Functional trait and phylogenetic tests of community assembly across spatial scales in an Amazonian forest

Despite a long history of the study of tropical forests, uncertainty about the importance of different ecological processes in shaping tropical tree species distributions persists. Trait- and phylogenetic-based tests of community assembly provide a powerful way to detect community assembly processes but have seldom been applied to the same community. Both methods are well suited to testing how the relative importance of different ecological processes changes with spatial scale. Here we apply both methods to the Yasuni Forest Dynamics Plot, a 25-ha Amazonian forest with >1100 tree species. We found evidence for habitat filtering from both trait and phylogenetic methods from small (25 m2) to intermediate (10 000 m2) spatial scales. Trait-based methods detected even spacing of strategies, a pattern consistent with niche partitioning or enemy-mediated density dependence, at smaller spatial scales (25–400 m2). Simulation modeling of community assembly processes suggests that low statistical power to detect eve...

[1]  J. Connell On the role of the natural enemies in preventing competitive exclusion in some marine animals and in rain forest trees , 1971 .

[2]  R. Condit,et al.  Tree species distributions and local habitat variation in the Amazon: large forest plot in eastern Ecuador , 2004 .

[3]  James Rosindell,et al.  Unified neutral theory of biodiversity and biogeography , 2010, Scholarpedia.

[4]  Dolph Schluter,et al.  Species diversity in ecological communities: historical and geographical perspectives. , 1993 .

[5]  Paul A. Keddy,et al.  Community Assembly Rules, Morphological Dispersion, and the Coexistence of Plant Species , 1998 .

[6]  M. Westoby,et al.  Plant height and evolutionary games , 2003 .

[7]  Michael D. Abràmoff,et al.  Image processing with ImageJ , 2004 .

[8]  P. Grubb THE MAINTENANCE OF SPECIES‐RICHNESS IN PLANT COMMUNITIES: THE IMPORTANCE OF THE REGENERATION NICHE , 1977 .

[9]  Sean C. Thomas,et al.  Ontogenetic changes in leaf size in Malaysian rain forest trees , 1995 .

[10]  Daniel R. Lynch,et al.  Nonrandom Processes Maintain Diversity in Tropical Forests , 2006 .

[11]  Janneke HilleRisLambers,et al.  The importance of niches for the maintenance of species diversity , 2009, Nature.

[12]  J. Bastow Wilson,et al.  Evidence for limiting similarity in a sand dune community , 2004 .

[13]  J. Terborgh The Vertical Component of Plant Species Diversity in Temperate and Tropical Forests , 1985, The American Naturalist.

[14]  D. A. King,et al.  The contribution of interspecific variation in maximum tree height to tropical and temperate diversity , 2005, Journal of Tropical Ecology.

[15]  Nathan J B Kraft,et al.  Functional Traits and Niche-Based Tree Community Assembly in an Amazonian Forest , 2008, Science.

[16]  David Tilman,et al.  Niche tradeoffs, neutrality, and community structure: a stochastic theory of resource competition, invasion, and community assembly. , 2004, Proceedings of the National Academy of Sciences of the United States of America.

[17]  M. Westoby,et al.  Alternative height strategies among 45 dicot rain forest species from tropical Queensland, Australia , 2005 .

[18]  K. Niklas Morphological evolution through complex domains of fitness. , 1994, Proceedings of the National Academy of Sciences of the United States of America.

[19]  Stephen P Hubbell,et al.  The phylogenetic structure of a neotropical forest tree community. , 2006, Ecology.

[20]  Paul A. Keddy,et al.  Assembly rules, null models, and trait dispersion : new questions from old patterns , 1995 .

[21]  B. Enquist,et al.  Opposing assembly mechanisms in a neotropical dry forest: implications for phylogenetic and functional community ecology. , 2009, Ecology.

[22]  T. Garland,et al.  TESTING FOR PHYLOGENETIC SIGNAL IN COMPARATIVE DATA: BEHAVIORAL TRAITS ARE MORE LABILE , 2003, Evolution; international journal of organic evolution.

[23]  Howard Rogers Tropical Forest Diversity and Dynamism , 2005 .

[24]  Campbell O. Webb,et al.  Phylogenies and Community Ecology , 2002 .

[25]  M. Lechowicz,et al.  Alternative Designs and the Evolution of Functional Diversity , 2005, The American Naturalist.

[26]  Campbell O. Webb,et al.  Phylodiversity-dependent seedling mortality, size structure, and disease in a Bornean rain forest. , 2006, Ecology.

[27]  J. Cavender-Bares,et al.  Phylogenetic Overdispersion in Floridian Oak Communities , 2004, The American Naturalist.

[28]  D. Ackerly,et al.  A trait-based test for habitat filtering: convex hull volume. , 2006, Ecology.

[29]  P. Reich,et al.  A handbook of protocols for standardised and easy measurement of plant functional traits worldwide , 2003 .

[30]  T. Kohyama,et al.  Size-structured tree populations in gap-dynamic forest-the forest architecture hypothesis for the stable coexistence of species , 1993 .

[31]  J. Shurin,et al.  Trait-based assembly and phylogenetic structure in northeast Pacific rockfish assemblages. , 2009, Ecology.

[32]  S. Hubbell,et al.  Recruitment Near Conspecific Adults and the Maintenance of Tree and Shrub Diversity in a Neotropical Forest , 1992, The American Naturalist.

[33]  N. Gotelli,et al.  NULL MODELS IN ECOLOGY , 1996 .

[34]  Peter B Adler,et al.  A niche for neutrality. , 2007, Ecology letters.

[35]  P. Dayton,et al.  Role of biological disturbance in maintaining diversity in the deep sea , 1972 .

[36]  Stuart J. Davies,et al.  Growth and mortality are related to adult tree size in a Malaysian mixed dipterocarp forest , 2006 .

[37]  R. Macarthur,et al.  The Limiting Similarity, Convergence, and Divergence of Coexisting Species , 1967, The American Naturalist.

[38]  Edward Naranjo,et al.  SAFETY IN DIVERSITY , 2008 .

[39]  Brian J Enquist,et al.  The influence of spatial and size scale on phylogenetic relatedness in tropical forest communities. , 2007, Ecology.

[40]  P. Coley,et al.  Divergent Defensive Strategies of Young Leaves in Two Species of Inga , 2005 .

[41]  J. Cavender-Bares,et al.  The merging of community ecology and phylogenetic biology. , 2009, Ecology letters.

[42]  Olivier J. Hardy,et al.  Testing the spatial phylogenetic structure of local communities: statistical performances of different null models and test statistics on a locally neutral community , 2008 .

[43]  Steven W Kembel,et al.  Disentangling niche and neutral influences on community assembly: assessing the performance of community phylogenetic structure tests. , 2009, Ecology letters.

[44]  Peter A. Abrams,et al.  The theory of limiting similarity , 1983 .

[45]  Barbara R. Holland,et al.  Analysis of Phylogenetics and Evolution with R , 2007 .

[46]  Jeannine Cavender-Bares,et al.  Phylogenetic structure of Floridian plant communities depends on taxonomic and spatial scale. , 2006, Ecology.

[47]  Campbell O. Webb,et al.  Bioinformatics Applications Note Phylocom: Software for the Analysis of Phylogenetic Community Structure and Trait Evolution , 2022 .

[48]  David D. Ackerly,et al.  Community assembly and shifts in plant trait distributions across an environmental gradient in coastal California , 2009 .

[49]  J. Connell Diversity in tropical rain forests and coral reefs. , 1978, Science.

[50]  Campbell O. Webb,et al.  Picante: R tools for integrating phylogenies and ecology , 2010, Bioinform..

[51]  Stephen P. Hubbell,et al.  Neutral theory in community ecology and the hypothesis of functional equivalence , 2005 .

[52]  S. Hubbell,et al.  Spatial and temporal variation of biomass in a tropical forest: results from a large census plot in Panama , 2003 .

[53]  A. G. Valk,et al.  Succession in Wetlands: A Gleasonian Appraoch , 1981 .

[54]  B. Emerson,et al.  Phylogenetic analysis of community assembly and structure over space and time. , 2008, Trends in ecology & evolution.

[55]  Lindsay A. Turnbull,et al.  Why equalising trade-offs aren't always neutral. , 2008, Ecology letters.

[56]  D. Janzen,et al.  Herbivores: Their Interaction With Secondary Plant Metabolites , 1982 .

[57]  S. Hubbell,et al.  A unified theory of biogeography and relative species abundance and its application to tropical rain forests and coral reefs , 1997, Coral Reefs.

[58]  Brian J. McGill,et al.  Null Versus Neutral Models: What's The Difference? , 2006 .

[59]  John P. O'Neill,et al.  A MORPHOLOGICAL APPROACH TO THE STUDY OF AVIAN COMMUNITY ORGANIZATION , 2003 .

[60]  Campbell O. Webb,et al.  Phylomatic: tree assembly for applied phylogenetics , 2005 .

[61]  J. A. Barone,et al.  HERBIVORY AND PLANT DEFENSES IN TROPICAL FORESTS , 1996 .

[62]  Robert John,et al.  Nonrandom Processes Maintain Diversity in Tropical Forests , 2006, Science.

[63]  A. Shmida,et al.  Biological determinants of species diversity , 1985 .

[64]  Campbell O. Webb,et al.  Emerging patterns in the comparative analysis of phylogenetic community structure , 2009, Molecular ecology.

[65]  Kyle E. Harms,et al.  Pervasive density-dependent recruitment enhances seedling diversity in a tropical forest , 2000, Nature.

[66]  Campbell O. Webb,et al.  Exploring the Phylogenetic Structure of Ecological Communities: An Example for Rain Forest Trees , 2000, The American Naturalist.

[67]  Campbell O. Webb,et al.  Regional and phylogenetic variation of wood density across 2456 Neotropical tree species. , 2006, Ecological applications : a publication of the Ecological Society of America.

[68]  Sean C. Thomas,et al.  The worldwide leaf economics spectrum , 2004, Nature.

[69]  J. X. Becerra The impact of herbivore–plant coevolution on plant community structure , 2007, Proceedings of the National Academy of Sciences.

[70]  O. Pellmyr Plant-Animal Interactions: Evolutionary Ecology in Tropical and Temperate Regions , 1991 .

[71]  S. Hubbell,et al.  Short-term dynamics of a neotropical forest: why ecological research matters to tropical conservation and management , 1992 .

[72]  B. Nicholas The ecology of trees in the tropical rain forest , 2004 .

[73]  M. Cadotte Ecological Niches: Linking Classical and Contemporary Approaches , 2004, Biodiversity & Conservation.

[74]  P. Abrams Limits to the Similarity of Competitors Under Hierarchical Lottery Competition , 1996, The American Naturalist.

[75]  J. Zimmerman,et al.  The problem and promise of scale dependency in community phylogenetics. , 2006, Ecology.

[76]  S. Pacala,et al.  Ecological drift in niche-structured communities: neutral pattern does not imply neutral process , 2005 .

[77]  David D. Ackerly,et al.  Limiting similarity and functional diversity along environmental gradients , 2005 .

[78]  Nathan J B Kraft,et al.  Response to Comment on “Functional Traits and Niche-Based Tree Community Assembly in an Amazonian Forest” , 2009, Science.

[79]  R. Dirzo,et al.  Herbivores. Their Interaction with Secondary Plant Metabolites. , 1981 .

[80]  M. Westoby,et al.  ECOLOGICAL STRATEGIES : Some Leading Dimensions of Variation Between Species , 2002 .

[81]  Stephen P Hubbell,et al.  Neutral theory and the evolution of ecological equivalence. , 2006, Ecology.

[82]  P. Reich,et al.  From tropics to tundra: global convergence in plant functioning. , 1997, Proceedings of the National Academy of Sciences of the United States of America.

[83]  B. McGill,et al.  Community inertia of Quaternary small mammal assemblages in North America. , 2005, Proceedings of the National Academy of Sciences of the United States of America.

[84]  H. A. Peters Neighbour‐regulated mortality: the influence of positive and negative density dependence on tree populations in species‐rich tropical forests , 2003 .

[85]  P. Chesson Mechanisms of Maintenance of Species Diversity , 2000 .

[86]  J. Svenning,et al.  Microhabitat specialization in a species‐rich palm community in Amazonian Ecuador , 1999 .

[87]  Campbell O. Webb,et al.  Trait Evolution, Community Assembly, and the Phylogenetic Structure of Ecological Communities , 2007, The American Naturalist.

[88]  P. Coley,et al.  Herbivores Promote Habitat Specialization by Trees in Amazonian Forests , 2004, Science.

[89]  Why Do Some Tropical Forests Have So Many Species of Trees? , 2004 .

[90]  Stephen P. Hubbell,et al.  Habitat associations of trees and shrubs in a 50‐ha neotropical forest plot , 2001 .

[91]  Campbell O. Webb,et al.  Phylogenetic signal in plant pathogen–host range , 2007, Proceedings of the National Academy of Sciences.

[92]  D. Janzen Herbivores and the Number of Tree Species in Tropical Forests , 1970, The American Naturalist.

[93]  D. Goldberg,et al.  Equivalence of competitors in plant communities: a null hypothesis and a field experimental approach. , 1983 .

[94]  Mark W. Chase,et al.  Evolution of the angiosperms: calibrating the family tree , 2001, Proceedings of the Royal Society of London. Series B: Biological Sciences.