Trait variation and functional diversity maintenance of understory herbaceous species coexisting along an elevational gradient in Yulong Mountain, Southwest China

doi:10.1016/j.pld.2016.11.002

Plant Diversity ›› 2016, Vol. 38 ›› Issue (06): 303-311.DOI: 10.1016/j.pld.2016.11.002

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Trait variation and functional diversity maintenance of understory herbaceous species coexisting along an elevational gradient in Yulong Mountain, Southwest China

Yahuang Luo^a,b,c, Jie Liu^a, Shaolin Tan^a,c, Marc W. Cadotte^d, Kun Xu^e, Lianming Gao^a, Dezhu Li^a,b,c

a. Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, China;
b. Germplasm Bank of Wild Species in Southwest China, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, China;
c. College of Life Science, University of Chinese Academy of Sciences, Kunming, Yunnan 650201, China;
d. Biological Sciences, University of Toronto-Scarborough & Ecology and Evolutionary Biology, University of Toronto, 1265 Military Trail, Toronto, ON M1C1A4, Canada;
e. Lijiang Forest Ecosystem Research Station, Kunming Institute of Botany, Chinese Academy of Sciences, Lijiang 674100, China

Received:2016-09-27 Revised:2016-11-07 Online:2021-11-05 Published:2016-12-25
Contact: Lianming Gao, Dezhu Li
Supported by:
We are grateful to Dr. Zhikun Wu, Detuan Liu, Zhifa Chen, Hua Huang, Weiwei Liu and Xiaoling Chen from Lijiang Forest Ecosystem Research Station for their help with field data collection. This study was supported by the National Key Basic Research Program of China (2014CB954100), the Ministry of Science and Technology of the People's Republic of China (2012FY110800), and the Applied Fundamental Research Foundation of Yunnan Province (2014GA003).

Trait variation and functional diversity maintenance of understory herbaceous species coexisting along an elevational gradient in Yulong Mountain, Southwest China

Yahuang Luo^a,b,c, Jie Liu^a, Shaolin Tan^a,c, Marc W. Cadotte^d, Kun Xu^e, Lianming Gao^a, Dezhu Li^a,b,c

a. Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, China;
b. Germplasm Bank of Wild Species in Southwest China, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, China;
c. College of Life Science, University of Chinese Academy of Sciences, Kunming, Yunnan 650201, China;
d. Biological Sciences, University of Toronto-Scarborough & Ecology and Evolutionary Biology, University of Toronto, 1265 Military Trail, Toronto, ON M1C1A4, Canada;
e. Lijiang Forest Ecosystem Research Station, Kunming Institute of Botany, Chinese Academy of Sciences, Lijiang 674100, China

通讯作者: Lianming Gao, Dezhu Li
基金资助:
We are grateful to Dr. Zhikun Wu, Detuan Liu, Zhifa Chen, Hua Huang, Weiwei Liu and Xiaoling Chen from Lijiang Forest Ecosystem Research Station for their help with field data collection. This study was supported by the National Key Basic Research Program of China (2014CB954100), the Ministry of Science and Technology of the People's Republic of China (2012FY110800), and the Applied Fundamental Research Foundation of Yunnan Province (2014GA003).

Abstract

Abstract: Characterizing trait variation across different ecological scales in plant communities has been viewed as a way to gain insights into the mechanisms driving species coexistence. However, little is known about how changes in intraspecific and interspecific traits across sites influence species richness and community assembly, especially in understory herbaceous communities. Here we partitioned the variance of four functional traits (maximum height, leaf thickness, leaf area and specific leaf area) across four nested biological scales: individual, species, plot, and elevation to quantify the scale-dependent distributions of understory herbaceous trait variance. We also integrated the comparison of the trait variance ratios to null models to investigate the effects of different ecological processes on community assembly and functional diversity along a 1200-m elevational gradient in Yulong Mountain. We found interspecific trait variation was the main trait variation component for leaf traits, although intraspecific trait variation ranged from 10% to 28% of total variation. In particular, maximum height exhibited high plasticity, and intraspecific variation accounted for 44% of the total variation. Despite the fact that species composition varied across elevation and species richness decreased dramatically along the elevational gradient, there was little variance at our largest (elevation) scale in leaf traits and functional diversity remained constant along the elevational gradient, indicating that traits responded to smaller scale influences. External filtering was only observed at high elevations. However, strong internal filtering was detected along the entire elevational gradient in understory herbaceous communities, possibly due to competition. Our results provide evidence that species coexistence in understory herbaceous communities might be structured by differential niche-assembled processes. This approach--integrating different biological scales of trait variation--may provide a better understanding of the mechanisms involved in the structure of communities.

Key words: Biological scale, Community assembly, Functional diversity, Intraspecific variation, Species richness, Trait variance ratio

摘要： Characterizing trait variation across different ecological scales in plant communities has been viewed as a way to gain insights into the mechanisms driving species coexistence. However, little is known about how changes in intraspecific and interspecific traits across sites influence species richness and community assembly, especially in understory herbaceous communities. Here we partitioned the variance of four functional traits (maximum height, leaf thickness, leaf area and specific leaf area) across four nested biological scales: individual, species, plot, and elevation to quantify the scale-dependent distributions of understory herbaceous trait variance. We also integrated the comparison of the trait variance ratios to null models to investigate the effects of different ecological processes on community assembly and functional diversity along a 1200-m elevational gradient in Yulong Mountain. We found interspecific trait variation was the main trait variation component for leaf traits, although intraspecific trait variation ranged from 10% to 28% of total variation. In particular, maximum height exhibited high plasticity, and intraspecific variation accounted for 44% of the total variation. Despite the fact that species composition varied across elevation and species richness decreased dramatically along the elevational gradient, there was little variance at our largest (elevation) scale in leaf traits and functional diversity remained constant along the elevational gradient, indicating that traits responded to smaller scale influences. External filtering was only observed at high elevations. However, strong internal filtering was detected along the entire elevational gradient in understory herbaceous communities, possibly due to competition. Our results provide evidence that species coexistence in understory herbaceous communities might be structured by differential niche-assembled processes. This approach--integrating different biological scales of trait variation--may provide a better understanding of the mechanisms involved in the structure of communities.

关键词: Biological scale, Community assembly, Functional diversity, Intraspecific variation, Species richness, Trait variance ratio

Yahuang Luo, Jie Liu, Shaolin Tan, Marc W. Cadotte, Kun Xu, Lianming Gao, Dezhu Li. Trait variation and functional diversity maintenance of understory herbaceous species coexisting along an elevational gradient in Yulong Mountain, Southwest China[J]. Plant Diversity, 2016, 38(06): 303-311.

References

Abràmoff, M.D., Magalhães, P.J., Ram, S.J., 2004. Image processing with ImageJ. Biophot. Int. 11, 36-43.
Ackerly, D.D., Cornwell, W., 2007. A trait-based approach to community assembly:partitioning of species trait values into within-and among-community components. Ecol. Lett. 10, 135-145.
Adler, P.B., HilleRisLambers, J., Levine, J.M., 2007. A niche for neutrality. Ecol. Lett. 10, 95-104.
Albert, C.H., Thuiller, W., Yoccoz, N.G., Douzet, R., Aubert, S., Lavorel, S., 2010.A multi-trait approach reveals the structure and the relative importance of intra-vs. interspecific variability in plant traits. Funct. Ecol. 24, 1192-1201.
Auger, S., Shipley, B., 2013. Inter-specific and intra-specific trait variation along short environmental gradients in an old-growth temperate forest. J. Veg. Sci. 24, 419-428.
Bagousse-Pinguet, L., Bello, F., Vandewalle, M., Leps, J., Sykes, M.T., 2014. Species richness of limestone grasslands increases with trait overlap: evidence from within-and between-species functional diversity partitioning. J. Ecol. 102, 466-474.
Bello, F.d., Lavorel, S., Lavergne, S., Albert, C.H., Boulangeat, I., Mazel, F., Thuiller, W., 2013. Hierarchical effects of environmental filters on the functional structure of plant communities: a case study in the French alps.Ecography 36, 393-402.
Bivand, R., 2012. Spdep: Spatial Dependence: Weighting Schemes, Statistics and Models. R package version. 0.5-53. http://CRAN.R-project.org/package=spdep.
Bland, J.M., Altman, D.G., 1996. Transformations, means, and confidence intervals.B. M. J. 312, 1079.
Carmona, C.P., de Bello, F., Mason, N., Lepš, J., 2016. Traits without borders: integrating functional diversity across scales. Trends Ecol. Evol. 31, 382-394.
Chao, A., Lee, S.M., 1992. Estimating the number of classes via sample coverage.J. Am. Stat. Assoc. 87, 210-217.
Chesson, P., 2000. Mechanisms of maintenance of species diversity. Annu. Rev. Ecol.Syst. 343-366.
Clark, J.S., 2010. Individuals and the variation needed for high species diversity in forest trees. Science 327, 1129-1132.
Clark, J.S., Dietze, M., Chakraborty, S., Agarwal, P.K., Ibanez, I., LaDeau, S., Wolosin, M., 2007. Resolving the biodiversity paradox. Ecol. Lett. 10, 647-659.
Cornelissen, J., Lavorel, S., Garnier, E., Diaz, S., Buchmann, N., Gurvich, D., et al., 2003.A handbook of protocols for standardised and easy measurement of plant functional traits worldwide. Aust. J. Bot. 51, 335-380.
Dıaz, S., Cabido, M., 2001. Vive la difference: plant functional diversity matters to ecosystem processes. Trends Ecol. Evol. 16, 646-655.
Diaz, S., Lavorel, S., de Bello, F., Quetier, F., Grigulis, K., Robson, M., 2007. Incorporating plant functional diversity effects in ecosystem service assessments. Proc.Natl. Acad. Sci. U. S. A. 104, 20684-20689.
Falster, D.S., Westoby, M., 2003. Plant height and evolutionary games. Trends Ecol.Evol. 18, 337-343.
Feng, J.M., Wang, X.P., Xu, C.D., Yang, Y.H., Fang, J.Y., 2006. Altitudinal patterns of plant species diversity and community structure on Yulong Mountains, Yunnan.China J. Mt. Sci. 24, 110-116.
Grime, J.P., 2006. Trait convergence and trait divergence in herbaceous plant communities: mechanisms and consequences. J. Veg. Sci. 17, 255-260.
Hulshof, C.M., Swenson, N.G., 2010. Variation in leaf functional trait values within and across individuals and species: an example from a Costa Rican dry forest.Funct. Ecol. 24, 217-223.
Jung, V., Violle, C., Mondy, C., Hoffmann, L., Muller, S., 2010. Intraspecific variability and trait-based community assembly. J. Ecol. 98, 1134-1140.
Körner, C., 2007. The use of ‘altitude’ in ecological research. Trends Ecol. Evol. 22, 569-574.
Kang, M., Chang, S.X., Yan, E.R., Wang, X.H., 2014. Trait variability differs between leaf and wood tissues across ecological scales in subtropical forests. J. Veg. Sci. 25, 703-714.
Kissling, W.D., Carl, G., 2007. Spatial autocorrelation and the selection of simultaneous autoregressive models. Glob. Ecol. Biogeogr. 17, 59-71.
Kraft, N.J., Godoy, O., Levine, J.M., 2015. Plant functional traits and the multidimensional nature of species coexistence. Proc. Natl. Acad. Sci. U. S. A. 112, 797-802.
Kraft, N.J., Valencia, R., Ackerly, D.D., 2008. Functional traits and niche-based tree community assembly in an Amazonian forest. Science 322, 580-582.
Laliberté, E., Legendre, P., 2010. A distance-based framework for measuring functional diversity from multiple traits. Ecology 91, 299-305.
Laliberté, E., Shipley, B., 2011. FD: Measuring Functional Diversity from Multiple Traits, and Other Tools for Functional Ecology. R package version 1.0-11. https://CRAN.R-project.org/package=FD.
Lepš, J., de Bello, F., Šmilauer, P., Doležal, J., 2011. Community trait response to environment: disentangling species turnover vs intraspecific trait variability effects. Ecography 34, 856-863.
Luo, Y.H., Liu, J., Tan, S.L., Cadotte, M.W., Wang, Y.H., Xu, K., Li, D.Z., Gao, L.M., 2016.Trait-based community assembly along an elevational gradient in subalpine forests: quantifying the roles of environmental factors in inter-and intraspecific variability. PLoS One 11, e0155749.
Mandle, L., Ticktin, T., 2015. Moderate land use changes plant functional composition without loss of functional diversity in India's Western Ghats. Ecol. Appl. 25, 1711-1724.
Mason, N.W., Mouillot, D., Lee, W.G., Wilson, J.B., 2005. Functional richness, functional evenness and functional divergence: the primary components of functional diversity. Oikos 111, 112-118.
Mayfield, M.M., Levine, J.M., 2010. Opposing effects of competitive exclusion on the phylogenetic structure of communities. Ecol. Lett. 13, 1085-1093.
McGill, B.J., Enquist, B.J., Weiher, E., Westoby, M., 2006. Rebuilding community ecology from functional traits. Trends Ecol. Evol. 21, 178-185.
Messier, J., McGill, B.J., Enquist, B.J., Lechowicz, M.J., 2016. Trait variation and integration across scales: is the leaf economic spectrum present at local scales? Ecography. http://dx.doi.org/10.1111/ecog.02006.
Messier, J., McGill, B.J., Lechowicz, M.J., 2010. How do traits vary across ecological scales? A case for trait-based ecology. Ecol. Lett. 13, 838-848.
Moraes, D.A., Cavalin, P.O., Moro, R.S., Oliveira, R.A., Carmo, M.R., Marques, M., 2016.Edaphic filters and the functional structure of plant assemblages in grasslands in southern Brazil. J. Veg. Sci. 27, 100-110.
Mouillot, D., Stubbs, W., Faure, M., Dumay, O., Tomasini, J.A., Wilson, J.B., Do Chi, T., 2005. Niche overlap estimates based on quantitative functional traits: a new family of non-parametric indices. Oecologia 145, 345-353.
Myers, N., Mittermeier, R.A., Mittermeier, C.G., Da Fonseca, G.A., Kent, J., 2000.Biodiversity hotspots for conservation priorities. Nature 403, 853-858.
Neyret, M., Bentley, L.P., Oliveras, I., Marimon, B.S., Marimon-Junior, B.H., Almeida de Oliveira, et al., 2016. Examining variation in the leaf mass per area of dominant species across two contrasting tropical gradients in light of community assembly. Ecol. Evol. 6, 5674-5689.
Oksanen, J., Blanchet, F.G., Kindt, R., Legendre, P., O'Hara, R., Simpson, G.L., et al., 2011. Vegan: Community Ecology Package. R package version 2.0-0. http://CRAN.R-project.org/package=vegan.
Pérez-Harguindeguy, N., Díaz, S., Garnier, E., Lavorel, S., Poorter, H., Jaureguiberry, P., et al., 2013. New handbook for standardised measurement of plant functional traits worldwide. Aust. J. Bot. 61, 167-234.
Pinheiro, J., Bates, D., DebRoy, S., Sarkar, D., 2015. R Core Team Nlme: Linear and Nonlinear Mixed Effects Models. R package version 3.1-120. http://CRAN.Rproject.org/package=nlme.
Qu, L., 2015. Package ‘varComp’. R package Version 0.1-360. http://CRAN.R-project.org/package=varComp.
R Development Core Team, 2015. R: a Language and Environment for Statistical Computing. R Foundation for Statistical Computing. Vienna, Austria. http://www.R-project.org/.
Roche, P., Díaz-Burlinson, N., Gachet, S., 2004. Congruency analysis of species ranking based on leaf traits: which traits are the more reliable? Plant Ecol. 174, 37-48.
Semchenko, M., Lepik, M., Götzenberger, L., Zobel, K., 2012. Positive effect of shade on plant growth: amelioration of stress or active regulation of growth rate? J. Ecol. 100, 459-466.
Siefert, A., 2012. Incorporating intraspecific variation in tests of trait-based community assembly. Oecologia 170, 767-775.
Siefert, A., Violle, C., Chalmandrier, L., Albert, C.H., Taudiere, A., Fajardo, A., et al., 2015. A global meta-analysis of the relative extent of intraspecific trait variation in plant communities. Ecol. Lett. 18, 1406-1419.
Spasojevic, M.J., Suding, K.N., 2012. Inferring community assembly mechanisms from functional diversity patterns: the importance of multiple assembly processes. J. Ecol. 100, 652-661.
Sutherland, W.J., Freckleton, R.P., Godfray, H.C.J., Beissinger, S.R., Benton, T., Cameron, D.D., et al., 2013. Identification of 100 fundamental ecological questions. J. Ecol. 101, 58-67.
Swenson, N.G., Enquist, B.J., Thompson, J., Zimmerman, J.K., 2007. The influence of spatial and size scale on phylogenetic relatedness in tropical forest communities. Ecology 88, 1770-1780.
Taudiere, A., Violle, C., 2015. cati: an R package using functional traits to detect and quantify multi-level community assembly processes. Ecography 39, 699-708.
Vilà-Cabrera, A., Martínez-Vilalta-Vilalta, J., Retana, J., 2015. Functional trait variation along environmental gradients in temperate and Mediterranean trees. Glob. Ecol.Biogeogr. 24, 1377-1389.
Violle, C., Enquist, B.J., McGill, B.J., Jiang, L., Albert, C.H., Hulshof, C., et al., 2012. The return of the variance: intraspecific variability in community ecology. Trends Ecol. Evol. 27, 244-252.
Wright, I.J., Reich, P.B., Cornelissen, J.H., Falster, D.S., Garnier, E., Hikosaka, K., et al., 2005. Assessing the generality of global leaf trait relationships. New Phytol. 166, 485-496.
Wright, I.J., Reich, P.B., Westoby, M., Ackerly, D.D., Baruch, Z., Bongers, F., et al., 2004.The worldwide leaf economics spectrum. Nature 428, 821-827.

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Trait variation and functional diversity maintenance of understory herbaceous species coexisting along an elevational gradient in Yulong Mountain, Southwest China

Trait variation and functional diversity maintenance of understory herbaceous species coexisting along an elevational gradient in Yulong Mountain, Southwest China

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