Difference in survival response of tree species to neighborhood crowding in a lower subtropical evergreen broad-leaved forest of Dinghushan

MA Qinhong1,2,3 LI Yanpeng 1,2,3 LIAN Juyu 1,2 YE Wanhui 1,2

(1.Key Laboratory of Vegetation Restoration and Management of Degraded Ecosystems, South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510650)
(2.Guangdong Provincial Key Laboratory of Applied Botany, South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510650)
(3.University of Chinese Academy of Sciences, Beijing 100049)

【Abstract】Identifying the mechanisms that drive community structure and dynamics is one of the most fundamental goals in ecology. The local neighborhood in which trees grow significantly influences species survival. In order to understand the mechanisms underlying the various survival responses to neighborhood crowding among species, we compared the survival responses of focal tree species through modeling tree survival in terms of neighborhood effects. These effects were quantified in different ways, based on the survival monitoring and functional trait data of 90 species commonly observed in a lower tropical evergreen broad-leaved forest 20-hm2 plot in the Dinghu Mountains. We found that among all species tested, 58% showed sensitivity to neighborhood effects, and that the survival responses of 50% were mediated by functional trait differences among co-occurring species. Shade tolerance of species is associated with species sensitivity to its neighborhood since the species with lower shade tolerance are intended to be sensitive to their neighbors. Lower specific leaf area, higher leaf dry matter content, wood density and maximum diameter at breast height indicate higher shade tolerance. Niche difference associated with light acquisition strategies may underlie species coexistence at the neighborhood scale. Our study provides new insights into quantifying neighborhood interaction and species coexistence at the local neighborhood scale.

【Keywords】 neighborhood crowding; survival response; trait difference; functional trait; shade tolerance;


【Funds】 National Natural Science Foundation of China (41371078) Strategic Priority Research Program of Chinese Academy of Sciences (XDPB0203) Forest Biodiversity Monitoring Network Construction Project of China

Download this article

(Translated by XU XY)


    Adler PB, Fajardo A, Kleinhesselink AR, Kraft NJ (2013) Trait-based tests of coexistence mechanisms. Ecology Letters, 16, 1294–1306.

    Bernard-Verdier M, Navas M, Vellend M, Violle C, Fayolle A, Garnier E (2012) Community assembly along a soil depth gradient: Contrasting patterns of plant trait convergence and divergence in a Mediterranean rangeland. Journal of Ecology, 100, 1422–1433.

    Cadotte MW, Tucker CM (2017) Should environmental filtering be abandoned? Trends in Ecology & Evolution, 32, 429–437.

    Canham CD, Papaik MJ, Uriarte M, McWilliams WH, Jenkins JC, Twery MJ (2006) Neighborhood analyses of canopy tree competition along environmental gradients in new England forests. Ecological Applications, 16, 540–554.

    Chen YX, Wright SJ, Muller-Landau HC, Hubbell SP, Wang YF, Yu SX (2016) Positive effects of neighborhood complementarity on tree growth in a Neotropical forest. Ecology, 97, 776–785.

    Comita LS, Muller-Landau HC, Aguilar S, Hubbell SP (2010) Asymmetric density dependence shapes species abundances in a tropical tree community. Science, 329, 330–332.

    Comita LS, Hubbell SP (2009) Local neighborhood and species’ shade tolerance influence survival in a diverse seedling bank. Ecology, 90, 328–334.

    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 (2003) A handbook of protocols for standardised and easy measurement of plant functional traits worldwide. Australian Journal of Botany, 51, 335–380.

    Díaz S, Kattge J, Cornelissen JH, Wright IJ, Lavorel S, Dray S, Reu B, Kleyer M, Wirth C, Prentice IC, Garnier E, Bonisch 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, Gunther A, Falczuk V, Ruger N, Mahecha MD, Gorne LD (2016) The global spectrum of plant form and function. Nature, 529, 167–171.

    Fortunel C, Valencia R, Wright SJ, Garwood NC, Kraft NJB (2016) Functional trait differences influence neighborhood interactions in a hyperdiverse Amazonian forest. Ecology Letters, 19, 1062–1070.

    Gravel D, Canham CD, Beaudet M, Messier C (2010) Shade tolerance, canopy gaps and mechanisms of coexistence of forest trees. Oikos, 119, 475–484.

    Han BC, Umaña MN, Mi XC, Liu XJ, Chen L, Wang YQ, Liang Y, Wei W, Ma KP (2017) The role of transcriptomes linked with responses to light environment on seedling mortality in a subtropical forest, China. Journal of Ecology, 105, 592–601.

    He FL, Duncan RP (2000) Density-dependent effects on tree survival in an old-growth Douglas fir forest. Journal of Ecology, 88, 676–688.

    HilleRisLambers J, Adler PB, Harpole WS, Levine JM, Mayfield MM (2012) Rethinking community assembly through the lens of coexistence theory. Annual Review of Ecology, Evolution, and Systematics, 43, 227–248.

    HilleRisLambers J, Clark JS, Beckage B (2002) Density-dependent mortality and the latitudinal gradient in species diversity. Nature, 417, 732–735.

    Hubbell SP, Ahumada JA, Condit R, Foster RB (2001) Local neighborhood effects on long-term survival of individual trees in a neotropical forest. Ecological Research, 16, 859–875.

    Johnson DJ, Bourg NA, Howe R, McShea WJ, Wolf A, Clay K (2014) Conspecific negative density-dependent mortality and the structure of temperate forests. Ecology, 95, 2493–2503.

    Kobe RK, Vriesendorp CF (2011) Conspecific density dependence in seedlings varies with species shade tolerance in a wet tropical forest. Ecology Letters, 14, 503–510.

    Kraft NJB, Ackerly DD (2010) Functional trait and phylogenetic tests of community assembly across spatial scales in an Amazonian forest. Ecological Monographs, 80, 401–422.

    Kraft NJB, Adler PB, Godoy O, James EC, Fuller S, Levine JM, Fox J (2015) Community assembly, coexistence and the environmental filtering metaphor. Functional Ecology, 29, 592–599.

    Kraft NJB, Valencia R, Ackerly DD (2008) Functional traits and niche-based tree community assembly in an Amazonian forest. Science, 322, 580–582.

    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 JH, Gourlet-Fleury S, Hanewinkel M, Herault B, Kattge J, Kurokawa H, Onoda Y, Penuelas J, Poorter H, Uriarte M, Richardson S, Ruiz-Benito P, Sun IF, Stahl G, Swenson NG, Thompson J, Westerlund B, Wirth C, Zavala MA, Zeng H, Zimmerman JK, Zimmermann NE, Westoby M (2016) Plant functional traits have globally consistent effects on competition. Nature, 529, 204–207.

    Kunstler G, Lavergne S, Courbaud B, Thuiller W, Vieilledent G, Zimmermann NE, Kattge J, Coomes DA (2012) Competitive interactions between forest trees are driven by species’ trait hierarchy, not phylogenetic or functional similarity: Implications for forest community assembly. Ecology Letters, 15, 831–840.

    Lasky JR, Uriarte M, Boukili VK, Chazdon RL (2014) Trait-mediated assembly processes predict successional changes in community diversity of tropical forests. Proceedings of the National Academy of Sciences, USA, 111, 5616–5621.

    Lasky JR, Bachelot B, Muscarella R, Schwartz N, Forero-Montaña J, Nytch CJ, Swenson NG, Thompson J, Zimmerman JK, Uriarte M (2015) Ontogenetic shifts in trait-mediated mechanisms of plant community assembly. Ecology, 96, 2157–2169.

    Ma L, Chen C, Shen Y, Wu LF, Huang ZL, Cao HL (2014) Determinants of tree survival at local scale in a sub-tropical forest. Ecological Research, 29, 69–80.

    Mayfield MM, Levine JM (2010) Opposing effects of competitive exclusion on the phylogenetic structure of communities. Ecology Letters, 13, 1085–1093.

    McCarthy-Neumann S, Kobe R (2008) Tolerance of soil pathogens co-varies with shade tolerance across species of tropical tree seedlings. Ecology, 89, 1883–1892.

    McGill BJ, Enquist BJ, Weiher E, Westoby M (2006) Rebuilding community ecology from functional traits. Trends in Ecology & Evolution, 21, 178–185.

    Pérez-Harguindeguy N, Díaz S, Garnier E, Lavorel S, Poorter H, Jaureguiberry P, Bret-Harte MS, Cornwell WK, Craine JM, Gurvich DE, Urcelay C, Veneklaas EJ, Reich PB, Poorter L, Wright IJ, Ray P, Enrico L, Pausas JG, de Vos AC, Buchmann N, Funes G, Quétier F, Hodgson JG, Thompson K, Morgan HD, ter Steege H, Sack L, Blonder B, Poschlod P, Vaieretti MV, Conti G, Staver AC, Aquino S, Cornelissen JHC (2013) New handbook for standardised measurement of plant functional traits worldwide. Australian Journal of Botany, 61, 167.

    Peters HA (2003) Neighbour-regulated mortality: The influence of positive and negative density dependence on tree populations in species-rich tropical forests. Ecology Letters, 6, 757–765.

    Poorter L, Wright SJ, Paz H, Ackerly DD, Condit R, Ibarra-Manriques G, Harms KE, Licona JC, Martinez-Ramos M, Mazer SJ, Muller-Landau HC, Pena-Claros M, Webb CO, Wright IJ (2008) Are functional traits good predictors of demographic rates? Evidence from five Neotropical forests. Ecology, 89, 1908–1920.

    R Core Team (2018) R: A Language and Environment for Statistical Computing. R foundation for statistical computing, Vienna, Austria. http://www.R-project.org/ (accessed on 2018-03-12).

    Shen Y, Santiago LS, Ma L, Lin GJ, Lian JY, Cao HL, Ye WH (2013) Forest dynamics of a subtropical monsoon forest in Dinghushan, China: Recruitment, mortality and the pace of community change. Journal of Tropical Ecology, 29, 131–145.

    Shen Y, Santiago LS, Shen H, Ma L, Lian JY, Cao HL, Lu HP, Ye WH (2014) Determinants of change in subtropical tree diameter growth with ontogenetic stage. Oecologia, 175, 1315–1324.

    Spasojevic MJ, Suding KN (2012) Inferring community assembly mechanisms from functional diversity patterns: The importance of multiple assembly processes. Journal of Ecology, 100, 652–661.

    Stoll P, Newbery DM (2005) Evidence of species-specific neighborhood effects in the dipterocarpaceae of a Bornean rain forest. Ecology, 86, 3048–3062.

    Uriarte M, Canham CD, Thompson J, Zimmerman JK (2004a) A neighborhood analysis of tree growth and survival in a hurricane-driven tropical forest. Ecological Monographs, 74, 591–614.

    Uriarte M, Condit R, Canham CD, Hubbell SP (2004b) A spatially explicit model of sapling growth in a tropical forest: Does the identity of neighbours matter? Journal of Ecology, 92, 348–360.

    Uriarte M, Swenson NG, Chazdon RL, Comita LS, Kress WJ, Erickson D, Forero-Montana J, Zimmerman JK, Thompson J (2010) Trait similarity, shared ancestry and the structure of neighborhood interactions in a subtropical wet forest: Implications for community assembly. Ecology Letters, 13, 1503–1514.

    Valladares F, Niinemets U (2008) Shade tolerance, a key plant feature of complex nature and consequences. Annual Review of Ecology, Evolution, and Systematics, 39, 237–257.

    Wang XG, Comita LS, Hao ZQ, Davies SJ, Ye J, Lin F, Yuan ZQ (2012) Local-scale drivers of tree survival in a temperate forest. PLoS ONE, 7, 10.

    Wiegand T, Uriarte M, Kraft NJB, Shen GC, Wang XG, He FL (2017) Spatially explicit metrics of species diversity, functional diversity, and phylogenetic diversity: Insights into plant community assembly processes. Annual Review of Ecology, Evolution, and Systematics, 48, 329–351.

    Wright SJ (2002) Plant diversity in tropical forests: A review of mechanisms of species coexistence. Oecologia, 130, 1–14.

    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 (2010) Functional traits and the growth-mortality trade-off in tropical trees. Ecology, 91, 3664–3674.

    Wright SJ, Muller-Landau HC, Condit R, Hubbell SP (2003) Gap-dependent recruitment, realized vital rates, and size distributions of tropical trees. Ecology, 84, 3174–3185.

    Ye WH, Cao HL, Huang ZL, Lian JY, Wang ZG, Li L, Wei SG, Wang ZM (2008) Community structure of a 20 hm2 lower subtropical evergreen broadleaved forest plot in Dinghushan, China. Journal of Plant Ecology (Chinese Version), 32, 274–286 (in Chinese with English abstract).

    Zambrano J, Marchand P, Swenson NG (2017) Local neighborhood and regional climatic contexts interact to explain tree performance. Proceedings of the Royal Society B: Biological Sciences, 284, 20170523.

    Zhu Y, Hogan JA, Cai HY, Xun YH, Jiang F, Jin GZ (2017) Biotic and abiotic drivers of the tree growth and mortality trade-off in an old-growth temperate forest. Forest Ecology and Management, 404, 354–360.

This Article



Vol 26, No. 06, Pages 535-544

June 2018


Article Outline


  • 1 Materials and methods
  • 2 Results
  • 3 Discussion
  • References