Abstract
Density dependence, environmental sorting and chance have been discussed for the purpose of understanding, predicting and explaining the species richness, composition and structural parameters of living communities. Different ecological mechanisms occur individually in an overlapping manner, so the structure of each local community is influenced by an independent mixture of these factors. To identify which of these factors prevails in organizing the species-rich tree community from 100 plots of 10 × 10 m in a primary forest patch (the Forest of Seu Nico – FSN, from the Atlantic Forest domain), we analyzed species-environment correlations via canonical correspondence analysis and identified two different pedo-environments. We analyzed the community’s phylogenetic structure using Phylocom 4.2 software to calculate the net relatedness index (NRI) and the nearest taxon index (NTI). Furthermore, we partitioned the total phylogenetic diversity into independent α and β components (ΠST). To reveal the overlap of ecological mechanisms such as neutrality, environmental filtering and density-dependent factors, we analyzed the phylogenetic structure in both pedo-environments. The species-environment correlations observed in the FSN are weak in comparison with those found in other studies, although the permanent plot presents a short environmental gradient, dividing the plot into an upper, more acidic hillside and a lower, more fertile bottom. The overall phylogenetic structure of the FSN community shows strong and significant phylogenetic overdispersion. This overdispersion indicates that density-dependent factors, such as interspecific competition, play an important role in maintaining the species richness and community structure in megadiverse ecosystems such as the FSN when we assume traits to be conserved within evolutionary lineages. The NRI and NTI are correlated positively with the soil pH and negatively with the soil’s aluminum concentration, so the bottom plots show higher phylogenetic overdispersion and lower ΠST values than the hillside plots. This pattern can be explained by the greater importance of environmental filters in more acidic soils that form less favorable habitats, while the influence of competition and therefore also the rate of competitive exclusion are higher in the more favorable, less acidic plots.
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References
APG (Angiosperm Phylogeny Group) III (2009) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Bot J Linn Soc 161:105–121
Baldeck CA, Harms KE, Yavitt YB, John R, Turner BL, Valencia R, Navarrete H, Davies SJ, Chuyong GB, Kenfack D, Thomas DW, Madawala S, Gunatilleke N, Gunatilleke S, Bunyavejchewin S, Kiratiprayoon S., Yaacob A, Nur Supardi MA, Dalling JW (2013) Soil resources and topography shape local tree community structure in tropical forests. Proc Roy Soc B 2013 280, 20122532, doi:10.1098/rspb.2012.2532
Baraloto C, Hardy OJ, Paine CET, Dexter KG, Cruaud C, Dunning LT, Gonzalez M-A, Molino J-F, Sabatier D, Savolainen V, Chave J (2012) Using functional traits and phylogenetic trees to examine the assembly of tropical tree communities. J Ecol 100:690–701
Botrel RT, Oliveira-Filho AT, Rodrigues LA, Curi N (2002) Influência do solo e topografia sobre as variações da composição florística e estrutura da comunidade arbóreo-arbustiva de uma Floresta Estacional Semidecidual em Ingaí, MG (Influence of soils and topography on the variations of species composition and structure of the community of trees and shrubs of a tropical semideciduous forest in Ingaí, southeastern Brazil). Brasil Rev Bot 25:195–213
Campos EP, Silva AF, Meira Neto JAA, Martins SV (2006) Floristica e estrutura horizontal da vegetação arbórea de uma ravina em um fragmento florestal no município de Viçosa, MG (Floristics and horizontal structure of the tree vegetation of a ravine in a forest fragment in the municipality of Viçosa, MG). R Árvore 30:1045–1054
Carvalho DAC, Oliveira-Filho AT, Vilela EA, Curi N, van den Berg E, Fontes MAL, Botezelli L (2005) Distribuição de espécies arbóreo-arbustivas ao longo de um gradiente de solos e topografia em um trecho de floresta ripária do Rio São Francisco em Três Marias, MG, Brasil (Distribution of tree and shrub species along a gradient of soils and topography in a strip of riparian forest of the São Francisco river in Três Marias, MG, Brazil). Brasil Rev Bot 28:329–345
Cavender-Bares J, Keen A, Miles B (2006) Phylogenetic structure of Floridian plant communities depends on taxonomic and spatial scale. Ecology 87:S109–S122
Cavender-Bares J, Ackerly DD, Baum DA, Bazzaz FA (2004) Phylogenetic overdispersion in Floridian oak communities. Amer Naturalist 163:823–843
Cavender-Bares J, Kozak KH, Fine PVA, Kembel SW (2009) The merging of community ecology and phylogenetic biology. Ecol Lett 12:693–715
Chave J (2009) Competition, Neutrality, and Community Organization. In Levin SA (ed) The Princeton guide to ecology. Princeton University Press, Princeton, pp 264–273
Chave J, Alonso D, Etienne RS (2006) Comparing models of species abundance. Nature 441:E1
Chave J, Riera B, Dubois MA (2001) Estimation of biomass in a neotropical forest of French Guiana: spatial and temporal variability. J Trop Ecol 17:79–96
Clayton DH, Bush SE (2006) The role of body size in host specificity: Reciprocal transfer experiments with feather lice. Evolution 60:2158–2167
Colwell RK, Coddington JA (1994) Estimating terrestrial biodiversity through extrapolation. Philos Trans, Ser B 345:101–108
Connell JH (1971) On the role of natural enemies in preventing competitive exclusion in some marine animals and in rain Forest trees. In Boer PJ, Gradwell GR (eds) Dynamics of populations. Center for Agricultural Publishing and Documentation, Wageningen, pp 198–310
Davies TJ, Barralough TG, Chase MW, Soltis PS, Soltis DE, Savolainen V (2004) Darwin’s abominable mystery: Insights from a supertree of the angiosperms. Proc Natl Acad Sci, USA 101:1904–1909
Diamond JM (1975) Assembly of species communities. In Cody ML, Diamond JM (eds) Ecology and evolution of communities. Harvard University, Press, Cambridge, Massachusetts, pp 342–444
Díaz S, Hodgson JG, Thompson K., Cabido M, Cornelissen JHC et al. (2004) The plant traits that drive ecosystems: Evidence from three continents. J Veg Sci 15:295–304
Elton CS (1927) Animal ecology. Sidgwick and Jackson, London
Emerson BC, Gillespie RG (2008) Phylogenetic analysis of community assembly and structure over space and time. Trends Ecol Evol 23:619–630
Ernest SKM, White EP, Brown JH (2009) Changes in a tropical forest support metabolic zero-sum dynamics. Ecol Lett 12:507–515
Ferreira Jr. WG, Silva AF, Meira-Neto JAA, Schaefer CEGR, Dias AS, Ignácio M, Medeiros MCMP (2007) Composição florística da vegetação arbórea de um trecho de Floresta Estacional Semidecídua em Viçosa, Minas Gerais, e espécies de maior ocorrência na região (Floristic composition of trees in a seasonal semideciduous forest in Viçosa, Minas Gerais, and species of greater occurrence in the region). R Árvore 31:1121–1130
Fine PVA, Mesones I, Coley PD (2004) Herbivores promote habitat specialization by trees in Amazonian forests. Science 305:663–665
Fine PVA, Miller ZJ, Mesones I, Irazuzta S, Appel HM, Stevens MHH et al. (2006). The growth-defense trade-off and habitat specialization by plants in Amazonian forests. Ecology 87:S150–S162
Forest F, Grenyer R, Rouget M, Davies TJ, Cowling RM, Faith DP et al. (2007) Preserving the evolutionary potential of floras in biodiversity hotspots. Nature 455:757–760
Forzza RC (coord) (2013) Lista de Espécies da Flora do Brasil. Online access at: http://floradobrasil.jbrj.gov.br/2012/. Accessed: March 2013
Friis EM, Crane PR, Petersen KR (2011) Early flowers and angiosperm evolution. Cambridge University Press, Cambridge
Gastauer M, Meira-Neto JAA (2013) Avoiding inaccuracies in tree calibration and phylogenetic community analysis using Phylocom 4.2. Ecol Informatics 15:85–90
Gause GF (1932) Experimental studies on the struggle for existence: 1. Mixed population of two species of yeast. J Exp Biol 9:389–402
Gause GF (1934) The struggle for existence. Williams & Wilkins, Baltimore
Grime JP (1979) Plant strategies and vegetation processes. John Wiley & Sons, Chichester
Grime JP (2001) Plant Strategies, vegetation processes, and ecosystem properties. Ed. 2. John Wiley & Sons, Chichester
Grinnell J (1917) The niche-relationships of the California Trasher. Auk 34:544–556
Haegeman B, Etienne RS (2011) Independent species in independent niches behave neutrally. Oikos 120:961–963
Hardin G (1960) The Competitive Exclusion Principle. Science 131:1292–1297
Hardy OJ (2008) Testing the spatial phylogenetic structure of local communities: statistical performances of different null models and test statistics on a locally neutral community. J Ecol 96:914–926
Hardy OJ (2009) SPACoDi 0.10: a program for spatial and phylogenetic analysis of community diversity. Available at: http://ebe.ulb.ac.be/ebe/Software.html
Hardy OJ, Jost L (2008) Interpreting and estimating measures of community phylogenetic structuring. J Ecol 96:849–852
Hardy OJ, Senterre B (2007) Characterizing the phylogenetic structure of communities by an additive partitioning of phylogenetic diversity. J Ecol 95:493–506
Harms KE, Condit R, Hubbell SP, Foster RB (2001) Habitat associations of trees and shrubs in a 50-ha neotropical forest plot. J Ecol 89:947–959
Hubbell SP (2001) The unified neutral theory of biodiversity and biogeography. Princeton University Press, Oxford
Hutchinson GE (1957) Concluding remarks. Cold Spring Harbor Symp Quant Biol 22:415–427
Janzen DH (1970) Herbivores and the number of tree species in tropical forests. Amer Naturalist 104:501–528
Kelly CK, Bowler MG, Pybus O, Harvey PH (2008) Phylogeny, niches, and relative abundance in natural communities. Ecology 89:962–970
Kembel SW, Hubbell SP (2006) The phylogenetic structure of a neotropical forest tree community. Ecology 87:S86–S99
Kochian LV, Piñeros MA, Hoekenga OA (2005) The physiology, genetics and molecular biology of plant aluminum resistance and toxicity. Pl & Soil 274:175–195
Kozak KH, Graham CH, Wiens JJ (2008) Integrating GIS-based environmental data into evolutionary biology. Trends Ecol Evol 23:141–148
Kress WJ, Erickson DL, Jones FA, Swenson NG, Perez R, Sanjur O, Bermingham E (2009) Plant DNA barcodes and a community phylogeny of a tropical forest dynamics plot in Panama. Proc Natl Acad Sci, USA 106:18621–18626
Leibold, MA (2009) Spatial and metacommunity dynamics in biodiversity. In Levin SA (ed) The Princeton guide to ecology. Princeton University Press, Princeton
Leibold MA, McPeek MA (2006) Coexistence of the niche and neutral perspectives in community ecology. Ecology 87:1399–1410
Letcher SG (2010) Phylogenetic structure of angiosperm communities during tropical forest succession. Proc Roy Soc London, Ser B, Biol Sci 277:97–104
MacArthur RH (1958) Population ecology of some warblers of northeastern coniferous forests. Ecology 39:599–619
MacArthur R, Levins R (1967) The limiting similarity, convergence, and divergence of coexisting species. Amer Naturalist 101:377–385
Mayfield MM, Levine JM (2010) Opposing effects of competitive exclusion on the phylogenetic structure of communities. Ecol Lett 13:1085–1093
McCune B, Mefford MJ (1999) PC-ORD version 4.0, multivariate analysis of ecological data: User’s guide. MjM Software Design, Glaneden Beach
Neri AV, Schaefer CEGR, Silva AF, Souza AL, Ferreira-Junior WG, Meira-Neto JAA (2012) The influence of soils on the floristic composition and community structure of an area of Brazilian Cerrado vegetation. Edinburgh J Bot 69:1–27
Newton AC (2007) Forest ecology and conservation: a handbook of techniques. Oxford University Press, Oxford
Oliveira-Filho AT, Fontes MAL (2000) Patterns of Floristic Differentiation among Atlantic Forests in Southeastern Brazil and the Influence of Climate. Biotropica 32:793–810
Paine CET, Baraloto C, Chave J, Hérault B (2011) Functional traits of individual trees reveal ecological constraints on community assembly in tropical rain forests. Oikos 120:720–727
Paine CET, Norden N, Chave J, Forget P-M, Fortunel C (2012) Phylogenetic density dependence and environmental filtering predict seedling mortality in a tropical forest. Ecol Lett 15:34–41
Paula AL, Silva AF, Marco-Jr. P, Santos FAM, Souza AL (2004) Sucessão ecológica da vegetação arbóres em uma Floresta Estacional Semidecidual, Viçosa, MG, Brasil (Ecological succession of a tree community of semideciduous seasonal forest in Viçosa, Minas Gerais State, Brazil). Acta Bot Brasil 18:407–423
Peel MC, Finlayson BL, McMahon TA (2007) Updated world map of the Köppen-Geiger climate classification. Hydrol Earth System Sci 11:1633–1644
Pei N, Lian J-Y, Erickson DL, Swenson NG, Kress WJ, Ye W-H, Ge X-J (2011) Exploring tree-habitat associations in a chinese subtropical forest plot using a molecular phylogeny generated from DNA barcode loci. PLoS ONE 6: e21273
Pinto SIP, Martins SV, Silva AG, Barros NF, Dias HCT, Scoss LM (2007) Estrutura de componente arbustivo-arbóreo de dois estádios successionais de Floresta Estacional Semidecidual na Reserva Florestal Mata do Paraíso, Viçosa, MG, Brasil (Structure of the tree-shrub component in two successional stages of semideciduous forest in the Mata do Paraíso Forest Reserve, Viçosa, MG, Brazil). R. Árvore 31:823–833
Punchi-Manage R, Getzin S, Wiegand T, Kanagaraj R, Savitri Gunatilleke SV, Gunatilleke IAUN, Wiegand K, Huth A (2013) Effects of topography on structuring local species assemblages in a Sri Lankan mixed dipterocarp forest. J Ecol 101:149–160
Quesada CA, Phillips OL, Schwarz M, Czimczik CI, Baker TR et al. (2012) Basin-wide variations in Amazon forest structure and function are mediated by both soils and climate. Biogeosciences 9:2203–2246
Ribas RF, Meira-Neto JAA, Silva AF, Souza AL (2003) Composição florística da dois trechos em diferentes etapas serais de uma Floresta Estacional Semidecidual em Viçosa, Minas Gerais (Floristic composition of two sites in different successional stages of Semideciduous Seasonal Montane Forest in Viçosa, Minas Gerais). R Árvore 27:821–830
Rosenzweig ML (1995) Species diversity in space and time. Cambridge University Press, Cambridge
Rosindell J, Hubbell SP, Etienne RS (2011) The unified neutral theory of biodiversity and biogeography at age ten. Trends Ecol Evol 26:340–348
Saporetti Jr AW, Schaefer CEGR, Souza AL, Soares MP, Araújo DSD, Meira Neto JAA (2012) Influence of soil physical properties on plants of the Mussununga ecosystem, Brazil. Folia Geobot 47:29–39
Sargent RD, Ackerly DD (2008) Plant-pollinator interactions and the assembly of plant communities. Trends Ecol Evol 23:123–130
Shipley B, Paine CET, Baraloto C (2012) Quantifying the importance of local niche-based and stochastic processes to tropical tree community assembly. Ecology 93:760–769
Simberloff DS (1970) Taxonomic diversity of island biotas. Evolution 24:23–47
Souza JS, Espírito-Santo FDB, Fontes MAL, Oliveira-Filho AT, Botezelli L (2003) Análise das variações florísticas e estruturais da comunidade arbórea de um fragmento de Floresta Semidecídua às margens do Rio Capivari, Lavras-MG (Analyses of the floristic and structural variations of the tree community in a fragment of Semidecidual Forest at the margin of the Capivari River, Lavras-MG). R Árvore 27:185–206
Sukri RSH, Wahab RA, Salim KA, Burslem DFRP (2012) Habitat associations and community structure of dipterocarps in response to environmental and soil conditions in Brunei Darussalam, northwest Borneo. Biotropica 44:595–605
Swenson NG (2009) Phylogenetic resolution and quantifying the phylogenetic diversity and dispersion of communities. PLoS ONE 4:e4390
Swenson NG, Enquist BJ (2009) Opposing assembly mechanisms in a Neotropical dry forest: implications for phylogenetic and functional community ecology. Ecology 90:2161–2170
Ter Braak CJF (1988) CANOCO – A FORTRAN program for canonical community ordination. Technical report LWA-88-2, TNO, Institute of Applied Computer Science, Wageningen
Tilman D (1985) The resource-ratio hypothesis of plant succession. Amer Naturalist 125:439–464
van der Valk AG (1981) Succession in wetlands – a Gleasonian approach. Ecology 62:688–696
Valiente-Banuet A, Verdú M (2007) Facilitation can increase the phylogenetic diversity of plant communities. Ecol Lett 10:1029–1036
Veloso HP, Rangel Filho ALR, Lima JCA (1991) Classificação da vegetação Brasileira, adaptada a um sistema universal. IBGE, Rio de Janeiro
Verdú M, Pausas JG (2007) Fire drives phylogenetic clustering in Mediterranean Basin woody plant communities. J Ecol 95:1316–1323
Webb CO (2000) Exploring the phylogenetic structure of ecological communities: an example for rain forest trees. Amer Naturalist 156:145–155
Webb CO, Donoghue MJ (2005) Phylomatic: tree assembly for applied phylogenetics. Molec Ecol Notes 5:181–183
Webb CO, Ackerly DD, Kembel SW (2008) Phylocom: software for the analysis of phylogenetic community structure and character evolution. Bioinformatics 24:2098–2100
Webb CO, Ackerly DD, Kembel S (2011) Software for the analysis of phylogenetic community structure and character evolution (with phylomatic and ecovolve). Users Manual. Available at: http://phylodiversity.net/phylocom/. Accessed: September 5, 2012
Webb CO, Ackerly DD, McPeek MA, Donaghue MJ (2002) Phylogenies and community ecology. Annual Rev Ecol Syst 33:475–505
Weiher E, Keddy PA (1995) Assembly rules, null models, and trait dispersion – new questions from old patterns. Oikos 74:159–164
Whitfeld TJS, Kress WJ, Erickson DL, Weiblen GD (2012) Change in community phylogenetic structure during tropical forest succession: evidence from New Guinea. Ecography 35:1–10
Wikstrom N, Savolainen V, Chase MW (2001) Evolution of angiosperms: Calibrating the family tree. Proc Roy Soc London, Ser B, Biol Sci 268:2211–2220
Zimmermann JK, Thompson J, Brakow N (2008) Large tropical forest dynamics plots: testing explication for the maintenance of species diversity. In Carson PC, Schnitzer AS (eds) Tropical forest community ecology. Blackwell Publishing Ltd, Chichester, pp 98–118
Acknowledgements
We are grateful to Suzano Pulp and Paper for a Ph.D. scholarship for one of the authors (MG) to carry out the presented research. We also thank FAPEMIG, Floresta Escola Project, MCTI, CAPES and CNPq for scholarship (JAAMN) and support. We thank David Irsigler for surveying the tree community in the FSN plot. For helping to identify plant taxa, we thank Gilmar E. Valente, Pedro Paulo de Souza, José Martins Fernandes, Victor Peçanha de Miranda Coelho, Priscila Bezerra de Souza, Michellia Pereira Soares, Walnir Gomes Ferreira Jr., Márcio Luiz Batista and Amilcar Walter Saporetti Jr. Joerg Ewald, Jerome Chave, C. E. Timothy Paine and two anonymous reviewers are gratefully acknowledged for valuable comments on the manuscript.
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Gastauer, M., Meira-Neto, J.A.A. Interactions, Environmental Sorting and Chance: Phylostructure of a Tropical Forest Assembly. Folia Geobot 49, 443–459 (2014). https://doi.org/10.1007/s12224-013-9181-1
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DOI: https://doi.org/10.1007/s12224-013-9181-1