006. “The Development of 1901–2000 Historical Monthly Climate Models for Canada and the United States.” Agricultural and Forest Meteorology 138: 69–81.

Messier, J. 2019. “juliemessier/Megantic‐Trait‐Data: Mont_Mégantic_Individual_Traits_2016–2017 (v1.0.0).” Zenodo. https://doi.org/10.5281/zenodo.3258342

Mitchell, R. M., J. P. Wright, and G. M. Ames. 2018. “Species' Traits Do Not Converge on Optimum Values in Preferred Habitats.” Oecologia 186: 719–729.

Moles, A. T., S. E. Perkins, S. W. Laffan, H. Flores‐Moreno, M. Awasthy, M. L. Tindall, L. Sack, et al. 2014. “Which Is a Better Predictor of Plant Traits: Temperature or Precipitation?” Journal of Vegetation Science 25: 1167–1180.

Nippert, J. B., and R. M. Holdo. 2015. “Challenging the Maximum Rooting Depth Paradigm in Grasslands and Savannas.” Functional Ecology 29: 739–745.

Oksanen, J., F. G. Blanchet, R. Kindt, P. Legendre, P. R. Minchin, R. B. O'hara, G. L. Simpson, et al. 2019. “vegan: Community Ecology Package.” https://cran.r-project.org/web/packages/vegan/vegan.pdf.

Olson, M. E., R. Aguirre‐Hernández, and J. A. Rosell. 2009. “Universal Foliage‐Stem Scaling across Environments and Species in Dicot Trees: Plasticity, Biomechanics and Corner's Rules.” Ecology Letters 12: 210–219.

Parmesan, C., and G. Yohe. 2003. “A Globally Coherent Fingerprint of Climate Change Impacts across Natural Systems.” Nature 421: 37–42.

Pauli, H., M. Gottfried, S. Dullinger, O. Abdaladze, M. Akhalkatsi, J. L. B. Alonso, G. Coldea, et al. 2012. “Recent Plant Diversity Changes on Europe's Mountain Summits.” Science 1979(336): 353–355.

Pérez‐Harguindeguy, N., S. Díaz, E. Garnier, S. Lavorel, H. Poorter, P. Jaureguiberry, M. S. Bret‐Harte, et al. 2013. “New Handbook for Standardised Measurement of Plant Functional Traits Worldwide.” Australian Journal of Botany 61: 167–234.

Perring, M. P., M. Bernhardt‐Römermann, L. Baeten, G. Midolo, H. Blondeel, L. Depauw, D. Landuyt, et al. 2018. “Global Environmental Change Effects on Plant Community Composition Trajectories depend upon Management Legacies.” Global Change Biology 24: 1722–1740.

Pickett, S. T. A. 1989. “Space‐for‐Time Substitution as an Alternative to Long‐Term Studies.” In Long‐Term Studies in Ecology, edited by G. E. Likens, 110–135. New York, NY: Springer. https://doi.org/10.1007/978-1-4615-7358-6_5.

Poorter, H., K. J. Niklas, P. B. Reich, J. Oleksyn, P. Poot, and L. Mommer. 2012. “Biomass Allocation to Leaves, Stems and Roots: Meta‐Analyses of Interspecific Variation and Environmental Control.” New Phytologist 193: 30–50.

Pregitzer, K. S., J. L. DeForest, A. J. Burton, M. F. Allen, R. W. Ruess, and R. L. Hendrick. 2002. “Fine Root Architecture of Nine North American Trees.” Ecological Monographs 72: 293–309.

R Core Team. 2020. R: A Language and Environment for Statistical Computing. Vienna: R Foundation for Statistical Computing.

Reich, P. B., D. S. Ellsworth, M. B. Walters, J. M. Vose, C. Gresham, J. C. Volin, and W. D. Bowman. 1999. “Generality of Leaf Trait Relationships: A Test across 6 Biomes.” Ecology 80: 1955–1969.

Ripley, B. 2019. “Tree: Classification and Regression Trees.” https://cran.r-project.org/web/packages/tree/tree.pdf.

Roumet, C., C. Urcelay, and S. Díaz. 2006. “Suites of Root Traits Differ between Annual and Perennial Species Growing in the Field.” New Phytologist 170: 357–368.

Rumpf, S. B., K. Hülber, G. Klonner, D. Moser, M. Schütz, J. Wessely, W. Willner, N. E. Zimmermann, and S. Dullinger. 2018. “Range Dynamics of Mountain Plants Decrease with Elevation.” Proceedings of the National Academy of Sciences of the United States of America 115: 1848–1853.

Salguero‐Gómez, R., C. Violle, O. Gimenez, and D. Childs. 2018. “Delivering the Promises of Trait‐Based Approaches to the Needs of Demographic Approaches.” Functional Ecology 32: 1424–1435.

Savage, J., and M. Vellend. 2014. “Elevational Shifts, Biotic Homogenization and Time Lags in Vegetation Change during 40 Years of Climate Warming.” Ecography 38: 546–555.

Schenk, H. J., and R. B. Jackson. 2002. “The Global Biogeography of Roots.” Ecological Monographs 72: 311–328.

Schulze, E. D., H. A. Mooney, O. E. Sala, E. Jobbagy, N. Buchmann, G. Bauer, J. Canadell, et al. 1996. “Rooting Depth, Water Availability, and Vegetation Cover along an Aridity Gradient in Patagonia.” Oecologia 108: 503–511.

Shipley, B., F. De Bello, J. H. C. Cornelissen, E. Laliberté, D. C. Laughlin, and P. B. Reich. 2016. “Reinforcing Loose Foundation Stones in Trait‐Based Plant Ecology.” Oecologia 180: 923–931.

Société des Établissements de Plein air du Québec (SEPAQ). 2007. “Synthèse des connaissances: Parc National du Mont‐Mégantic.”

Sonnier, G., S. E. Johnson, K. L. Amatangelo, D. A. Rogers, and D. M. Waller. 2014. “Is Taxonomic Homogenization Linked to Functional Homogenization in Temperate Forests?” Global Ecology and Biogeography 23: 894–902.

Swenson, N. G., B. J. Enquist, J. Pither, A. J. Kerkhoff, B. Boyle, M. D. Weiser, J. J. Elser, et al. 2012. “The Biogeography and Filtering of Woody Plant Functional Diversity in North and South America.” Global Ecology and Biogeography 21: 798–808.

Swenson, N. G., and M. D. Weiser. 2010. “Plant Geography upon the Basis of Functional Traits: An Example from Eastern North American Trees.” Ecology 91: 2234–2241.

Talluto, M. V., I. Boulangeat, S. Vissault, W. Thuiller, and D. Gravel. 2017. “Extinction Debt and Colonization Credit Delay Range Shifts of Eastern North American Trees.” Nature Ecology and Evolution 1: 1–6.

ter Braak, C. J. F. 2019. “New Robust Weighted Averaging‐ and Model‐Based Methods for Assessing Trait–Environment Relationships.” Methods in Ecology and Evolution 10: 1962–1971.

Thuiller, W., S. Lavorel, M. T. Sykes, and M. B. Araújo. 2006. “Using Niche‐Based Modelling to Assess the Impact of Climate Change on Tree Functional Diversity in Europe.” Diversity and Distributions 12: 49–60.

Tilman, D. 1988. Plant Strategies and the Dynamics and Structure of Plant Communities. Princeton, NJ: Princeton University Press.

Treseder, K. K., and P. M. Vitousek. 2001. “Effects of Soil Nutrient Availability on Investment in Acquisition of N and P in Hawaiian Forests.” Ecology 82: 946–954.

Umaña, M. N., M. Cao, L. Lin, N. G. Swenson, and C. Zhang. 2021. “Trade‐Offs in Above‐ and Below‐Ground Biomass Allocation Influencing Seedling Growth in a Tropical Forest.” Journal of Ecology 109: 1184–1193.

Vanneste, T., A. Valdés, K. Verheyen, M. P. Perring, M. Bernhardt‐Römermann, E. Andrieu, J. Brunet, et al. 2019. “Functional Trait Variation of Forest Understorey Plant Communities across Europe.” Basic and Applied Ecology 34: 1–14.

Vellend, M., L. Baeten, A. Becker‐Scarpitta, V. Boucher‐Lalonde, J. L. McCune, J. Messier, I. H. Myers‐Smith, and D. F. Sax. 2017. “Plant Biodiversity Change across Scales during the Anthropocene.” Annual Review of Plant Biology 68: 563–586.

Vellend, M., L. Baeten, I. H. Myers‐Smith, S. C. Elmendorf, R. Beauséjour, C. D. Brown, P. De Frenne, K. Verheyen, and S. Wipf. 2013. “Global Meta‐Analysis Reveals No Net Change in Local‐Scale Plant Biodiversity over Time.” Proceedings of the National Academy of Sciences of the United States of America 110: 19456–19459.

Weathers, K. C., S. M. Simkin, G. M. Lovett, and S. E. Lindberg. 2006. “Empirical Modeling of Atmospheric Deposition in Mountainous Landscapes.” Ecological Applications 16: 1590–1607.

Weemstra, M., L. Mommer, E. J. W. Visser, J. van Ruijven, T. W. Kuyper, G. M. J. Mohren, and F. J. Sterck. 2016. “Towards a Multidimensional Root Trait Framework: a Tree Root Review.” New Phytol 211: 1159–1169.

Weemstra, M., O. J. Valverde‐Barrantes, M. L. McCormack, and D. Kong. 2023. “Root Traits and Functioning: From Individual Plants to Ecosystems.” Oikos 2023: 1–9.

Westoby, M. 1998. “A Leaf‐Height‐Seed Plant Ecology Strategy Scheme.” Plant Soil 199: 213–227.

Westoby, M., D. Falster, A. Moles, P. Vesk, and I. Wright. 2002. “Plant Ecological Strategies: Some Leading Dimensions of Variation between Species.” Annual Review of Ecology, Evolution, and Systematics 33: 125–159.

White, P. S. 1983. “Corner's Rules in Eastern Deciduous Trees: Allometry and Its Implications for the Adaptive Architecture of Trees.” Bulletin of the Torrey Botanical Club 110: 203–212.

Wieczynskiak, D. J., B. Boyle, V. Buzzard, S. M. Duran, A. N. Henderson, C. M. Hulshof, A. J. Kerkhoff, et al. 2019. “Climate Shapes and Shifts Functional Biodiversity in Forests Worldwide.” Proceedings of the National Academy of Sciences of the United States of America 116: 587–592.

Withington, J. M., P. B. Reich, J. Oleksyn, and D. M. Eissenstat. 2006. “Comparisons of Structure and Life Span in Roots and Leaves.” Ecological Monographs 76: 381–397.

Wright, I. J., P. B. Reich, M. Westoby, D. D. Ackerly, Z. Baruch, F. Bongers, J. Cavender‐Bares, et al. 2004. “The Worldwide Leaf Economics Spectrum.” Nature 428: 821–827.

Wurzburger, N., and S. J. Wright. 2015. “Fine‐Root Responses to Fertilization Reveal Multiple Nutrient Limitation in a Lowland Tropical Forest.” Ecology 96: 2137–2146.

Xing, J., J. Pleim, R. Mathur, G. Pouliot, C. Hogrefe, C. M. Gan, and C. Wei. 2013. “Historical Gaseous and Primary Aerosol Emissions in the United States from 1990 to 2010.” Atmospheric Chemistry and Physics 13: 7531–7549.

Zelený, D. 2018. “Which Results of the Standard Test for Community‐Weighted Mean Approach Are Too Optimistic?” Journal of Vegetation Science 29: 953–966.

Zuur, A. F., E. N. Ieno, and C. S. Elphick. 2010. “A Protocol for Data Exploration to Avoid Common Statistical Problems.” Methods in Ecology and Evolution 1: 3–14.