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Inferring community assembly processes from trait diversity across environmental gradients

Published online by Cambridge University Press:  13 June 2016

Yong Shen
Department of Ecology, School of Life Sciences/State Key Laboratory of Biocontrol, Sun Yat-sen University, Guangzhou 510275, PR China
Shi-Xiao Yu*
Department of Ecology, School of Life Sciences/State Key Laboratory of Biocontrol, Sun Yat-sen University, Guangzhou 510275, PR China
Ju-Yu Lian
Key Laboratory of Vegetation Restoration and Management of Degraded Ecosystems, South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510650, PR China
Hao Shen
Key Laboratory of Vegetation Restoration and Management of Degraded Ecosystems, South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510650, PR China
Hong-Lin Cao
Key Laboratory of Vegetation Restoration and Management of Degraded Ecosystems, South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510650, PR China
Huan-Ping Lu
Ecological Meteorological Center of Guangdong Province, Guangzhou 510640, PR China
Wan-Hui Ye*
Key Laboratory of Vegetation Restoration and Management of Degraded Ecosystems, South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510650, PR China
1Corresponding authors. Email:,
1Corresponding authors. Email:,


Environmental filtering and competitive interactions are important ecological processes in community assembly. The contribution of the two processes to community assembly can be evaluated by shifts in functional diversity patterns. We examined the correlations between functional diversity of six traits (leaf chlorophyll concentration, dry matter content, size, specific leaf area, thickness and wood density) and environmental gradients (topography and soil) for 92 species in the 20-ha Dinghushan forest plot in China. A partial Mantel test showed that most of the community-weighted mean trait values changed with terrain convexity and soil fertility, which implied that environmental filtering was occurring. Functional diversity of many traits significantly increased with increasing terrain convexity and soil fertility, which was associated with increased light and below-ground resources respectively. These results suggest that co-occurring species are functionally convergent in regions of strong abiotic stress under the environmental filtering, but functionally divergent in more benign environments due to resource partitioning and competitive interactions. Single-trait diversity and multivariate functional diversity had different relationships with environmental factors, indicating that traits were related to different niche axes, and associated with different ecological processes, which demonstrated the importance of focusing niche axes in traits selection. Between 9% and 41% of variation in functional diversity of different traits was explained by environmental factors in stepwise multiple regression models. Terrain convexity and soil fertility were the best predictors of functional diversity, which contributed 30.5% and 29.0% of total R2 to the model. These provided essential evidence that different environmental factors had distinguishing impacts on regulating diversity of traits.

Research Article
Copyright © Cambridge University Press 2016 

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BIN, Y., LIAN, J., WANG, Z., YE, W. & CAO, H. 2011. Tree mortality and recruitment in a subtropical broadleaved monsoon forest in South China. Journal of Tropical Forest Science 23:5766.Google Scholar
BIN, Y., YE, W. H., MULLER-LANDAU, H. C., WU, L. F., LIAN, J. Y. & CAO, H. L. 2012. Unimodal tree size distributions possibly result from relatively strong conservatism in intermediate size classes. PLoS ONE 7 (12).CrossRefGoogle ScholarPubMed
CARRENO-ROCABADO, G., PENA-CLAROS, M., BONGERS, F., ALARCON, A., LICONA, J. C. & POORTER, L. 2012. Effects of disturbance intensity on species and functional diversity in a tropical forest. Journal of Ecology 100:14531463.CrossRefGoogle Scholar
CHALMANDRIER, L., MUNKEMULLER, T., GALLIEN, L., DE BELLO, F., MAZEL, F., LAVERGNE, S. & THUILLER, W. 2013. A family of null models to distinguish between environmental filtering and biotic interactions in functional diversity patterns. Journal of Vegetation Science 24:853864.CrossRefGoogle ScholarPubMed
CHATURVEDI, R. K., RAGHUBANSHI, A. S. & SINGH, J. S. 2011. Leaf attributes and tree growth in a tropical dry forest. Journal of Vegetation Science 22:917931.CrossRefGoogle Scholar
CIELO, R., GNERI, M. A. & MARTINS, F. R. 2007. Position on slope, disturbance, and tree species coexistence in a seasonal semideciduous forest in SE Brazil. Plant Ecology 190:189203.CrossRefGoogle Scholar
COMITA, L. S. & ENGELBRECHT, B. M. J. 2009. Seasonal and spatial variation in water availability drive habitat associations in a tropical forest. Ecology 90:27552765.CrossRefGoogle Scholar
CONTI, G. & DIAZ, S. 2013. Plant functional diversity and carbon storage – an empirical test in semi-arid forest ecosystems. Journal of Ecology 101:1828.CrossRefGoogle Scholar
CORNELISSEN, J. H. C., LAVOREL, S., GARNIER, E., DIAZ, S., BUCHMANN, N., GURVICH, D. E., REICH, P. B., TER STEEGE, H., MORGAN, H. D., VAN DER HEIJDEN, M. G. A., PAUSAS, J. G. & POORTER, H. 2003. A handbook of protocols for standardised and easy measurement of plant functional traits worldwide. Australian Journal of Botany 51:335380.CrossRefGoogle Scholar
CORNWELL, W. K. & ACKERLY, D. D. 2009. Community assembly and shifts in plant trait distributions across an environmental gradient in coastal California. Ecological Monographs 79:109126.CrossRefGoogle Scholar
CORNWELL, W. K., SCHWILK, D. W. & ACKERLY, D. D. 2006. A trait-based test for habitat filtering: convex hull volume. Ecology 87:14651471.CrossRefGoogle ScholarPubMed
DETTO, M., MULLER-LANDAU, H. C., MASCARO, J. & ASNER, G. P. 2013. Hydrological networks and associated topographic variation as templates for the spatial organization of tropical forest vegetation. PLoS ONE 8 (10).CrossRefGoogle ScholarPubMed
ENOKI, T. & ABE, A. 2004. Saplings distribution in relation to topography and canopy openness in an evergreen broad-leaved forest. Plant Ecology 173:283291.CrossRefGoogle Scholar
FORTUNEL, C., PAINE, C. E. T., FINE, P. V. A., KRAFT, N. J. B. & BARALOTO, C. 2014. Environmental factors predict community functional composition in Amazonian forests. Journal of Ecology 102:145155.CrossRefGoogle Scholar
GARNIER, E., CORTEZ, J., BILLES, G., NAVAS, M. L., ROUMET, C., DEBUSSCHE, M., LAURENT, G., BLANCHARD, A., AUBRY, D., BELLMANN, A., NEILL, C. & TOUSSAINT, J. P. 2004. Plant functional markers capture ecosystem properties during secondary succession. Ecology 85:26302637.CrossRefGoogle Scholar
GROMPING, U. 2006. Relative importance for linear regression in R: the package relaimpo. Journal of Statistical Software 17 (1).CrossRefGoogle Scholar
HARMS, K. E., CONDIT, R., HUBBELL, S. P. & FOSTER, R. B. 2001. Habitat associations of trees and shrubs in a 50-ha neotropical forest plot. Journal of Ecology 89:947959.CrossRefGoogle Scholar
HOULE, G., MCKENNA, M. F. & LAPOINTE, L. 2001. Spatiotemporal dynamics of Floerkea proserpinacoides (Limnanthaceae), an annual plant of the deciduous forest of eastern North America. American Journal of Botany 88:594607.CrossRefGoogle Scholar
JANECEK, S., DE BELLO, F., HORNIK, J., BARTOS, M., CERNY, T., DOLEZAL, J., DVORSKY, M., FAJMON, K., JANECKOVA, P., JIRASKA, S., MUDRAK, O. & KLIMESOVA, J. 2013. Effects of land-use changes on plant functional and taxonomic diversity along a productivity gradient in wet meadows. Journal of Vegetation Science 24:898909.CrossRefGoogle Scholar
KATABUCHI, M., KUROKAWA, H., DAVIES, S. J., TAN, S. & NAKASHIZUKA, T. 2012. Soil resource availability shapes community trait structure in a species-rich dipterocarp forest. Journal of Ecology 100:643651.CrossRefGoogle Scholar
KAZAKOU, E., VILE, D., SHIPLEY, B., GALLET, C. & GARNIER, E. 2006. Co-variations in litter decomposition, leaf traits and plant growth in species from a Mediterranean old-field succession. Functional Ecology 20:2130.CrossRefGoogle Scholar
KING, D. A., DAVIES, S. J., SUPARDI, M. N. N. & TAN, S. 2005. Tree growth is related to light interception and wood density in two mixed dipterocarp forests of Malaysia. Functional Ecology 19:445453.CrossRefGoogle Scholar
LALIBERTE, E. & LEGENDRE, P. 2010. A distance-based framework for measuring functional diversity from multiple traits. Ecology 91:299305.CrossRefGoogle ScholarPubMed
LAN, G. Y., HU, Y. H., CAO, M. & ZHU, H. 2011. Topography related spatial distribution of dominant tree species in a tropical seasonal rain forest in China. Forest Ecology and Management 262:15071513.CrossRefGoogle Scholar
LEGENDRE, P. & FORTIN, M. J. 1989. Spatial pattern and ecological analysis. Vegetatio 80:107138.CrossRefGoogle Scholar
LEGENDRE, P., MI, X. C., REN, H. B., MA, K. P., YU, M. J., SUN, I. F. & HE, F. L. 2009. Partitioning beta diversity in a subtropical broad-leaved forest of China. Ecology 90:663674.CrossRefGoogle Scholar
LIN, G. J., STRALBERG, D., GONG, G. Q., HUANG, Z. L., YE, W. H. & WU, L. F. 2013. Separating the effects of environment and space on tree species distribution: from population to community. PLoS ONE 8 (2).Google Scholar
LIU, X., MENG, W., LIANG, G., LI, K., XU, W., HUANG, L. & YAN, J. 2014. Available phosphorus in forest soil increases with soil nitrogen but not total phosphorus: evidence from subtropical forests and a pot experiment. PLoS ONE 9 (2).Google ScholarPubMed
LOH, F. C. W., GRABOSKY, J. C. & BASSUK, N. L. 2002. Using the SPAD 502 meter to assess chlorophyll and nitrogen content of benjamin fig and cottonwood leaves. Horttechnology 12:682686.Google Scholar
MAC NALLY, R. 2002. Multiple regression and inference in ecology and conservation biology: further comments on identifying important predictor variables. Biodiversity and Conservation 11:13971401.CrossRefGoogle Scholar
MARTÍNEZ-VILALTA, J., MENCUCCINI, M., VAYREDA, J. & RETANA, J. 2010. Interspecific variation in functional traits, not climatic differences among species ranges, determines demographic rates across 44 temperate and Mediterranean tree species. Journal of Ecology 98:14621475.CrossRefGoogle Scholar
MASON, N. W. H., DE BELLO, F., DOLEZAL, J. & LEPS, J. 2011. Niche overlap reveals the effects of competition, disturbance and contrasting assembly processes in experimental grassland communities. Journal of Ecology 99:788796.CrossRefGoogle Scholar
MASON, N. W. H., RICHARDSON, S. J., PELTZER, D. A., DE BELLO, F., WARDLE, D. A., & ALLEN, R. B. 2012. Changes in coexistence mechanisms along a long-term soil chronosequence revealed by functional trait diversity. Journal of Ecology 100:678689.CrossRefGoogle Scholar
MASON, N. W. H., DE BELLO, F., MOUILLOT, D., PAVOINE, S. & DRAY, S. 2013. A guide for using functional diversity indices to reveal changes in assembly processes along ecological gradients. Journal of Vegetation Science 24:794806.CrossRefGoogle Scholar
MAYFIELD, M. M. & LEVINE, J. M. 2010. Opposing effects of competitive exclusion on the phylogenetic structure of communities. Ecology Letters 13:10851093.CrossRefGoogle ScholarPubMed
MCEWAN, R. W., LIN, Y. C., SUN, I. F., HSIEH, C. F., SU, S. H., CHANG, L. W., SONG, G. Z. M., WANG, H. H., HWONG, J. L., LIN, K. C., YANG, K. C. & CHIANG, J. M. 2011. Topographic and biotic regulation of aboveground carbon storage in subtropical broad-leaved forests of Taiwan. Forest Ecology and Management 262:18171825.CrossRefGoogle Scholar
MOUCHET, M. A., VILLEGER, S., MASON, N. W. H. & MOUILLOT, D. 2010. Functional diversity measures: an overview of their redundancy and their ability to discriminate community assembly rules. Functional Ecology 24:867876.CrossRefGoogle Scholar
NOGUCHI, H., ITOH, A., MIZUNO, T., SRI-NGERNYUANG, K., KANZAKI, M., TEEJUNTUK, S., SUNGPALEE, W., HARA, M., OHKUBO, T., SAHUNALU, P., DHANMMANONDA, P. & YAMAKURA, T. 2007. Habitat divergence in sympatric Fagaceae tree species of a tropical montane forest in northern Thailand. Journal of Tropical Ecology 23:549558.CrossRefGoogle Scholar
POORTER, L. & BONGERS, F. 2006. Leaf traits are good predictors of plant performance across 53 rain forest species. Ecology 87:17331743.CrossRefGoogle Scholar
POORTER, L., BONGERS, F., STERCK, F. J. & WOLL, H. 2005. Beyond the regeneration phase: differentiation of height-light trajectories among tropical tree species. Journal of Ecology 93:256267.CrossRefGoogle Scholar
POORTER, L., WRIGHT, S. J., PAZ, H., ACKERLY, D. D., CONDIT, R., IBARRA-MANRIQUES, G., HARMS, K. E., LICONA, J. C., MARTINEZ-RAMOS, M., MAZER, S. J., MULLER-LANDAU, H. C., PENA-CLAROS, M., WEBB, C. O. & WRIGHT, I. J. 2008. Are functional traits good predictors of demographic rates? Evidence from five Neotropical forests. Ecology 89:19081920.CrossRefGoogle ScholarPubMed
PURSCHKE, O., SCHMID, B. C., SYKES, M. T., POSCHLOD, P., MICHALSKI, S. G., DURKA, W., KUHN, I., WINTER, M. & PRENTICE, H. C. 2013. Contrasting changes in taxonomic, phylogenetic and functional diversity during a long-term succession: insights into assembly processes. Journal of Ecology 101:857866.CrossRefGoogle Scholar
ROSCHER, C., SCHUMACHER, J., LIPOWSKY, A., GUBSCH, M., WEIGELT, A., POMPE, S., KOLLE, O., BUCHMANN, N., SCHMID, B. & SCHULZE, E. D. 2013. A functional trait-based approach to understand community assembly and diversity-productivity relationships over 7 years in experimental grasslands. Perspectives in Plant Ecology Evolution and Systematics 15:139149.CrossRefGoogle Scholar
RUBINO, D. L. & MCCARTHY, B. C. 2003. Evaluation of coarse woody debris and forest vegetation across topographic gradients in a southern Ohio forest. Forest Ecology and Management 183:221238.CrossRefGoogle Scholar
RUGER, N., HUTH, A., HUBBELL, S. P. & CONDIT, R. 2009. Response of recruitment to light availability across a tropical lowland rain forest community. Journal of Ecology 97:13601368.CrossRefGoogle Scholar
SHEN, Y., SANTIAGO, L. S., MA, L., LIN, G. J., LIAN, J. Y., CAO, H. L. & YE, W. H. 2013. Forest dynamics of a subtropical monsoon forest in Dinghushan, China: recruitment, mortality and the pace of community change. Journal of Tropical Ecology 29:131145.CrossRefGoogle Scholar
SHEN, Y., SANTIAGO, L. S., SHEN, H., MA, L., LIAN, J. Y., CAO, H. L., LU, H. P. & YE, W. H. 2014. Determinants of change in subtropical tree diameter growth with ontogenetic stage. Oecologia 175:13151324.CrossRefGoogle ScholarPubMed
SIEFERT, A., RAVENSCROFT, C., WEISER, M. D. & SWENSON, N. G. 2013. Functional beta-diversity patterns reveal deterministic community assembly processes in eastern North American trees. Global Ecology and Biogeography 22:682691.CrossRefGoogle Scholar
SPASOJEVIC, M. J. & SUDING, K. N. 2012. Inferring community assembly mechanisms from functional diversity patterns: the importance of multiple assembly processes. Journal of Ecology 100:652661.CrossRefGoogle Scholar
STERCK, F. J., POORTER, L. & SCHIEVING, F. 2006. Leaf traits determine the growth-survival trade-off across rain forest tree species. American Naturalist 167:758765.Google ScholarPubMed
SUTTON-GRIER, A. E., WRIGHT, J. P., MCGILL, B. M. & RICHARDSON, C. 2011. Environmental conditions influence the plant functional diversity effect on potential denitrification. PLoS ONE 6 (2).CrossRefGoogle ScholarPubMed
SWENSON, N. G. & ENQUIST, B. J. 2009. Opposing assembly mechanisms in a Neotropical dry forest: implications for phylogenetic and functional community ecology. Ecology 90:21612170.CrossRefGoogle Scholar
TSUJINO, R., TAKAFUMI, H., AGETSUMA, N. & YUMOTO, T. 2006. Variation in tree growth, mortality and recruitment among topographic positions in a warm temperate forest. Journal of Vegetation Science 17:281290.CrossRefGoogle Scholar
VILLEGER, S., MASON, N. W. H. & MOUILLOT, D. 2008. New multidimensional functional diversity indices for a multifaceted framework in functional ecology. Ecology 89:22902301.CrossRefGoogle ScholarPubMed
WANG, Z., YE, W. H., CAO, H. L., HUANG, Z. L., LIAN, J. Y., LI, L., WEI, S. G. & SUN, I. F. 2009. Species-topography association in a species-rich subtropical forest of China. Basic and Applied Ecology 10:648655.CrossRefGoogle Scholar
WEIHER, E. & KEDDY, P. A. 1995. Assembly rules, null models, and trait dispersion – new questions from old patterns. Oikos 74:159164.CrossRefGoogle Scholar
WRIGHT, I. J., REICH, P. B., WESTOBY, M., ACKERLY, D. D., BARUCH, Z., BONGERS, F., CAVENDER-BARES, J., CHAPIN, T., CORNELISSEN, J. H. C., DIEMER, M., FLEXAS, J., GARNIER, E., GROOM, P. K., GULIAS, J., HIKOSAKA, K., LAMONT, B. B., LEE, T., LEE, W., LUSK, C., MIDGLEY, J. J., NAVAS, M. L., NIINEMETS, U., OLEKSYN, J., OSADA, N., POORTER, H., POOT, P., PRIOR, L., PYANKOV, V. I., ROUMET, C., THOMAS, S. C., TJOELKER, M. G., VENEKLAAS, E. J. & VILLAR, R. 2004. The worldwide leaf economics spectrum. Nature 428:821827.CrossRefGoogle Scholar
WRIGHT, I. J., ACKERLY, D. D., BONGERS, F., HARMS, K. E., IBARRA-MANRIQUEZ, G., MARTINEZ-RAMOS, M., MAZER, S. J., MULLER-LANDAU, H. C., PAZ, H., PITMAN, N. C. A., POORTER, L., SILMAN, M. R., VRIESENDORP, C. F., WEBB, C. O., WESTOBY, M. & WRIGHT, S. J. 2007. Relationships among ecologically important dimensions of plant trait variation in seven Neotropical forests. Annals of Botany 99:10031015.CrossRefGoogle ScholarPubMed
WRIGHT, S. J., KITAJIMA, K., KRAFT, N. J. B., REICH, P. B., WRIGHT, I. J., BUNKER, D. E., CONDIT, R., DALLING, J. W., DAVIES, S. J., DIAZ, S., ENGELBRECHT, B. M. J., HARMS, K. E., HUBBELL, S. P., MARKS, C. O., RUIZ-JAEN, M. C., SALVADOR, C. M. & ZANNE, A. E. 2010. Functional traits and the growth-mortality trade-off in tropical trees. Ecology 91:36643674.CrossRefGoogle ScholarPubMed