Beyond the continuum: a multi-dimensional phase space for neutral–niche community assembly

Neutral and niche processes are generally considered to interact in natural communities along a continuum, exhibiting community patterns bounded by pure neutral and pure niche processes. The continuum concept uses niche separation, an attribute of the community, to test the hypothesis that communities are bounded by pure niche or pure neutral conditions. It does not accommodate interactions via feedback between processes and the environment. By contrast, we introduce the Community Assembly Phase Space (CAPS), a multi-dimensional space that uses community processes (such as dispersal and niche selection) to define the limiting neutral and niche conditions and to test the continuum hypothesis. We compare the outputs of modelled communities in a heterogeneous landscape, assembled by pure neutral, pure niche and composite processes. Differences in patterns under different combinations of processes in CAPS reveal hidden complexity in neutral–niche community dynamics. The neutral–niche continuum only holds for strong dispersal limitation and niche separation. For weaker dispersal limitation and niche separation, neutral and niche processes amplify each other via feedback with the environment. This generates patterns that lie well beyond those predicted by a continuum. Inferences drawn from patterns about community assembly processes can therefore be misguided when based on the continuum perspective. CAPS also demonstrates the complementary information value of different patterns for inferring community processes and captures the complexity of community assembly. It provides a general tool for studying the processes structuring communities and can be applied to address a range of questions in community and metacommunity ecology.

[1]  R Core Team,et al.  R: A language and environment for statistical computing. , 2014 .

[2]  S. Hubbell,et al.  The case for ecological neutral theory. , 2012, Trends in ecology & evolution.

[3]  Helmut Hillebrand,et al.  Empirical approaches to metacommunities: a review and comparison with theory. , 2011, Trends in ecology & evolution.

[4]  S. Hubbell The Unified Neutral Theory of Biodiversity and Biogeography (MPB-32) , 2011 .

[5]  T. Hughes,et al.  Coral reef diversity refutes the neutral theory of biodiversity , 2006, Nature.

[6]  Wilfried Thuiller,et al.  From diversity indices to community assembly processes: a test with simulated data , 2012 .

[7]  Y. Carmel,et al.  Environmental heterogeneity affects the location of modelled communities along the niche–neutrality continuum , 2014, Proceedings of the Royal Society B: Biological Sciences.

[8]  D. Wardle,et al.  Spatial soil ecology , 2002 .

[9]  B. Mcrae,et al.  ISOLATION BY RESISTANCE , 2006, Evolution; international journal of organic evolution.

[10]  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.

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

[12]  A. Peterson,et al.  INTERPRETATION OF MODELS OF FUNDAMENTAL ECOLOGICAL NICHES AND SPECIES' DISTRIBUTIONAL AREAS , 2005 .

[13]  L. Jost Partitioning diversity into independent alpha and beta components. , 2007, Ecology.

[14]  Lael Parrott,et al.  Levels of emergence in individual based models: Coping with scarcity of data and pattern redundancy , 2011 .

[15]  C. M. Mutshinda,et al.  Integrating the niche and neutral perspectives on community structure and dynamics , 2011, Oecologia.

[16]  C. Stokes,et al.  Niche differentiation and neutral theory: an integrated perspective on shrub assemblages in a parkland savanna. , 2010, Ecology.

[17]  S. Levin,et al.  Comparing Classical Community Models: Theoretical Consequences for Patterns of Diversity , 2002, The American Naturalist.

[18]  J. D. Haan,et al.  How emergence arises , 2006 .

[19]  Michel Loreau,et al.  A mathematical synthesis of niche and neutral theories in community ecology. , 2011, Journal of theoretical biology.

[20]  John L. Harper,et al.  Population Biology of Plants. , 1978 .

[21]  Jonathan M. Chase,et al.  Disentangling the importance of ecological niches from stochastic processes across scales , 2011, Philosophical Transactions of the Royal Society B: Biological Sciences.

[22]  Paul A. Keddy,et al.  Assembly and response rules: two goals for predictive community ecology , 1992 .

[23]  Dominique Gravel,et al.  Reconciling niche and neutrality: the continuum hypothesis. , 2006, Ecology letters.

[24]  Lindsay A. Turnbull,et al.  Biology, chance, or history? The predictable reassembly of temperate grassland communities. , 2010, Ecology.

[25]  Mark A. Davis,et al.  Fluctuating resources in plant communities: a general theory of invasibility , 2000 .

[26]  Neil J. Anderson,et al.  Wolverine gene flow across a narrow climatic niche. , 2009, Ecology.

[27]  Ryan A Chisholm,et al.  Niche and neutral models predict asymptotically equivalent species abundance distributions in high-diversity ecological communities , 2010, Proceedings of the National Academy of Sciences.

[28]  M. Cadotte Concurrent niche and neutral processes in the competition–colonization model of species coexistence , 2007, Proceedings of the Royal Society B: Biological Sciences.

[29]  Shurong Zhou,et al.  Hybrid niche-neutral models outperform an otherwise equivalent neutral model for fitting coral reef data. , 2013, Journal of theoretical biology.

[30]  E. Tjørve,et al.  Scale‐dependence in species‐area relationships , 2005 .

[31]  R. Whittaker,et al.  Neutral theory and the species abundance distribution: recent developments and prospects for unifying niche and neutral perspectives , 2014, Ecology and evolution.

[32]  S. Hubbell,et al.  The Unified Neutral Theory of Biodiversity and Biogeography , 2001 .

[33]  J. Harwood,et al.  Web‐location by linyphiid spiders: prey‐specific aggregation and foraging strategies , 2003 .

[34]  M. Uriarte,et al.  Multispecies coexistence of trees in tropical forests: spatial signals of topographic niche differentiation increase with environmental heterogeneity , 2013, Proceedings of the Royal Society B: Biological Sciences.

[35]  J. Dengler,et al.  Which function describes the species–area relationship best? A review and empirical evaluation , 2009 .

[36]  T. Hughes,et al.  Coral reef diversity refutes the neutral theory of biodiversity , 2006, Nature.

[37]  James S Clark,et al.  The coherence problem with the Unified Neutral Theory of Biodiversity. , 2012, Trends in ecology & evolution.

[38]  Pierre Legendre,et al.  SPECIES DIVERSITY PATTERNS DERIVED FROM SPECIES–AREA MODELS , 2002 .