Functional basis of architectural differentiation among co-occurring tree species in a tropical seasonal rainforest: allometric intercepts and ontogenetic shifts

Abstract

Tree architecture shapes forest structure and function. Functional traits reflecting ecological strategies may influence allometric relationships between architectural parameters across species and life stages, which remains poorly understood. We measured architectural parameters (tree height, crown width, crown depth) for 1309 individuals of 41 co-occurring tree species in a 20-ha tropical seasonal rainforest dynamics plot in Xishuangbanna, southwest China. Using Bayesian hierarchical models, we quantified species-specific allometric relationships, with which we further tested how functional traits covering mechanical, life-history, and leaf-economic dimensions correlated and how these correlations varied across ontogeny. Interspecific variation in tree architecture was primarily driven by allometric intercepts rather than slopes. Species with higher adult stature (H_max) and crown exposure developed slenderer stature but narrower, shallower crowns at given sizes. While denser wood consistently supported greater height and deeper crowns across ontogeny, the correlation between traits and crown dimensions tended to weaken from the juvenile to the adult stage. Our results uncover a trade-off between height growth and crown expansion, and reveal that ecological strategy may depend on life stages. Overall, the relationships between functional traits and tree architecture reveal multiple dimensions of species differentiation arising from variation in architectural and leaf-economic strategies among co-occurring tree species in tropical seasonal rainforests.

Keywords

Architectural allometry / Architectural strategy / Bayesian hierarchical model / Crown exposure / Niche differentiation

References

[1] Abràmoff MD, Magalhães PJ, Ram SJ. Image processing with ImageJ. Biophotonics Int, 2004, 11(7): 36-42

[2] Aiba M, Nakashizuka T. Architectural differences associated with adult stature and wood density in 30 temperate tree species. Funct Ecol, 2009, 23(2): 265-273

[3] Bartlett MK, Zhang Y, Yang J, Kreidler N, Sun SW, Lin L, Hu YH, Cao KF, Sack L. Drought tolerance as a driver of tropical forest assembly: resolving spatial signatures for multiple processes. Ecology, 2016, 97(2): 503-514

[4] Bohlman S, O’Brien S. Allometry, adult stature and regeneration requirement of 65 tree species on barro Colorado Island. Panama J Trop Ecol, 2006, 22(2): 123-136

[5] Bürkner PC. brms: an R package for Bayesian multilevel models using Stan. J Stat Soft, 2017, 80(1): 1-28

[6] Cao M, Zhu H, Wang H, Lan G, Hu Y, Zhou S, Deng X, Cui J. Xishuangbanna tropical seasonal rainforest dynamics plot: tree distribution maps, diameter tables and species documentation, 2008, Kunming, Yunnan Science and Technology Press

[7] Carpenter B, Gelman A, Hoffman MD, Lee D, Goodrich B, Betancourt M, Brubaker M, Guo JQ, Li P, Riddell A. Stan: a probabilistic programming language. J Stat Soft, 2017, 76(1): 1-32

[8] Chave J, Muller-Landau HC, Baker TR, Easdale TA, ter Steege H, Webb CO. Regional and phylogenetic variation of wood density across 2456 neotropical tree species. Ecol Appl, 2006, 16(6): 2356-2367

[9] Chave J, Réjou-Méchain M, Búrquez A, Chidumayo E, Colgan MS, Delitti WBC, Duque A, Eid T, Fearnside PM, Goodman RC, Henry M, Martínez-Yrízar A, Mugasha WA, Muller-Landau HC, Mencuccini M, Nelson BW, Ngomanda A, Nogueira EM, Ortiz-Malavassi O, Pélissier R, Ploton P, Ryan CM, Saldarriaga JG, Vieilledent G. Improved allometric models to estimate the aboveground biomass of tropical trees. Glob Change Biol, 2014, 20(10): 3177-3190

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

[11] Cornelissen JHC, Lavorel S, Garnier E, Díaz S, Buchmann N, Gurvich DE, Reich PB, ter Steege H, Morgan HD, van der Heijden MGA, Pausas JG, Poorter H. A handbook of protocols for standardised and easy measurement of plant functional traits worldwide. Aust J Bot, 2003, 51(4): 335-380

[12] Díaz S, Kattge J, Cornelissen JHC, Wright IJ, Lavorel S, Dray S, Reu B, Kleyer M, Wirth C, Prentice IC, Garnier E, Bönisch G, Westoby M, Poorter H, Reich PB, Moles AT, Dickie J, Gillison AN, Zanne AE, Chave J, Wright SJ, Sheremet'ev SN, Jactel H, Baraloto C, Cerabolini B, Pierce S, Shipley B, Kirkup D, Casanoves F, Joswig JS, Günther A, Falczuk V, Rüger N, Mahecha MD, Gorné LD. The global spectrum of plant form and function. Nature, 2016, 529(7585): 167-171

[13] Dietze MC, Wolosin MS, Clark JS. Capturing diversity and interspecific variability in allometries: a hierarchical approach. For Ecol Manag, 2008, 256(11): 1939-1948

[14] Dong SX. The structure and dynamics of tropical forests in relation to climate variability (Doctoral dissertation), 2011, Cambridge, Harvard University

[15] Falster DS, Brännström Å, Westoby M, Dieckmann U. Multitrait successional forest dynamics enable diverse competitive coexistence. Proc Natl Acad Sci U S A, 2017, 114(13): E2719-E2728

[16] Gelman A, Carlin JB, Stern HS, Dunson DB, Vehtari A, Rubin DB. Bayesian data analysis, 20133rd ednBoca Raton, Chapman & Hall/CRC Press

[17] Hardiman B, Bohrer G, Gough C, Curtis P. Canopy structural changes following widespread mortality of canopy dominant trees. Forests, 2013, 4(3): 537-552

[18] Iida Y, Kohyama TS, Kubo T, Kassim AR, Poorter L, Sterck F, Potts MD. Tree architecture and life-history strategies across 200 co-occurring tropical tree species. Funct Ecol, 2011, 25(6): 1260-1268

[19] Iida Y, Poorter L, Sterck FJ, Kassim AR, Kubo T, Potts MD, Kohyama TS. Wood density explains architectural differentiation across 145 co-occurring tropical tree species. Funct Ecol, 2012, 26(1): 274-282

[20] Jucker T, Fischer FJ, Chave J, Coomes DA, Caspersen J, Ali A, Loubota Panzou GJ, Feldpausch TR, Falster D, Usoltsev VA, Jackson TD, Adu-Bredu S, Alves LF, Aminpour M, Angoboy Ilondea B, Anten NPR, Antin C, Askari Y, Ayyappan N, Banin LF, Barbier N, Battles JJ, Beeckman H, Bocko YE, Bond-Lamberty B, Bongers F, Bowers S, van Breugel M, Chantrain A, Chaudhary R, Dai JY, Dalponte M, Dimobe K, Domec JC, Doucet JL, Dupuy Rada JM, Duursma RA, Enríquez M, van Ewijk KY, Farfán-Rios W, Fayolle A, Ferretti M, Forni E, Forrester DI, Gilani H, Godlee JL, Haeni M, Hall JS, He JK, Hemp A, Hernández-Stefanoni JL, Higgins SI, Holdaway RJ, Hussain K, Hutley LB, Ichie T, Iida Y, Jiang HS, Joshi PR, Kaboli H, Kazempour Larsary M, Kenzo T, Kloeppel BD, Kohyama TS, Kunwar S, Kuyah S, Kvasnica J, Lin SL, Lines ER, Liu HY, Lorimer C, Loumeto JJ, Malhi Y, Marshall PL, Mattsson E, Matula R, Meave JA, Mensah S, Mi XC, Momo ST, Moncrieff GR, Mora F, Muñoz R, Nissanka SP, Nur Hajar ZS, O’Hara KL, Pearce S, Pelissier R, Peri PL, Ploton P, Poorter L, Pour MJ, Pourbabaei H, Ribeiro SC, Ryan C, Sanaei A, Sanger J, Schlund M, Sellan G, Shenkin A, Sonké B, Sterck FJ, Svátek M, Takagi K, Trugman AT, Vadeboncoeur MA, Valipour A, Vanderwel MC, Vovides AG, Waldner P, Wang WW, Wang LQ, Wirth C, Woods M, Xiang WH, de Aquino XF, Xu YZ, Yamada T, Zavala MA, Zimmermann NE. The global spectrum of tree crown architecture. Nat Commun, 2025, 16: 4876

[21] Kassambara A, Mundt F (2020) factoextra: extract and visualize the results of multivariate data analyses, version 1.0.7. CRAN. https://CRAN.R-project.org/package=factoextra

[22] King DA, Davies SJ, Tan S, Noor NSM. The role of wood density and stem support costs in the growth and mortality of tropical trees. J Ecol, 2006, 94(3): 670-680

[23] Kitajima K, Poorter L. Carson WP, Schnitzer SA. Functional basis for resource niche partitioning by tropical trees. Tropical forest community ecology, 2008Wiley-Blackwell: 160-181

[24] Kohyama T. Significance of architecture and allometry in saplings. Funct Ecol, 1987, 1(4): 399-404

[25] Kohyama T, Suzuki E, Partomihardjo T, Yamada T, Kubo T. Tree species differentiation in growth, recruitment and allometry in relation to maximum height in a Bornean mixed dipterocarp forest. J Ecol, 2003, 91(5): 797-806

[26] Lê S, Josse J, Husson F. FactoMineR: an R package for multivariate analysis. J Stat Soft, 2008, 25(1): 1-18

[27] Leishman MR, Westoby M. The role of seed size in seedling establishment in dry soil conditions—experimental evidence from semi-arid species. J Ecol, 1994, 82(2): 249-258

[28] Loubota Panzou GJ, Ligot G, Gourlet-Fleury S, Doucet JL, Forni E, Loumeto JJ, Fayolle A. Architectural differences associated with functional traits among 45 coexisting tree species in Central Africa. Funct Ecol, 2018, 32(11): 2583-2593

[29] Martínez Cano I, Muller-Landau HC, Wright SJ, Bohlman SA, Pacala SW. Tropical tree height and crown allometries for the Barro Colorado Nature Monument, Panama: a comparison of alternative hierarchical models incorporating interspecific variation in relation to life history traits. Biogeosciences, 2019, 16(4): 847-862

[30] Matsuo T, Hiura T, Onoda Y. Vertical and horizontal light heterogeneity along gradients of secondary succession in cool- and warm-temperate forests. J Veg Sci, 2022, 33(3 e13135

[31] McMahon T. Size and shape in biology: elastic criteria impose limits on biological proportions, and consequently on metabolic rates. Science, 1973, 1794079): 1201-1204

[32] Moles AT, Westoby M. Seedling survival and seed size: a synthesis of the literature. J Ecol, 2004, 92(3): 372-383

[33] Moravie MA, Durand M, Houllier F. Ecological meaning and predictive ability of social status, vigour and competition indices in a tropical rain forest (India). For Ecol Manag, 1999, 117(1–3): 221-240

[34] Niklas KJ. Plant biomechanics: an engineering approach to plant form and function, 1992, Chicago, University of Chicago Press

[35] Niklas KJ. Plant allometry: the scaling of form and process, 1994, Chicago, University of Chicago Press

[36] Poorter L, Bongers F, Sterck FJ, Wöll H. Architecture of 53 rain forest tree species differing in adult stature and shade tolerance. Ecology, 2003, 84(3): 602-608

[37] Poorter L, Bongers F, Sterck FJ. Beyond the regeneration phase: differentiation of height-light trajectories among tropical tree species. J Ecol, 2005, 93(2): 256-267

[38] Poorter L, Bongers L, Bongers F. Architecture of 54 moist-forest tree species: traits, trade-offs, and functional groups. Ecology, 2006, 87(5): 1289-1301

[39] Purves DW, Lichstein JW, Pacala SW. Crown plasticity and competition for canopy space: a new spatially implicit model parameterized for 250 North American tree species. PLoS ONE, 2007, 2(9 e870

[40] R Core Team (2024) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. https://www.R-project.org/

[41] Reich PB. The world-wide ‘fast–slow’ plant economics spectrum: a traits manifesto. J Ecol, 2014, 1022): 275-301

[42] Rüger N, Comita LS, Condit R, Purves D, Rosenbaum B, Visser MD, Wright SJ, Wirth C. Beyond the fast–slow continuum: demographic dimensions structuring a tropical tree community. Ecol Lett, 2018, 21(7): 1075-1084

[43] Rüger N, Condit R, Dent DH, DeWalt SJ, Hubbell SP, Lichstein JW, Lopez OR, Wirth C, Farrior CE. Demographic trade-offs predict tropical forest dynamics. Science, 2020, 368(6487): 165-168

[44] Saccenti E, Hendriks MHWB, Smilde AK. Corruption of the Pearson correlation coefficient by measurement error and its estimation, bias, and correction under different error models. Sci Rep, 2020, 10(1 438

[45] Seelig HD, Stoner RJ, Linden JC. Irrigation control of cowpea plants using the measurement of leaf thickness under greenhouse conditions. Irrig Sci, 2012, 304): 247-257

[46] Sheil D, Salim A, Chave J, Vanclay J, Hawthorne WD. Illumination–size relationships of 109 coexisting tropical forest tree species. J Ecol, 2006, 94(2): 494-507

[47] Slowikowski K (2024) ggrepel: automatically position non-overlapping text labels with 'ggplot2', version 0.9.3. CRAN. https://CRAN.R-project.org/package=ggrepel

[48] Sterck FJ, Bongers F, Newbery DM. Tree architecture in a Bornean lowland rain forest: intraspecific and interspecific patterns. Plant Ecol, 2001, 153(1): 279-292

[49] Thomas SC. Asymptotic height as a predictor of growth and allometric characteristics in Malaysian rain forest trees. Am J Bot, 1996, 83(5): 556-566

[50] Turner IM. The ecology of trees in the tropical rain forest, 2001, Cambridge, Cambridge University Press

[51] Vehtari A, Gelman A, Gabry J. Practical Bayesian model evaluation using leave-one-out cross-validation and WAIC. Stat Comput, 2017, 27(5): 1413-1432

[52] Whitmore TC. An introduction to tropical rain forests, 19982Oxford, Oxford University Press

[53] Wickham H. Data analysis. Ggplot2: elegant graphics for data analysis, 2016, Cham, Springer: 189-201

[54] Wright IJ, Reich PB, Westoby M, Ackerly DD, Baruch Z, Bongers F, Cavender-Bares J, Chapin T, Cornelissen JHC, Diemer M, Flexas J, Garnier E, Groom PK, Gulias J, Hikosaka K, Lamont BB, Lee T, Lee W, Lusk C, Midgley JJ, Navas ML, Niinemets U, Oleksyn J, Osada N, Poorter H, Poot P, Prior L, Pyankov VI, Roumet C, Thomas SC, Tjoelker MG, Veneklaas EJ, Villar R. The worldwide leaf economics spectrum. Nature, 2004, 428(6985): 821-827

[55] Wright SJ, Kitajima K, Kraft NJB, Reich PB, Wright IJ, Bunker DE, Condit R, Dalling JW, Davies SJ, Díaz S, Engelbrecht BMJ, Harms KE, Hubbell SP, Marks CO, Ruiz-Jaen MC, Salvador CM, Zanne AE. Functional traits and the growth–mortality trade-off in tropical trees. Ecology, 2010, 91(12): 3664-3674

[56] Wu ZY, Raven PH, Hong DY (2013) Flora of China. Science Press, Beijing; Missouri Botanical Garden Press, St Louis

[57] Yamada T, Ngakan OP, Suzuki E. Differences in growth trajectory and strategy of two sympatric congeneric species in an Indonesian floodplain forest. Am J Bot, 2005, 92(1): 45-52

[58] Yang J, Swenson NG. Height and crown allometries and their relationship with functional traits: an example from a subtropical wet forest. Ecol Evol, 2023, 13(2 e9804

[59] Zhang CC, Gu R, Lin LX, Russo SE. Functional traits and ecological niches as correlates of the interspecific growth–mortality trade-off among seedlings of 14 tropical tree species. Funct Ecol, 2024, 389): 1888-1901

Funding

Joint Fund of the National Natural Science Foundation of China-Yunnan Province(U1902203)

RIGHTS & PERMISSIONS

Northeast Forestry University