Spatially Explicit Metrics of Species Diversity, Functional Diversity, and Phylogenetic Diversity: Insights into Plant Community Assembly Processes

Spatial processes underlie major species coexistence mechanisms. A range of spatial analysis techniques are increasingly applied to data of fully mapped communities to quantify spatial structures in species and phylogenetic and functional diversity at some given spatial scale with the goal of gaining insights into processes of community assembly and dynamics. We review these techniques, including spatial point pattern analysis, quadrat-based analyses, and individual-based neighborhood models, and provide a practical roadmap for ecologists in the analysis of local spatial structures in species and phylogenetic and functional diversity. We show how scale-dependent metrics of spatial diversity can be used in concert with ecological null models, statistical models, and dynamic community simulation models to detect spatial patterns, reveal the influence of the biotic neighborhood on plant performance, and quantify the relative contribution of species interactions, habitat heterogeneity, and stochastic processe...

[1]  David L. Erickson,et al.  Trait similarity, shared ancestry and the structure of neighbourhood interactions in a subtropical wet forest: implications for community assembly. , 2010, Ecology letters.

[2]  G. Perry,et al.  Stochastic geometry best explains spatial associations among species pairs and plant functional types in species‐rich shrublands , 2014 .

[3]  Stephen P Hubbell,et al.  Asymmetric Density Dependence Shapes Species Abundances in a Tropical Tree Community , 2010, Science.

[4]  M. Fortin,et al.  Spatial pattern and ecological analysis , 1989, Vegetatio.

[5]  Virgilio Gómez-Rubio,et al.  Spatial Point Patterns: Methodology and Applications with R , 2016 .

[6]  S. Levin THE PROBLEM OF PATTERN AND SCALE IN ECOLOGY , 1992 .

[7]  David Kenfack,et al.  A general framework for the distance–decay of similarity in ecological communities , 2008, Ecology letters.

[8]  B. McGill Towards a unification of unified theories of biodiversity. , 2010, Ecology letters.

[9]  Widespread density-dependent seedling mortality promotes species coexistence in a highly diverse Amazonian rain forest. , 2010, Ecology.

[10]  Andreas Huth,et al.  Statistical inference for stochastic simulation models--theory and application. , 2011, Ecology letters.

[11]  Matthew R. Helmus,et al.  Separating the determinants of phylogenetic community structure. , 2007, Ecology letters.

[12]  S. Reed,et al.  Tree species control rates of free-living nitrogen fixation in a tropical rain forest. , 2008, Ecology.

[13]  C. Canham,et al.  A neighborhood analysis of canopy tree competition : effects of shading versus crowding , 2004 .

[14]  Andreas Huth,et al.  Moving beyond abundance distributions: neutral theory and spatial patterns in a tropical forest , 2015, Proceedings of the Royal Society B: Biological Sciences.

[15]  M. Uriarte,et al.  Effects of forest fragmentation on the seedling recruitment of a tropical herb: assessing seed vs. safe-site limitation. , 2010, Ecology.

[16]  David W. Winkler,et al.  20. A Null Model for Null Models in Biogeography , 1984 .

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

[18]  Stephen P. Hubbell,et al.  Local neighborhood effects on long-term survival of individual trees in a neotropical forest , 2001, Ecological Research.

[19]  Nathan J B Kraft,et al.  Plant functional traits and the multidimensional nature of species coexistence , 2015, Proceedings of the National Academy of Sciences.

[20]  Werner Ulrich,et al.  Intransitive competition is widespread in plant communities and maintains their species richness. , 2015, Ecology letters.

[21]  Kenichiro Shimatani Multivariate point processes and spatial variation of species diversity , 2001 .

[22]  Stephen P. Hubbell,et al.  Beta-Diversity in Tropical Forest Trees , 2002, Science.

[23]  Charles D. Canham,et al.  Natural disturbance and human land use as determinants of tropical forest dynamics: results from a forest simulator , 2009 .

[24]  D. M. Newbery,et al.  Evidence of species-specific neighborhood effects in the dipterocarpaceae of a Bornean rain forest , 2005 .

[25]  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 .

[26]  Thorsten Wiegand,et al.  An evaluation of the state of spatial point pattern analysis in ecology , 2016 .

[27]  S. Hubbell,et al.  A systematic comparison of summary characteristics for quantifying point patterns in ecology , 2013 .

[28]  J. Illian,et al.  Success of spatial statistics in determining underlying process in simulated plant communities , 2016 .

[29]  J. Zimmerman,et al.  Life‐history trade‐offs during the seed‐to‐seedling transition in a subtropical wet forest community , 2013 .

[30]  François Goreaud,et al.  ads Package for R: A Fast Unbiased Implementation of the K-function Family for Studying Spatial Point Patterns in Irregular-Shaped Sampling Windows , 2015 .

[31]  J. Connell Diversity and the coevolution of competitors, or the ghost of competition past , 1980 .

[32]  J. Chave,et al.  Phylogenetic density dependence and environmental filtering predict seedling mortality in a tropical forest. , 2012, Ecology letters.

[33]  Robin L. Chazdon,et al.  A trait‐mediated, neighbourhood approach to quantify climate impacts on successional dynamics of tropical rainforests , 2016 .

[34]  Simon A. Levin,et al.  Biologically generated spatial pattern and the coexistence of competing species , 1997 .

[35]  K. R. Clarke,et al.  A taxonomic distinctness index and its statistical properties , 1998 .

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

[37]  G. Perry,et al.  Community‐level spatial structure supports a model of stochastic geometry in species‐rich shrublands , 2017 .

[38]  Fei Lin,et al.  Mechanisms underlying local functional and phylogenetic beta diversity in two temperate forests. , 2015, Ecology.

[39]  D. Lieberman,et al.  Nearest‐neighbor tree species combinations in tropical forest: the role of chance, and some consequences of high diversity , 2007 .

[40]  M. Mayfield,et al.  Opposing effects of competitive exclusion on the phylogenetic structure of communities. , 2010, Ecology letters.

[41]  L. Webb,et al.  Compensatory Recruitment, Growth, and Mortality as Factors Maintaining Rain Forest Tree Diversity , 1984 .

[42]  J. Chave,et al.  Prevalence of phylogenetic clustering at multiple scales in an African rain forest tree community , 2014 .

[43]  P. Brussard,et al.  Matters of scale. , 1993, Science.

[44]  Nathan J B Kraft,et al.  Functional trait differences influence neighbourhood interactions in a hyperdiverse Amazonian forest. , 2016, Ecology letters.

[45]  A. Escudero,et al.  Beyond the classical nurse species effect: diversity assembly in a Mediterranean semi‐arid dwarf shrubland , 2016 .

[46]  Andreas Huth,et al.  Neighborhood diversity of large trees shows independent species patterns in a mixed dipterocarp forest in Sri Lanka. , 2015, Ecology.

[47]  Functional diversity through the mean trait dissimilarity: resolving shortcomings with existing paradigms and algorithms , 2016, Oecologia.

[48]  Roger Bivand,et al.  Community ecology in the age of multivariate multiscale spatial analysis , 2012 .

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

[50]  Peter Stoll,et al.  INTRASPECIFIC AGGREGATION ALTERS COMPETITIVE INTERACTIONS IN EXPERIMENTAL PLANT COMMUNITIES , 2001 .

[51]  J. Plotkin,et al.  Species-area curves, spatial aggregation, and habitat specialization in tropical forests. , 2000, Journal of theoretical biology.

[52]  Claudia Neuhauser,et al.  Spatial Dynamics in Model Plant Communities: What Do We Really Know? , 2003, The American Naturalist.

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

[54]  Xiangcheng Mi,et al.  Quantifying spatial phylogenetic structures of fully stem‐mapped plant communities , 2013 .

[55]  Susanne A. Fritz,et al.  A guide to phylogenetic metrics for conservation, community ecology and macroecology , 2016, Biological reviews of the Cambridge Philosophical Society.

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

[57]  D. Farine,et al.  Phylogenetic community structure metrics and null models: a review with new methods and software , 2015, bioRxiv.

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

[59]  Stephan Getzin,et al.  Testing the independent species’ arrangement assertion made by theories of stochastic geometry of biodiversity , 2012, Proceedings of the Royal Society B: Biological Sciences.

[60]  F. Valladares,et al.  Fine-scale spatial patterns and genetic structure of two co-occurring plant congeners: fingerprints of coexistence? , 2011 .

[61]  N. Swenson,et al.  A Phylogenetic Perspective on the Individual Species-Area Relationship in Temperate and Tropical Tree Communities , 2013, PloS one.

[62]  M. Uriarte,et al.  Analysis of neighborhood dynamics of forest ecosystems using likelihood methods and modeling. , 2006, Ecological applications : a publication of the Ecological Society of America.

[63]  Fei Lin,et al.  Phylogenetic and functional diversity area relationships in two temperate forests , 2013 .

[64]  Jonathan M. Chase,et al.  Navigating the multiple meanings of β diversity: a roadmap for the practicing ecologist. , 2011, Ecology letters.

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

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

[67]  J. Wiens Spatial Scaling in Ecology , 1989 .

[68]  Uta Berger,et al.  Pattern-Oriented Modeling of Agent-Based Complex Systems: Lessons from Ecology , 2005, Science.

[69]  N. Gotelli Null model analysis of species co-occurrence patterns , 2000 .

[70]  S. Hubbell,et al.  Strong density- and diversity-related effects help to maintain tree species diversity in a neotropical forest. , 1997, Proceedings of the National Academy of Sciences of the United States of America.

[71]  Andreas Huth,et al.  How individual species structure diversity in tropical forests , 2007, Proceedings of the National Academy of Sciences.

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

[73]  S. Hubbell,et al.  A spatially explicit model of sapling growth in a tropical forest: does the identity of neighbours matter? , 2004 .

[74]  Stefano Allesina,et al.  A competitive network theory of species diversity , 2011, Proceedings of the National Academy of Sciences.

[75]  T. Wiegand,et al.  Spatial patterns of tree species richness in two temperate forests , 2011 .

[76]  Andreas Huth,et al.  Fast calibration of a dynamic vegetation model with minimum observation data , 2015 .

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

[78]  W. Bossert,et al.  The Measurement of Diversity , 2001 .

[79]  S. Levin The problem of pattern and scale in ecology , 1992 .

[80]  P. R. van Gardingen,et al.  An individual-based spatially explicit tree growth model for forests in East Kalimantan (Indonesian Borneo) , 2003 .

[81]  J. Zimmerman,et al.  A NEIGHBORHOOD ANALYSIS OF TREE GROWTH AND SURVIVAL IN A HURRICANE‐DRIVEN TROPICAL FOREST , 2004 .

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

[83]  Norman A. Bourg,et al.  CTFS‐ForestGEO: a worldwide network monitoring forests in an era of global change , 2015, Global change biology.

[84]  Andreas Huth,et al.  Do abundance distributions and species aggregation correctly predict macroecological biodiversity patterns in tropical forests? , 2016, Global ecology and biogeography : a journal of macroecology.

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

[86]  Nathan J B Kraft,et al.  Stochastic dilution effects weaken deterministic effects of niche-based processes in species rich forests. , 2016, Ecology.

[87]  Nathan J B Kraft,et al.  The Assembly of Plant Communities , 2014 .

[88]  Eliot J. B. McIntire,et al.  Beyond description: the active and effective way to infer processes from spatial patterns. , 2009, Ecology.

[89]  Jonathan M. Chase,et al.  The metacommunity concept: a framework for multi-scale community ecology , 2004 .

[90]  O. Hardy,et al.  Characterizing the phylogenetic structure of communities by an additive partitioning of phylogenetic diversity , 2007 .

[91]  P. Legendre Spatial Autocorrelation: Trouble or New Paradigm? , 1993 .

[92]  Peter Kareiva,et al.  Spatial ecology : the role of space in population dynamics and interspecific interactions , 1998 .

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

[94]  Hans-Hermann Gerdes,et al.  Functional Traits and Niche-Based Tree Community Assembly in an Amazonian Forest , 2008 .

[95]  M. Uriarte,et al.  Trait-mediated assembly processes predict successional changes in community diversity of tropical forests , 2014, Proceedings of the National Academy of Sciences.

[96]  Richard Condit,et al.  Tropical Forest Census Plots , 1998, Environmental Intelligence Unit.

[97]  Nathan J B Kraft,et al.  Community assembly, coexistence and the environmental filtering metaphor , 2015 .

[98]  Calum Brown,et al.  Linking ecological processes with spatial and non‐spatial patterns in plant communities , 2011 .

[99]  Xiangcheng Mi,et al.  Species-area relationships explained by the joint effects of dispersal limitation and habitat heterogeneity. , 2009, Ecology.

[100]  Daniel Simberloff,et al.  The Assembly of Species Communities: Chance or Competition? , 1979 .

[101]  J. Zimmerman,et al.  Linking spatial patterns of leaf litterfall and soil nutrients in a tropical forest: a neighborhood approach. , 2015, Ecological applications : a publication of the Ecological Society of America.

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

[103]  P. Diggle,et al.  Spatial patterns reveal negative density dependence and habitat associations in tropical trees. , 2011, Ecology.

[104]  F. Valladares,et al.  Competition may explain the fine‐scale spatial patterns and genetic structure of two co‐occurring plant congeners , 2011 .

[105]  Calyampudi R. Rao Diversity and dissimilarity coefficients: A unified approach☆ , 1982 .

[106]  D. Coomes,et al.  Competitive interactions between forest trees are driven by species' trait hierarchy, not phylogenetic or functional similarity: implications for forest community assembly. , 2012, Ecology letters.

[107]  N. Swenson Phylogenetic Beta Diversity Metrics, Trait Evolution and Inferring the Functional Beta Diversity of Communities , 2011, PloS one.