Explaining variation in tropical plant community composition: influence of environmental and spatial data quality

The degree to which variation in plant community composition (beta-diversity) is predictable from environmental variation, relative to other spatial processes, is of considerable current interest. We addressed this question in Costa Rican rain forest pteridophytes (1,045 plots, 127 species). We also tested the effect of data quality on the results, which has largely been overlooked in earlier studies. To do so, we compared two alternative spatial models [polynomial vs. principal coordinates of neighbour matrices (PCNM)] and ten alternative environmental models (all available environmental variables vs. four subsets, and including their polynomials vs. not). Of the environmental data types, soil chemistry contributed most to explaining pteridophyte community variation, followed in decreasing order of contribution by topography, soil type and forest structure. Environmentally explained variation increased moderately when polynomials of the environmental variables were included. Spatially explained variation increased substantially when the multi-scale PCNM spatial model was used instead of the traditional, broad-scale polynomial spatial model. The best model combination (PCNM spatial model and full environmental model including polynomials) explained 32% of pteridophyte community variation, after correcting for the number of sampling sites and explanatory variables. Overall evidence for environmental control of beta-diversity was strong, and the main floristic gradients detected were correlated with environmental variation at all scales encompassed by the study (c. 100–2,000 m). Depending on model choice, however, total explained variation differed more than fourfold, and the apparent relative importance of space and environment could be reversed. Therefore, we advocate a broader recognition of the impacts that data quality has on analysis results. A general understanding of the relative contributions of spatial and environmental processes to species distributions and beta-diversity requires that methodological artefacts are separated from real ecological differences.

[1]  David B. Clark,et al.  EDAPHIC FACTORS AND THE LANDSCAPE-SCALE DISTRIBUTIONS OF TROPICAL RAIN FOREST TREES , 1999 .

[2]  Kalle Ruokolainen,et al.  Dispersal, Environment, and Floristic Variation of Western Amazonian Forests , 2003, Science.

[3]  Petr Šmilauer,et al.  CANOCO 4.5 Reference Manual and CanoDraw for Windows User's Guide: Software for Canonical Community Ordination , 2002 .

[4]  R. Busing,et al.  The Unified Neutral Theory of Biodiversity and Biogeography , 2002 .

[5]  B. Gilbert,et al.  FERN COMMUNITY ASSEMBLY: THE ROLES OF CHANCE AND THE ENVIRONMENT AT LOCAL AND INTERMEDIATE SCALES , 2005 .

[6]  Stephen P. Hubbell,et al.  Soil nutrients influence spatial distributions of tropical tree species , 2007, Proceedings of the National Academy of Sciences.

[7]  J. Svenning,et al.  The relative roles of environment and history as controls of tree species composition and richness in Europe , 2005 .

[8]  P. Legendre,et al.  vegan : Community Ecology Package. R package version 1.8-5 , 2007 .

[9]  Stephen P. Hubbell,et al.  Role of dispersal in the recruitment limitation of neotropical pioneer species , 2002 .

[10]  Karl Cottenie,et al.  Integrating environmental and spatial processes in ecological community dynamics. , 2005, Ecology letters.

[11]  P. Legendre,et al.  Variation partitioning of species data matrices: estimation and comparison of fractions. , 2006, Ecology.

[12]  Hanna Tuomisto,et al.  DISSECTING THE SPATIAL STRUCTURE OF ECOLOGICAL DATA AT MULTIPLE SCALES , 2004 .

[13]  Oliver L. Phillips,et al.  Habitat association among Amazonian tree species: a landscape‐scale approach , 2003 .

[14]  Kalle Ruokolainen,et al.  LINKING FLORISTIC PATTERNS WITH SOIL HETEROGENEITY AND SATELLITE IMAGERY IN ECUADORIAN AMAZONIA , 2003 .

[15]  H. Tuomisto Edaphic niche differentiation among Polybotrya ferns in western Amazonia: implications for coexistence and speciation , 2006 .

[16]  J. Plotkin,et al.  HABITAT PATTERNS IN TROPICAL RAIN FORESTS: A COMPARISON OF 105 PLOTS IN NORTHWEST BORNEO , 2002 .

[17]  Pierre Legendre,et al.  All-scale spatial analysis of ecological data by means of principal coordinates of neighbour matrices , 2002 .

[18]  P. Legendre,et al.  Spatial distribution and habitats of useful plants: an initial assessment for conservation on an indigenous territory, Panama , 2002, Biodiversity & Conservation.

[19]  Guillem Chust,et al.  Determinants and spatial modeling of tree β-diversity in a tropical forest landscape in Panama , 2006 .

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

[21]  H. Tuomisto,et al.  Edaphic and Floristic Variation within a 1‐ha Plot of Lowland Amazonian Rain Forest 1 , 2006 .

[22]  R. Dirzo,et al.  The effects of gap size and age on the understorey herb community of a tropical Mexican rain forest , 1992 .

[23]  T. Gregoire,et al.  HABITAT CHARACTERIZATIONS UNDERESTIMATE THE ROLE OF EDAPHIC FACTORS CONTROLLING THE DISTRIBUTION OF ENTANDROPHRAGMA , 2004 .

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

[25]  J. Denslow TROPICAL RAINFOREST GAPS AND TREE SPECIES DIVERSITY , 1987 .

[26]  Hanna Tuomisto,et al.  Dissecting Amazonian Biodiversity , 1995, Science.

[27]  S. Hubbell,et al.  The unified neutral theory of biodiversity and biogeography at age ten. , 2011, Trends in ecology & evolution.

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

[29]  M. Austin Spatial prediction of species distribution: an interface between ecological theory and statistical modelling , 2002 .

[30]  Á. Duque,et al.  Different floristic patterns of woody understorey and canopy plants in Colombian Amazonia , 2002, Journal of Tropical Ecology.

[31]  P. Legendre,et al.  Forward selection of explanatory variables. , 2008, Ecology.

[32]  C.J.F. ter Braak,et al.  CANOCO Reference Manual and User's Guide to Canoco for Windows: Software for Canonical Community Ordination (Version 4) , 1998 .

[33]  M. Silman,et al.  Distance-dependence in two Amazonian palms: effects of spatial and temporal variation in seed predator communities , 2004, Oecologia.

[34]  P. Balvanera,et al.  Patterns of β-diversity in a Mexican tropical dry forest , 2002 .

[35]  Jens-Christian Svenning,et al.  ECOLOGICAL DETERMINISM IN PLANT COMMUNITY STRUCTURE ACROSS A TROPICAL FOREST LANDSCAPE , 2004 .

[36]  Ter Braak,et al.  Canoco reference manual and CanoDraw for Windows user''s guide: software for canonical community ord , 2002 .

[37]  J. Duivenvoorden,et al.  Patterns of plant species composition on Amazonian sandstone outcrops in Colombia , 2004 .

[38]  Benjamin Gilbert,et al.  Neutrality, niches, and dispersal in a temperate forest understory. , 2004, Proceedings of the National Academy of Sciences of the United States of America.

[39]  Stéphane Dray,et al.  Spatial modelling: a comprehensive framework for principal coordinate analysis of neighbour matrices (PCNM) , 2006 .

[40]  J. Nabe‐Nielsen,et al.  An improved method for the rapid assessment of forest understorey light environments , 2000 .

[41]  Jens-Christian Svenning,et al.  Diversity and dominance in palm (Arecaceae) communities in terra firme forests in the western Amazon basin , 2004 .

[42]  C. Cannon,et al.  Tree species distributions across five habitats in a Bornean rain forest , 2004 .

[43]  P. Legendre,et al.  Partialling out the spatial component of ecological variation , 1992 .

[44]  D. Clark,et al.  Edaphic and Human Effects on Landscape‐Scale Distributions of Tropical Rain Forest Palms , 1995 .

[45]  Hanna Tuomisto,et al.  Effects of mesoscale environmental heterogeneity and dispersal limitation on floristic variation in rain forest ferns , 2006 .

[46]  P. Legendre,et al.  ANALYZING BETA DIVERSITY: PARTITIONING THE SPATIAL VARIATION OF COMMUNITY COMPOSITION DATA , 2005 .

[47]  R. Marrs,et al.  Relationships between the species composition of forest field‐layer vegetation and environmental drivers, assessed using a national scale survey , 2006 .

[48]  Ralph Dubayah,et al.  Validation of Vegetation Canopy Lidar sub-canopy topography measurements for a dense tropical forest , 2002 .

[49]  H. Balslev,et al.  Overstorey Control of Understorey Species Composition in a Near-natural Temperate Broadleaved Forest in Denmark , 2005, Plant Ecology.

[50]  Gerrit Davidse,et al.  Psilotaceae a salviniaceae , 1995 .

[51]  Joost E. Duivenvoorden,et al.  Tree species composition and rain forest-environment relationships in the middle Caquetá area, Colombia, NW Amazonia , 1995, Vegetatio.