Drought tolerance modulates the effects of neighborhood diversity and competition on tree growth in a tropical forest

Abstract

Neighborhood diversity and competition are key drivers of tree growth in tropical forests, but their responses to drought remain uncertain. It is also unclear whether species-level drought tolerance regulates the strength of these neighborhood effects. Using 33 years of individual-level census data from the 50-ha ForestGEO plot on Barro Colorado Island (BCI), Panama, we quantified the effects of neighborhood species richness (NSR) and neighborhood competition index (NCI) on tree basal area growth across seven census intervals from 1982 to 2015. To distinguish long-term and short-term competitive effects, NCI was decomposed into a between-individual component representing each tree’s long-term mean competitive environment and a within-individual component representing temporal deviations from that mean. We used linear mixed-effects models to test whether climatic water balance, represented by the standardized precipitation-evapotranspiration index (SPEI), and focal-tree drought tolerance, represented by xylem water potential at 50% loss of hydraulic conductivity (P50), modified these effects. Among long-term surviving focal trees, NSR had a significant positive effect on basal area growth rate, although sensitivity analyses including pre-mortality observations suggest that this diversity benefit is conditional on individual tree vigor. However, both NCI components significantly reduced growth, with the between-individual component exhibiting a substantially stronger negative effect than the within-individual component. Climatic water balance modified persistent and temporal components of neighborhood competition in contrasting ways: long-term competitive environments exhibited a statistically significant but modest increase in competitive suppression under dry conditions, whereas temporal fluctuations in competition had stronger growth consequences under wetter conditions. In contrast, drought did not significantly modify the positive effect of NSR, suggesting that competitive suppression rather than diversity-mediated complementarity dominated tree growth under water-limited conditions. Among the 29 species with available P50 data, which represented 39.4% of the focal-tree observations, drought tolerance significantly modulated the neighborhood effects. Drought-sensitive species, with less negative P50 values, showed stronger positive responses to NSR. However, they also suffered greater growth suppression from both competition components. Drought-tolerant species, with more negative P50 values, responded weaklier overall. Whether this pattern holds for the broader BCI community remains untested, particularly for rarer and less-studied species. These findings show that long-term competitive environment and short-term competitive fluctuations influence tree growth differently depending on moisture conditions, and that hydraulic traits mediate how trees respond to neighborhood interactions. Incorporating hydraulic traits and decomposed competition effects into neighborhood models could be a promising direction for future work.

Keywords

Neighborhood competition / Neighborhood species richness / Tree growth / Hydraulic traits / Drought tolerance / SPEI / Tropical forest

References

[1] Bagchi R, Swinfield T, Gallery RE, Lewis OT, Gripenberg S, Narayan L, Freckleton RP. Testing the Janzen-Connell mechanism: pathogens cause overcompensating density dependence in a tropical tree. Ecol Lett, 2010, 131012621269.

[2] Bates D, Mächler M, Bolker B, Walker S. Fitting linear mixed-effects models using lme4. J Stat Softw, 2015, 671148.

[3] Bauman D, Fortunel C, Cernusak LA, Bentley LP, McMahon SM, Rifai SW, Aguirre-Gutiérrez J, Oliveras I, Bradford M, Laurance SGW, Delhaye G, Hutchinson MF, Dempsey R, McNellis BE, Santos-Andrade PE, Ninantay-Rivera HR, Chambi Paucar JR, Phillips OL, Malhi Y. Tropical tree growth sensitivity to climate is driven by species intrinsic growth rate and leaf traits. Glob Change Biol, 2022, 28414141432.

[4] Beauchamp N, Kunstler G, Touzot L, Ruiz-Benito P, Cienciala E, Dahlgren J, Hawryło P, Klopčič M, Lehtonen A, Šebeň V, Socha J, Zavala MA, Courbaud B. Light competition affects how tree growth and survival respond to climate. J Ecol, 2025, 1133672688.

[5] Beguería S (2017) sbegueria/SPEIbase: Version 2.5.1. 10.5281/ZENODO.834462

[6] Behroozian M. Partial ecological niche differentiation suggests incipient ecological divergence in Dianthus polylepis subspecies. Evol Ecol, 2025, 395633656.

[7] Bell A, Jones K. Explaining fixed effects: random effects modeling of time-series cross-sectional and panel data. Polit Sci Res Methods, 2015, 31133153.

[8] Canham CD, LePage PT, Coates KD. A neighborhood analysis of canopy tree competition: effects of shading versus crowding. Can J for Res, 2004, 344778787.

[9] Castagneri D, Vacchiano G, Hacket-Pain A, DeRose RJ, Klein T, Bottero A. Meta-analysis reveals different competition effects on tree growth resistance and resilience to drought. Ecosystems, 2022, 2513043.

[10] Chen YX, Uriarte M, Wright SJ, Yu SX. Effects of neighborhood trait composition on tree survival differ between drought and postdrought periods. Ecology, 2019, 1009e02766

[11] Chin ARO, Hille Ris Lambers J, Franklin JF. Context matters: natural tree mortality can lead to neighbor growth release or suppression. For Ecol Manag, 2023, 529. 120735

[12] Choat B, Jansen S, Brodribb TJ, Cochard H, Delzon S, Bhaskar R, Bucci SJ, Feild TS, Gleason SM, Hacke UG, Jacobsen AL, Lens F, Maherali H, Martínez-Vilalta J, Mayr S, Mencuccini M, Mitchell PJ, Nardini A, Pittermann J, Pratt RB, Sperry JS, Westoby M, Wright IJ, Zanne AE. Global convergence in the vulnerability of forests to drought. Nature, 2012, 4917426752755.

[13] Cinoğlu D, Rüger N, Decker RR, Farrior CE. Small disturbances and subsequent competition for light can maintain a diversity of demographic strategies in a neotropical forest: results from model–data integration. J Ecol, 2025, 113926252638.

[14] Comita LS, Muller-Landau HC, Aguilar S, Hubbell SP. Asymmetric density dependence shapes species abundances in a tropical tree community. Science, 2010, 3295989330332.

[15] Comita LS, Aguilar S, Hubbell SP, Pérez R. Long-term seedling and small sapling census data from the Barro Colorado Island 50 ha Forest Dynamics Plot, Panama. Ecology, 2023, 1049e4140

[16] Condit R. Tropical forest census plots: methods and results from Barro Colorado Island, Panama and a comparison with other plots, 1998, Berlin: Springer-Verlag

[17] Condit R, Pérez R, Aguilar S, Lao S, Foster R, Hubbell S (2019) Complete data from the Barro Colorado 50-ha plot: 423617 trees, 35 years, 2019 version. https://doi.org/10.15146/5xcp-0d46

[18] Curzon MT, D’Amato AW, Fraver S, Huff ES, Palik BJ. Succession, climate and neighbourhood dynamics influence tree growth over time: an 87-year record of change in a Pinus resinosa-dominated forest, Minnesota, USA. J Veg Sci, 2017, 2818292.

[19] Durgante FM, Higuchi N, Ohashi S, Householder JE, Lima AJN, Ishizuka M, Trumbore S. Soil fertility and drought interact to determine large variations in wood production for a hyperdominant Amazonian tree species. Frontiers Forests Global Change, 2023, 5: 1065645.

[20] Enders CK, Tofighi D. Centering predictor variables in cross-sectional multilevel models: a new look at an old issue. Psychol Methods, 2007, 122121138.

[21] Fichtner A, Schnabel F, Bruelheide H, Kunz M, Mausolf K, Schuldt A, Härdtle W, von Oheimb G. Neighbourhood diversity mitigates drought impacts on tree growth. J Ecol, 2020, 1083865875.

[22] Fox J, Weisberg S. An R companion to applied regression, 20193rd ednThousand OaksSage

[23] Georgi L, Kunz M, Fichtner A, Reich KF, Bienert A, Maas HG, von Oheimb G. Effects of local neighbourhood diversity on crown structure and productivity of individual trees in mature mixed-species forests. For Ecosyst, 2021, 8. 26

[24] Gilbert GS. Evolutionary ecology of plant diseases in natural ecosystems. Annu Rev Phytopathol, 2002, 40: 13-43.

[25] Gómez-Aparicio L, Domínguez-Begines J, Kardol P, Ávila JM, Ibáñez B, García LV. Plant-soil feedbacks in declining forests: implications for species coexistence. Ecology, 2017, 98719081921.

[26] Grossiord C. Having the right neighbors: how tree species diversity modulates drought impacts on forests. New Phytol, 2020, 22814249.

[27] Guillemot J, Kunz M, Schnabel F, Fichtner A, Madsen CP, Gebauer T, Härdtle W, von Oheimb G, Potvin C. Neighbourhood-mediated shifts in tree biomass allocation drive overyielding in tropical species mixtures. New Phytol, 2020, 228412561268.

[28] Haberstroh S, Werner C. The role of species interactions for forest resilience to drought. Plant Biol, 2022, 24710981107.

[29] Harris I, Jones PD, Osborn TJ, Lister DH. Updated high-resolution grids of monthly climatic observations–the CRU TS3.10 dataset. Int J Climatol, 2014, 343623642.

[30] He Q, Bertness MD. Extreme stresses, niches, and positive species interactions along stress gradients. Ecology, 2014, 95614371443.

[31] Hegyi F. A simulation model for managing jack-pine stands. Growth models for tree and stand simulation, 1974, Stockholm: Royal College of Forestry, 74-90

[32] Huang YY, Chen YX, Castro-Izaguirre N, Baruffol M, Brezzi M, Lang A, Li Y, Härdtle W, von Oheimb G, Yang XF, Liu XJ, Pei KQ, Both S, Yang B, Eichenberg D, Assmann T, Bauhus J, Behrens T, Buscot F, Chen XY, Chesters D, Ding BY, Durka W, Erfmeier A, Fang JY, Fischer M, Guo LD, Guo DL, Gutknecht JLM, He JS, He CL, Hector A, Hönig L, Hu RY, Klein AM, Kühn P, Liang Y, Li S, Michalski S, Scherer-Lorenzen M, Schmidt K, Scholten T, Schuldt A, Shi XZ, Tan MZ, Tang ZY, Trogisch S, Wang ZW, Welk E, Wirth C, Wubet T, Xiang WH, Yu MJ, Yu XD, Zhang JY, Zhang SR, Zhang NL, Zhou HZ, Zhu CD, Zhu L, Bruelheide H, Ma KP, Niklaus PA, Schmid B. Impacts of species richness on productivity in a large-scale subtropical forest experiment. Science, 2018, 36264108083.

[33] Hubbell SP, Foster RB. Diversity of canopy trees in a neotropical forest and implications for conservation. Tropical rain forest: ecology and management, 1983, Oxford: Blackwell Scientific Publications, 25-41

[34] Hubbell SP, Ahumada JA, Condit R, Foster RB. Local neighborhood effects on long-term survival of individual trees in a neotropical forest. Ecol Res, 2001, 165859875.

[35] Isbell F, Craven D, Connolly J, Loreau M, Schmid B, Beierkuhnlein C, Bezemer TM, Bonin C, Bruelheide H, de Luca E, Ebeling A, Griffin JN, Guo QF, Hautier Y, Hector A, Jentsch A, Kreyling J, Lanta V, Manning P, Meyer ST, Mori AS, Naeem S, Niklaus PA, Polley HW, Reich PB, Roscher C, Seabloom EW, Smith MD, Thakur MP, Tilman D, Tracy BF, van der Putten WH, van Ruijven J, Weigelt A, Weisser WW, Wilsey B, Eisenhauer N. Biodiversity increases the resistance of ecosystem productivity to climate extremes. Nature, 2015, 5267574574577.

[36] Jansen K, von Oheimb G, Bruelheide H, Härdtle W, Fichtner A. Tree species richness modulates water supply in the local tree neighbourhood: evidence from wood δ¹³C signatures in a large-scale forest experiment. Proc R Soc B, 2021, 288194620203100

[37] Jiang P, Liu HY, Piao SL, Ciais P, Wu XC, Yin Y, Wang HY. Enhanced growth after extreme wetness compensates for post-drought carbon loss in dry forests. Nat Commun, 2019, 10: 195.

[38] John R, Dalling JW, Harms KE, Yavitt JB, Stallard RF, Mirabello M, Hubbell SP, Valencia R, Navarrete H, Vallejo M, Foster RB. Soil nutrients influence spatial distributions of tropical tree species. Proc Natl Acad Sci USA, 2007, 1043864869.

[39] Kunstler G, Falster D, Coomes DA, Hui F, Kooyman RM, Laughlin DC, Poorter L, Vanderwel M, Vieilledent G, Wright SJ, Aiba M, Baraloto C, Caspersen J, Cornelissen JHC, Gourlet-Fleury S, Hanewinkel M, Herault B, Kattge J, Kurokawa H, Onoda Y, Peñuelas J, Poorter H, Uriarte M, Richardson S, Ruiz-Benito P, Sun IF, Ståhl G, Swenson NG, Thompson J, Westerlund B, Wirth C, Zavala MA, Zeng HC, Zimmerman JK, Zimmermann NE, Westoby M. Plant functional traits have globally consistent effects on competition. Nature, 2016, 5297585204207.

[40] Kuznetsova A, Brockhoff PB, Christensen RHB. lmerTest Package: tests in linear mixed effects models. J Stat Softw, 2017, 8213126.

[41] Lebrija-Trejos E, Hernández A, Wright SJ. Effects of moisture and density-dependent interactions on tropical tree diversity. Nature, 2023, 6157950100104.

[42] Liang Y, Duveneck MJ, Gustafson EJ, Serra-Diaz JM, Thompson JR. How disturbance, competition, and dispersal interact to prevent tree range boundaries from keeping pace with climate change. Glob Change Biol, 2018, 241e335e351.

[43] Liang JJ, Crowther TW, Picard N, Wiser S, Zhou M, Alberti G, Schulze ED, McGuire AD, Bozzato F, Pretzsch H, de-Miguel S, Paquette A, Hérault B, Scherer-Lorenzen M, Barrett CB, Glick HB, Hengeveld GM, Nabuurs GJ, Pfautsch S, Viana H, Vibrans AC, Ammer C, Schall P, Verbyla D, Tchebakova N, Fischer M, Watson JV, Chen HYH, Lei XD, Schelhaas MJ, Lu HC, Gianelle D, Parfenova EI, Salas C, Lee E, Lee B, Kim HS, Bruelheide H, Coomes DA, Piotto D, Sunderland T, Schmid B, Gourlet-Fleury S, Sonké B, Tavani R, Zhu J, Brandl S, Vayreda J, Kitahara F, Searle EB, Neldner VJ, Ngugi MR, Baraloto C, Frizzera L, Bałazy R, Oleksyn J, Zawiła-Niedźwiecki T, Bouriaud O, Bussotti F, Finér L, Jaroszewicz B, Jucker T, Valladares F, Jagodzinski AM, Peri PL, Gonmadje C, Marthy W, O’Brien T, Martin EH, Marshall AR, Rovero F, Bitariho R, Niklaus PA, Alvarez-Loayza P, Chamuya N, Valencia R, Mortier F, Wortel V, Engone-Obiang NL, Ferreira LV, Odeke DE, Vasquez RM, Lewis SL, Reich PB (2016) Positive biodiversity-productivity relationship predominant in global forests. Science 354(6309): aaf8957. https://doi.org/10.1126/science.aaf8957

[44] Liu S, McVicar TR, Wu X, Cao X, Liu Y. Assessing the relative importance of dry-season incoming solar radiation and water storage dynamics during the 2005, 2010 and 2015 southern Amazon droughts: not all droughts are created equal. Environ Res Lett, 2024, 193034027

[45] Lu KL, Chen N, Zhang CK, Dong XX, Zhao CM. Drought enhances the role of competition in mediating the relationship between tree growth and climate in semi-arid areas of northwest China. Forests, 2019, 109804.

[46] Lüdecke D, Ben-Shachar MS, Patil I, Waggoner P, Makowski D. Performance: an R package for assessment, comparison and testing of statistical models. J Open Source Softw, 2021, 6603139

[47] Maestre FT, Martínez I, Escolar C, Escudero A. On the relationship between abiotic stress and co-occurrence patterns: an assessment at the community level using soil lichen communities and multiple stress gradients. Oikos, 2009, 118710151022.

[48] Magalhães JGDS, Amoroso MM, Larson BC. What evidence exists on the effects of competition on trees’ responses to climate change? Syst Map Protocol. Environ Evid, 2021, 10134.

[49] Meinzer FC, Johnson DM, Lachenbruch B, McCulloh KA, Woodruff DR. Xylem hydraulic safety margins in woody plants: coordination of stomatal control of xylem tension with hydraulic capacitance. Funct Ecol, 2009, 235922930.

[50] Mello FNA, Estrada-Villegas S, DeFilippis DM, Schnitzer SA. Can functional traits explain plant coexistence? A case study with tropical lianas and trees. Diversity, 2020, 1210397.

[51] Nemetschek D, Derroire G, Marcon E, Aubry-Kientz M, Auer J, Badouard V, Baraloto C, Bauman D, Le Blaye Q, Boisseaux M, Bonal D, Coste S, Dardevet E, Heuret P, Hietz P, Levionnois S, Maréchaux I, McMahon SM, Stahl C, Vleminckx J, Wanek W, Ziegler C, Fortunel C. Climate anomalies and neighbourhood crowding interact in shaping tree growth in old-growth and selectively logged tropical forests. J Ecol, 2024, 1123590612.

[52] Nemetschek D, Fortunel C, Marcon E, Auer J, Badouard V, Baraloto C, Boisseaux M, Bonal D, Coste S, Dardevet E, Heuret P, Hietz P, Levionnois S, Maréchaux I, Stahl C, Vleminckx J, Wanek W, Ziegler C, Derroire G. Love thy neighbour? Tropical tree growth and its response to climate anomalies is mediated by neighbourhood hierarchy and dissimilarity in carbon- and water-related traits. Ecol Lett, 2025, 284e70028

[53] Pretzsch H, del Río M, Grote R, Klemmt HJ, Ordóñez C, Oviedo FB. Tracing drought effects from the tree to the stand growth in temperate and Mediterranean forests: insights and consequences for forest ecology and management. Eur J for Res, 2022, 1414727751.

[54] Pustejovsky JE, Tipton E. Small-sample methods for cluster-robust variance estimation and hypothesis testing in fixed effects models. J Bus Econ Stat, 2018, 364672683.

[55] Pustejovsky JE (2022) clubSandwich: cluster-robust (sandwich) variance estimators with small-sample corrections. R package version 0.5.8. https://CRAN.R-project.org/package=clubSandwich

[56] Rao AS, Chhawri R, Chauhan A, Yadav SS, Meena KC, Bansal P. Plant functional traits: a key framework for understanding and managing ecosystem responses to global environmental challenges. Plant functional traits for improving productivity, 2024Springer Nature Singapore287299.

[57] Rowland L, da Costa ACL, Oliveira RS, Bittencourt PRL, Giles AL, Coughlin I, de Britto Costa P, Bartholomew D, Domingues TF, Miatto RC, Ferreira LV, Vasconcelos SS, Junior JAS, Oliveira AAR, Mencuccini M, Meir P. The response of carbon assimilation and storage to long-term drought in tropical trees is dependent on light availability. Funct Ecol, 2021, 3514353.

[58] Rozendaal DMA, Phillips OL, Lewis SL, Affum-Baffoe K, Alvarez-Davila E, Andrade A, Aragão LEOC, Araujo-Murakami A, Baker TR, Bánki O, Brienen RJW, Camargo JLC, Comiskey JA, Djuikouo Kamdem MN, Fauset S, Feldpausch TR, Killeen TJ, Laurance WF, Laurance SGW, Lovejoy T, Malhi Y, Marimon BS, Marimon BHJrMarshall AR, Neill DA, Núñez Vargas P, Pitman NCA, Poorter L, Reitsma J, Silveira M, Sonké B, Sunderland T, Taedoumg H, ter Steege H, Terborgh JW, Umetsu RK, van der Heijden GMF, Vilanova E, Vos V, White LJT, Willcock S, Zemagho L, Vanderwel MC. Competition influences tree growth, but not mortality, across environmental gradients in Amazonia and tropical Africa. Ecology, 2020, 1017e03052

[59] Rüger N, Berger U, Hubbell SP, Vieilledent G, Condit R. Growth strategies of tropical tree species: disentangling light and size effects. PLoS ONE, 2011, 69e25330

[60] Sachsenmaier L, Schnabel F, Dietrich P, Eisenhauer N, Ferlian O, Quosh J, Richter R, Wirth C. Forest growth resistance and resilience to the 2018–2020 drought depend on tree diversity and mycorrhizal type. J Ecol, 2024, 112817871803.

[61] Sanchez-Martinez P, Mencuccini M, García-Valdés R, Hammond WM, Serra-Diaz JM, Guo WY, Segovia RA, Dexter KG, Svenning JC, Allen C, Martínez-Vilalta J. Increased hydraulic risk in assemblages of woody plant species predicts spatial patterns of drought-induced mortality. Nat Ecol Evol, 2023, 71016201632.

[62] Schielzeth H, Dingemanse NJ, Nakagawa S, Westneat DF, Allegue H, Teplitsky C, Réale D, Dochtermann NA, Garamszegi LZ, Araya-Ajoy YG. Robustness of linear mixed-effects models to violations of distributional assumptions. Methods Ecol Evol, 2020, 11911411152.

[63] Schnabel F, Barry KE, Eckhardt S, Guillemot J, Geilmann H, Kahl A, Moossen H, Bauhus J, Wirth C. Neighbourhood species richness and drought-tolerance traits modulate tree growth and δ¹³C responses to drought. Plant Biol, 2024, 262330345.

[64] Schwinning S, Weiner J. Mechanisms determining the degree of size asymmetry in competition among plants. Oecologia, 1998, 1134447455.

[65] Serrano-León H, Blondeel H, Glenz P, Steurer J, Schnabel F, Baeten L, Guillemot J, Martin-StPaul N, Skiadaresis G, Scherer-Lorenzen M, Bonal D, Boone M, Decarsin R, Druel A, Godbold DL, Gong JL, Hajek P, Jactel H, Koricheva J, Mereu S, Ponette Q, Rewald B, Sandén H, Van den Bulcke J, Verheyen K, Werner R, Bauhus J. Multiyear drought strengthens positive and negative functional diversity effects on tree growth response. Glob Change Biol, 2025, 319e70394

[66] Sha SL, Cai GC, Liu SR, Ali Ahmed M. Roots to the rescue: how plants harness hydraulic redistribution to survive drought across contrasting soil textures. Adv Biotechnol, 2024, 2443.

[67] Smith-Martin CM, Muscarella R, Ankori-Karlinsky R, Delzon S, Farrar SL, Salva-Sauri M, Thompson J, Zimmerman JK, Uriarte M. Hurricanes increase tropical forest vulnerability to drought. New Phytol, 2022, 235310051017.

[68] Uriarte M, Canham CD, Thompson J, Zimmerman JK. A neighborhood analysis of tree growth and survival in a hurricane-driven tropical forest. Ecol Monogr, 2004, 744591614.

[69] Velázquez E, Wiegand T. Competition for light and persistence of rare light-demanding species within tree-fall gaps in a moist tropical forest. Ecology, 2020, 1017e03034

[70] Vicente-Serrano SM, Beguería S, López-Moreno JI. A multiscalar drought index sensitive to global warming: the standardized precipitation evapotranspiration index. J Clim, 2010, 23716961718.

[71] Watson JV, Liang JJ, Tobin PC, Lei XD, Rentch JS, Artis CE. Large-scale forest inventories of the United States and China reveal positive effects of biodiversity on productivity. For Ecosyst, 2015, 2: 22.

[72] Weiner J. Asymmetric competition in plant populations. Trends Ecol Evol, 1990, 511360364.

[73] Weng XX, Guo YP, Tang ZY. Spatial-temporal dependence of the neighborhood interaction in regulating tree growth in a tropical rainforest. For Ecol Manag, 2022, 508. 120032

[74] Xu H, Detto M, Hogan JA, Alonso A, Birch JD, Bissiengou P, Chu CJ, Davies SJ, Fischer GA, Hau BCH, Kenfack D, Li BH, Lian JY, Lin MX, Liu WD, Liu Y, Liu ZF, Lutz JA, Memiaghe HR, Mi XC, Novotny V, Ren HB, Su JR, Thompson J, Uriarte M, Valencia R, Yao TL, Yap SL, Zhang YC, Zimmerman JK, Weiblen GD, Li YD, Fang SQ, He FL. The importance of competition and facilitation for global tree diversity. Nature, 2026, 653811610851091.

[75] Yeo I, Johnson RA. A new family of power transformations to improve normality or symmetry. Biometrika, 2000, 874954959.

[76] Yi XX, Wang NN, Ren HB, Yu JP, Hu TY, Su YJ, Mi XC, Guo QH, Ma KP. From canopy complementarity to asymmetric competition: the negative relationship between structural diversity and productivity during succession. J Ecol, 2022, 1102457465.

[77] Zambrano J, Beckman NG, Marchand P, Thompson J, Uriarte M, Zimmerman JK, Umaña MN, Swenson NG. The scale dependency of trait-based tree neighborhood models. J Veg Sci, 2020, 314581593.

[78] Zhang Y, Zheng QZ, Gao XX, Ma YD, Liang KM, Yue HT, Huang XX, Wu KT, Wang XR. Land degradation changes the role of above- and belowground competition in regulating plant biomass allocation in an Alpine meadow. Front Plant Sci, 2022, 13. 822594

[79] Zhang HN, Gao YY, Zheng X, Hu YP, Zhou X, Fang YM, Li Y, Xie L, Ding H. Neighborhood diversity promotes tree growth in a secondary forest: the interplay of intraspecific competition, interspecific competition, and spatial scale. Plants (Basel), 2024, 13141994

[80] Zhang HT, Gheyret G, Bai YH, Guo YP, Li S, Schmid B, Bruelheide H, Ma KP, Tang ZY. Functional dissimilarity in mixed forests promotes stem radial growth by mitigating tree water deficit. Natl Sci Rev, 2024, 113nwad320

[81] Zhang HT, Ovaskainen O, Chi XL, Guo Q, Tang ZY. Spatiotemporal variation in the negative effect of neighbourhood crowding on stem growth. J Ecol, 2024, 112511401149.

[82] Zheng LT, Ibáñez I, Williams LJ, Zhu K, Serrano-León H, Jensen J, Eisenhauer N, Verheyen K, Scherer-Lorenzen M, Schnabel F, Kreft H, Guerrero-Ramírez NR, Hölscher D, Paterno GB, Irawan B, Ponette Q, Messier C, Paquette A, Stefanski A, Mereu S, Bauhus J, Hajek P, Nock CA, Cavender-Bares J, Parker WC, Quosh J, Ferlian O, Auge H, Potvin C, Yan ER, Yang BY, Zhang L, Zhao Z, Sinacore K, Hall JS, Guillemot J, Robin A, Brancalion PHS, Sundawati L, Reich PB. Neighbourhood diversity increases tree growth in experimental forests more in wetter climates but not in wetter years. Nat Ecol Evol, 2025, 91018121824.

[83] Zuur AF, Ieno EN, Elphick CS. A protocol for data exploration to avoid common statistical problems. Methods Ecol Evol, 2010, 11314.

Funding

National Natural Science Foundation(32201306)

Xinjiang Normal University Young Talent Program(XJNUQB2024-13)

Rights & permissions

Northeast Forestry University