Community assembly: alternative stable states or alternative transient states?

The concept of alternative stable states has long been a dominant framework for studying the influence of historical contingency in community assembly. This concept focuses on stable states, yet many real communities are kept in a transient state by disturbance, and the utility of predictions for stable states in explaining transient states remains unclear. Using a simple model of plant community assembly, we show that the conditions under which historical contingency affects community assembly can differ greatly for stable versus transient states. Differences arise because the contribution of such factors as mortality rate, environmental heterogeneity and plant-soil feedback to historical contingency changes as community assembly proceeds. We also show that transient states can last for a long time relative to immigration rate and generation time. These results argue for a conceptual shift of focus from alternative stable states to alternative transient states for understanding historical contingency in community assembly.

[1]  P. White,et al.  The Ecology of Natural Disturbance and Patch Dynamics , 1986 .

[2]  Anthony R. Taylor,et al.  Multiple successional pathways of boreal forest stands in central Canada. , 2011 .

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

[4]  Sharon K Collinge,et al.  Transient patterns in the assembly of vernal pool plant communities. , 2009, Ecology.

[5]  Peter J. Bellingham,et al.  Contrasting impacts of a native and an invasive exotic shrub on flood-plain succession , 2005 .

[6]  Jonathan M. Chase,et al.  Alternative stable states and regional community structure. , 2004, Journal of theoretical biology.

[7]  Richard J. Hobbs,et al.  Models for Ecosystem Dynamics as Frameworks for Restoration Ecology , 2009 .

[8]  G. Bell.  Neutral macroecology. , 2001, Science.

[9]  M. Scheffer,et al.  IMPLICATIONS OF SPATIAL HETEROGENEITY FOR CATASTROPHIC REGIME SHIFTS IN ECOSYSTEMS , 2005 .

[10]  A. D. Roos,et al.  Direct experimental evidence for alternative stable states: a review , 2005 .

[11]  A. Hastings.  Timescales, dynamics, and ecological understanding1 , 2010 .

[12]  George C. Hurtt,et al.  The consequences of recruitment limitation: reconciling chance, history and competitive differences between plants , 1995 .

[13]  P. White,et al.  The Ecology of Natural Disturbance and Patch Dynamics , 1986 .

[14]  R. Whittaker.  Evolution and measurement of species diversity , 1972 .

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

[16]  David L. Verbyla,et al.  Relative importance of different secondary successional pathways in an Alaskan boreal forest , 2008 .

[17]  G. van der Velde,et al.  Ecological niches. Linking classical and contemporary approaches , 2008 .

[18]  R. Lewontin,et al.  The meaning of stability. , 2020, Brookhaven symposia in biology.

[19]  T. Fukami,et al.  Assembly history dictates ecosystem functioning: evidence from wood decomposer communities. , 2010, Ecology letters.

[20]  Kevin J. Gaston,et al.  Measuring beta diversity for presence–absence data , 2003 .

[21]  S. Carpenter,et al.  Early-warning signals for critical transitions , 2009, Nature.

[22]  C. Peterson.  Does a Rigorous Criterion for Environmental Identity Preclude the Existence of Multiple Stable Points? , 1984, The American Naturalist.

[23]  T. Mitchell-Olds,et al.  REGRESSION ANALYSIS OF NATURAL SELECTION: STATISTICAL INFERENCE AND BIOLOGICAL INTERPRETATION , 1987, Evolution; international journal of organic evolution.

[24]  Mathew A. Leibold,et al.  CYCLIC ASSEMBLY TRAJECTORIES AND SCALE‐DEPENDENT PRODUCTIVITY–DIVERSITY RELATIONSHIPS , 2004 .

[25]  S. Levin.  Population Dynamic Models in Heterogeneous Environments , 1976 .

[26]  D. DeAngelis,et al.  Equilibrium and Nonequilibrium Concepts in Ecological Models , 1987 .

[27]  T. Fukami.  ASSEMBLY HISTORY INTERACTS WITH ECOSYSTEM SIZE TO INFLUENCE SPECIES DIVERSITY , 2004 .

[28]  Paul Kardol,et al.  Temporal variation in plant-soil feedback controls succession. , 2006, Ecology letters.

[29]  A. Hastings.  Transients: the key to long-term ecological understanding? , 2004, Trends in ecology & evolution.

[30]  T. Fukami,et al.  Immigration history controls diversification in experimental adaptive radiation , 2007, Nature.

[31]  R. Law.  Theoretical Aspects of Community Assembly , 2009 .

[32]  Nicolas Mouquet,et al.  Mechanisms of Coexistence in Competitive Metacommunities , 2004, The American Naturalist.

[33]  David Tilman,et al.  Limiting Similarity in Mechanistic and Spatial Models of Plant Competition in Heterogeneous Environments , 1994, The American Naturalist.

[34]  I. Noy-Meir,et al.  Stability of Grazing Systems: An Application of Predator-Prey Graphs , 1975 .

[35]  Tadashi Fukami,et al.  Community assembly along a species pool gradient: implications for multiple-scale patterns of species diversity , 2004, Population Ecology.

[36]  N. Knowlton.  Thresholds and Multiple Stable States in Coral Reef Community Dynamics , 1992 .

[37]  James A. Drake,et al.  Community-Assembly Mechanics and the Structure of an Experimental Species Ensemble , 1991, The American Naturalist.

[38]  David A. Norton,et al.  Are systems with strong underlying abiotic regimes more likely to exhibit alternative stable states , 2005 .

[39]  T. O. Crist,et al.  Partitioning species diversity across landscapes and regions: a hierarchical analysis of alpha, beta, and gamma diversity. , 2003, The American naturalist.

[40]  P. Petraitis,et al.  Experimental evidence for the origin of alternative communities on rocky intertidal shores , 1999 .

[41]  Elizaveta Pachepsky,et al.  Plant-soil feedbacks and invasive spread. , 2006, Ecology letters.

[42]  Jonathan M. Chase.  Productive Environments Stochastic Community Assembly Causes Higher Biodiversity in More , 2012 .

[43]  M. Scheffer,et al.  Long Transients Near the Ghost of a Stable State in Eutrophic Shallow Lakes with Fluctuating Water Levels , 2007, Ecosystems.

[44]  James A. Drake,et al.  The mechanics of community assembly and succession , 1990 .

[45]  Marten Scheffer,et al.  Slow Recovery from Perturbations as a Generic Indicator of a Nearby Catastrophic Shift , 2007, The American Naturalist.

[46]  Jeffrey Clary,et al.  The ecology of restoration: historical links, emerging issues and unexplored realms , 2005 .

[47]  R. May.  Thresholds and breakpoints in ecosystems with a multiplicity of stable states , 1977, Nature.

[48]  H. Tuomisto.  A diversity of beta diversities: straightening up a concept gone awry. Part 1. Defining beta diversity as a function of alpha and gamma diversity , 2010 .

[49]  Margaret J Eppstein,et al.  Invasiveness in plant communities with feedbacks. , 2007, Ecology letters.

[50]  J. Bever,et al.  Soil community feedback and the coexistence of competitors: conceptual frameworks and empirical tests. , 2003, The New phytologist.

[51]  T. M. Bezemer,et al.  Species divergence and trait convergence in experimental plant community assembly , 2005 .

[52]  M. Loreau,et al.  Plant species richness and community productivity: why the mechanism that promotes coexistence matters , 2002 .

[53]  K. Anderson.  Temporal Patterns in Rates of Community Change during Succession , 2007, The American Naturalist.

[54]  M. Westoby,et al.  Opportunistic management for rangelands not at equilibrium. , 1989 .

[55]  J. Connell,et al.  On the Evidence Needed to Judge Ecological Stability or Persistence , 1983, The American Naturalist.

[56]  Roger del Moral,et al.  Transition Dynamics in Succession : Implications for Rates , Trajectories and Restoration , 2008 .

[57]  L. Jost,et al.  Independence of alpha and beta diversities. , 2010, Ecology.

[58]  T. Schoener,et al.  Alternatives to Lotka-Volterra competition: models of intermediate complexity. , 1976, Theoretical population biology.

[59]  D. Wardle,et al.  How understanding aboveground-belowground linkages can assist restoration ecology. , 2010, Trends in ecology & evolution.

[60]  Wim H. van der Putten,et al.  Microbe-mediated plant-soil feedback causes historical contingency effects in plant community assembly , 2007 .

[61]  Peter Chesson,et al.  Coexistence of Competitors in Spatially and Temporally Varying Environments: A Look at the Combined Effects of Different Sorts of Variability , 1985 .

[62]  J. L. Reid,et al.  New Models for Ecosystem Dynamics and Restoration , 2009 .

[63]  Nancy Knowlton,et al.  Multiple “stable” states and the conservation of marine ecosystems , 2004 .

[64]  M. Gilpin,et al.  Multiple domains of attraction in competition communities , 1976, Nature.

[65]  James C. Stegen,et al.  PERSPECTIVE Navigating the multiple meanings of b diversity: a roadmap for the practicing ecologist , 2010 .

[66]  Brian J Wilsey.  An empirical comparison of beta diversity indices in establishing prairies. , 2010, Ecology.

[67]  D. Haydon,et al.  Alternative stable states in ecology , 2003 .

[68]  Jonathan M. Chase.  Community assembly: when should history matter? , 2003, Oecologia.

[69]  Lin Jiang,et al.  Community assembly in the presence of disturbance: a microcosm experiment. , 2008, Ecology.

[70]  Henry Chandler Cowles,et al.  The Ecological Relations of the Vegetation on the Sand Dunes of Lake Michigan [Continued] , 1899, Botanical Gazette.

[71]  R. Jackson,et al.  A state-transition approach to understanding nonequilibrium plant community dynamics in Californian grasslands , 2002, Plant Ecology.

[72]  J. Sutherland,et al.  Multiple Stable Points in Natural Communities , 1974, The American Naturalist.

[73]  T. O. Crist,et al.  Diversity partitioning without statistical independence of alpha and beta. , 2010, Ecology.

[74]  S. Carpenter,et al.  Catastrophic shifts in ecosystems , 2001, Nature.

[75]  C. Fastie,et al.  Causes and Ecosystem Consequences of Multiple Pathways of Primary Succession at Glacier Bay, Alaska , 1995 .

[76]  R. Lande.  Statistics and partitioning of species diversity, and similarity among multiple communities , 1996 .

[77]  Henry C. Cowles,et al.  The Ecological Relations of the Vegetation on the Sand Dunes of Lake Michigan [Continued] , 1899, Botanical Gazette.

[78]  G. Almany,et al.  PRIORITY EFFECTS IN CORAL REEF FISH COMMUNITIES , 2003 .

[79]  R. Whittaker.  Vegetation of the Siskiyou Mountains, Oregon and California , 1960 .

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

[81]  Lewontin Rc,et al.  The Meaning of Stability , 2020, The Early Mubarak Years 1982–1988.