Changes in species composition during the succession of ecological communities potentially reflect the differential effects of environmental filters and limiting similarity on structuring communities. As ecological succession can represent community assembly in action, understanding how successional time affects the functional and phylogenetic structure of communities can reveal the influence of different factors on the assembly process. We analysed functional patterns of multiple traits related to the succession of tropical forests to answer if there are trait convergence and/or divergence with regeneration age, and if functional and phylogenetic diversity can be predicted by forest age. We compiled checklists from studies of 23 successional forests in Brazilian Atlantic Forest, ranging from 4 to 120 years old. We also compiled functional traits for a total of 355 species. We analysed the data by a method that includes scaling-up trait-based data to the community level and matrix correlations of multiple traits. We built linear models to show the relationship between each trait and diversity (taxonomic, functional and phylogenetic) with successional time. We found no phylogenetic signal at the species pool and metacommunity levels, but significant trait divergence (tree guild, leaf slenderness, leaf area, pollination entomophilous generalist and pollination by vertebrate) and trait convergence (arboreal habitus, tree guild, leaf compoundness, pollination entomophilous generalist) patterns related to the successional gradient. Also, functional diversity increased during succession, with a significant increase in leaf slenderness and zoochoric dispersal and decrease in tree guild. Phylogenetic diversity also increased along the successional gradient. We found that the communities in the studied successional gradient are structured by both environmental (measured by trait convergence) and biotic (measured by trait divergence) filtering. The species turnover and diversification at taxonomic level are followed by well-defined patterns of trait turnover, revealing that community assembly is constrained by environmental filters at the beginning and by limiting similarity at the advanced stages of the succession.
Trait Convergence Assembly Pattern
Trait Divergence Assembly Pattern
A.P.G. III. 2009. An update of the Angiosperm Phylogeny Group classification for the orders and families of fowering plants: APG III. Bot. J. Linn. Soc. 161: 105–121.
Bazzaz, F.A., and S.T.A. Pickett. 1980. Physiological ecology of tropical succession: a comparative review. Annu. Rev. Ecol. Syst. 11: 287–310.
Bell, C. D., D. E. Soltis and P.S. Soltis. 2010. The age and diversification of the angiosperms re-revisited. Am. J. Bot. 97: 1296–1303.
Bhaskar, R., T. E. Dawson and P. Balvanera. 2014. Community assembly and functional diversity along sucession post-management. Funct. Ecol. 28: 1256–1265.
Blomberg, S.P. and T. Garland. 2002. Tempo and mode in evolution: phylogenetic inertia, adaptation and comparative methods. J. Evol. Biol. 15: 899–910.
Botta-Dukát, Z. 2005. Rao’s quadratic entropy as a measure of functional diversity based on multiple traits. J. Veg. Sci. 16: 533–540.
Campetella, G., Z. Botta-Dukát, C. Wellstein, R. Canullo, S. Gatto, S. Chelli, L. Mucina and S. Bartha. 2011. Patterns of plant trait-environment relationships along a forest succession chronose-quence. Agr. Ecosyst. Environ. 145: 38–48.
Carvalho, G.H., M.V. Cianciaruso and M.A. Batalha. 2010. Plantminer: A web tool for checking and gathering plant species taxonomic information. Environ. Modell. Softw. 25: 815–816.
Cavender-Bares, J., K.H. Kozak, P.V.A. Fine and S.W. Kembel. 2009. The merging of community ecology and phylogenetic biology. Ecol. Lett. 12, 693–715.
Chazdon: R.L. 2008. Chance and determinism in tropical forest succession. In: W.P. Carson and S.A. Schnitzer. (eds.), Tropical Forest Community Ecology. Wiley-Backwell, Oxford. pp. 384–408.
Clements, F.E. 1916. Plant Succession: An Analysis of the Development of Vegetation. Carnegie Institution of Washington, Washington.
Cornelissen, J.H.C., S. Lavorel, E. Garnier, S. Díaz, N. Buchmann, D.E. Gurvich, P.B. Reich, H. ter Steege, H.D. Morgan, M.G.A van der Heijden, J.G. Pausas and H. Poorter. 2003. A handbook of protocols for standardised and easy measurement of plant functional traits worldwide. Aust. J. Bot. 51: 335–380.
Devictor, V., D. Mouillot, C. Meynard, F. Jiguet, W. Thuiller and N. Mouquet. 2010. Spatial mismatch and congruence between taxonomic, phylogenetic and functional diversity: the need for integrative conservation strategies in a changing world. Ecol. Lett. 13: 1030–1040.
Díaz, S., M. Cabido and F. Casanoves. 1998. Plant functional traits and environmental filters at the regional scale. J. Veg. Sci. 9: 113–122.
Ding, Y., R. Zang, S.G. Letcher, S. Liu, and F. He. 2012. Disturbance regime changes the trait distribution, phylogenetic structure and community assembly of tropical rain forests. Oikos 121: 1263–1270.
Duarte, L.D.S., R.S. Bergamin, V. Marcilio-Silva, G.D.D.S. Seger and M.C.M. Marques. 2014. Phylobetadiversity among forest types in the Brazlian Atlantic forest complex. PLoS One 9: e105043.
Fukami, T., T.M. Bezemer, S.R. Mortimer, and W.H. Van der Putten. 2005. Species divergence and trait convergence in experimental plant community assembly. Ecol. Lett. 8: 1283–1290.
Garnier, E., J. Cortez, G. Billes, M.L. Navas, C. Roumet, M. Debussche, G. Laurent, A. Blanchard, D. Aubry, A. Bellmann, C. Neill and J.P. Toussaint. 2004. Plant functional markers capture ecosystem properties during secondary succession. Ecology 85: 2630–2637.
Gerhold, P., J.F. Cahill Jr., M. Winter, I.V. Bartish and A. Prinzing. 2015. Phylogenetic patterns are not proxy of community assembly mechanisms (they are far better). Funct. Ecol. 29: 600–614.
Girão, L.C., A.V. Lopes, M. Tabarelli and E.M. Bruna. 2007. Changes in tree reproductive traits reduce functional diversity in a fragmented Atlantic forest landscape. PLoS ONE 2, e908.
Götzenberger L., F. de Bello, K.A. Bråthen, et al. 2011. Ecological assembly rules in plant communities–approaches, patterns and prospects. Biol. Rev. 87: 111–27.
Guariguata, M.R. and R. Ostertag. 2001. Neotropical secondary forest succession: changes in structural and functional characteristics. Forest Ecol. Manag. 148: 185–206.
Herben, T. and D.E. Goldberg. 2014. Community assembly by limiting similarity vs. competitive hierarchies: testing the consequences of dispersion of individual traits. J. Ecol. 102: 156–166.
Herben, T., Z. Novakova, J. Klimesova and L. Hrouda. 2012. Species traits and plant performance: functional trade-offs in a large set of species in a botanical garden. J. Ecol. 100: 1522–1533.
Keddy, P.A. 1992. Assembly and response rules: two goals for predictive community ecology. J. Veg. Sci. 3: 157–164.
Kembel, S. and S.P. Hubbell. 2006. The phylogenetic structure of a neotropical forest tree community. Ecology 87: 86–99.
Lack, D. 1942. Ecological features of the bird faunas of British smallislands. J. Anim. Ecol. 11: 9–36.
Lebrija-Trejos, E., E.A. Pérez-García, J.A. Meave, F. Bongers and L. Poorter. 2010. Functional traits and environmental filtering drive community assembly in a species- rich tropical system. Ecology 91: 386–398.
Letcher, S.G. 2009. Phylogenetic structure of angiosperm communities during tropical forest succession. P. Roy. Soc. Lond. B. Bio. 277: 97–104.
Letcher, S.G., R.L. Chazdon, A.C.S. Andrade,F. Bongers, M. van Breugel, B. Finegan, S.G. Laurance, R.C.G. Mesquita, M. Martínez-Ramos and G.B. Williamson. 2012. Phylogenetic community structure during succession: Evidence from three Neotropical forest sites. Perspect. Plant Ecol. Evol. Syst. 14: 79–87.
Liebsch, D., M.C.M Marques and R. Goldenberg. 2008. How long does the Atlantic Rain Forest take to recover after a disturbance? Changes in species composition and ecological features during secondary succession. Biol. Conserv. 141: 1717–1725.
Lohbeck, M., L. Poorter, H. Paz, L. Plac, M. van Breugel, M. Martínez-Ramos and F. Bongers. 2012. Functional diversity changes during tropical forest succession. Perspect. Plant Ecol. Evol. Syst. 14: 89–96.
Lohbeck, M., L. Poorter, E. Lebrija-Trejos, M. Martínez-Ramos, J.A. Meave, H. Paz, E.A. Pérez-García, E. Romero-Pérez, A. Tauro and F. Bongers. 2013. Successional changes in functional composition contrast for dry and wet tropical forest. Ecology 94: 1211–1216.
Losos, J.B. 2008. Phylogenetic niche conservatism, phylogenetic signal andthe relationship between phylogenetic relatedness and ecological similarity among species. Ecol. Lett. 11: 995–1003.
MacArthur, R. and R. Levins. 1967. The limiting similarity, convergence, and divergence of coexisting species. Am. Nat. 101: 377–385.
Marques, M.C.M., M. Swaine and D. Liebsch. 2011. Diversity distribution and floristic differentiation of the coastal lowland vegetation: implications for the conservation of the Brazilian Atlantic Forest. Biodivers. Conserv. 20: 153–168.
Mayfield, M.M. and J.M. Levine. 2010. Opposing effects of competitive exclusion on the phylogenetic structure of communities. Ecol. Lett. 13: 1085–1093.
Meiners, S.J., M.W. Cadotte, J.D. Fridley, S.T.A. Pickett and L.R. Walker. 2015. Is successional research nearing its climax? New approaches for understanding dynamic communities. Funct. Ecol. 29: 154–164.
Mouquet, N., V. Devictor, C.N. Meynard,F. Munoz, L.F. Bersier, J. Chave, J. et al. 2012. Ecophylogenetics: advances and perspectives. Biol. Rev. 87: 769–785.
Mouquet, J.L. Moore and M. Loreau. 2002. Plant species richness and community productivity: why the mechanism that promotes coexistence matters. Ecol. Lett. 5: 56–65.
Oliveira-Filho, A.T. and M.A.L. Fontes. 2000. Patterns of floristic differentiation among Atlantic forests in south-eastern Brazil, and the influence of climate. Biotropica 32: 793–810.
Opler, P.A., H.G. Baker and G.W. Frankie. 1980. Plant reproductive characteristics during secondary succession in neotropical low-land forest ecosystems. Biotropica 12: 40–46.
Peterson, A.T. 2011. Ecological niche conservatism: a time-structure-dreview of evidence. J. Biogeogr. 38: 817–827.
Pillar, V.D. and L.D.S Duarte. 2010. A framework for metacommunity analysis of phylogenetic structure. Ecol. Lett. 13: 587–596.
Pillar, V.D., L.S. Duarte, E.E. Sosinski and F. Joner. 2009. Discriminating trait-convergence and trait-divergence assembly patterns in ecological community gradients. J. Veg. Sci. 20: 334–348.
Pillar, V.D.and E.E. Sosinski. 2003. An improved method for searching plant functional types by numerical analysis. J. Veg. Sci. 14: 323–332.
Prinzing, A., R. Reiffers, W.G. Braakhekke, S.M. Hennekens, O. Tackenberg, W.A. Ozinga et al. 2008. Less lineages – more trait variation:phylogenetically clustered plant communities are functionally morediverse. Ecol. Lett. 11: 809–819.
Purschke O., B.C. Schmid, M.T. Sykes, P. Poschlod, S.G. Michalski, W. Durka et al. 2013. Contrasting changes in taxonomic, phylogenetic and functional diversity during a long-term succession: insights into assembly processes. J. Ecol. 101: 857–866.
Ribeiro, M.C., J.P. Metzger, A.C. Martensen, F.J. Ponzoni and M.M. Hirota. 2009. The Brazilian Atlantic Forest: How much is left, and how is the remaining forest distributed? Implications for conservation. Biol. Conserv. 142: 1141–1153.
Ricotta, C. and M. Moretti. 2011. CWM and Rao’s quadratic diversity: a unified framework for functional ecology. Oecologia 167: 181–188.
Rozendaal, D.M.A., V.H. Hurtado and L. Poorter. 2006. Plasticity in leaf traits of 38 tropical tree species in response to light; relationships with light demand and adult stature. Funct. Ecol. 20: 207–216.
Scarano, F.R. 2002. Structure, function and floristic relationships of plants communities in stressful habitats marginal to Brazilian Atlantic Rainforest. Ann. Bot. 90: 517–524.
Shooner, S., C. Chisholm and T.J. Davies. 2015. The phylogenetics of succession can guide restoration: an example from abandoned mine sites in the subarctic. J. Appl. Ecol. 52: 1509–1517.
Swenson, N.G. 2013. The assembly of tropical tree communities – the advances and shortcomings of phylogenetic and functional trait analyses. Ecography 36: 264–276.
Srivastava, D.S., M.W. Cadotte,A.A.M. MacDonald, R.G. Marushia and N. Mirotchnick. 2012. Phylogenetic diversity and the functioning ofecosystems. Ecol. Lett. 15: 637–648.
Webb, C.O., D.D. Ackerly and S.W. Kembel. 2008. Phylocom: software for the analysis of phylogenetic community structure and character evolution. Bioinformatics 24: 2098–2100.
Webb, C.O. and D.R. Peart. 2000. Habitat associations of trees and seedlings in a Bornean rain forest. J. Ecol. 88: 464–78.
Webb, C.O., D.D. Ackerly, M.A. McPeek and M.J. Donoghue. 2002. Phylogenies and community ecology. Annu. Rev. Ecol. Syst. 33: 475–505.
Weiher, E. and P.A. Keddy. 1999. Ecological Assembly Rules: Perspectives, Advances, Retreats. Cambridge University Press, Cambridge.
Weiher, E., D. Freund, T. Bunton, A. Stefanski, T. Lee and S. Bentivenga. 2011. Advances, challenges and a developing synthesis of ecological community assembly theory. Philos. T. Roy. Soc. B 366: 2403–2413.
Whitfield, T. J. S., W.J. Kress, D.L. Erickson and G.D. Weiblen. 2012. Change in community phylogenetic structure during tropical forest succession: evidences from New Guinea. Ecography 35: 1–10.
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Marcilio-Silva, V., Pillar, V.D. & Marques, M.C.M. Functional turnover and community assemblage during tropical forest succession. COMMUNITY ECOLOGY 17, 88–97 (2016). https://doi.org/10.1556/168.2016.17.1.11
- Community assembly
- Environmental filtering
- Functional traits
- Trait convergence
- Trait divergence
- Tropical forest
- A.P.G. III (2009)