Constructing Random Matrices to Represent Real Ecosystems

Models of complex systems with n components typically have order n2 parameters because each component can potentially interact with every other. When it is impractical to measure these parameters, one may choose random parameter values and study the emergent statistical properties at the system level. Many influential results in theoretical ecology have been derived from two key assumptions: that species interact with random partners at random intensities and that intraspecific competition is comparable between species. Under these assumptions, community dynamics can be described by a community matrix that is often amenable to mathematical analysis. We combine empirical data with mathematical theory to show that both of these assumptions lead to results that must be interpreted with caution. We examine 21 empirically derived community matrices constructed using three established, independent methods. The empirically derived systems are more stable by orders of magnitude than results from random matrices. This consistent disparity is not explained by existing results on predator-prey interactions. We investigate the key properties of empirical community matrices that distinguish them from random matrices. We show that network topology is less important than the relationship between a species’ trophic position within the food web and its interaction strengths. We identify key features of empirical networks that must be preserved if random matrix models are to capture the features of real ecosystems.

[1]  P. Erdos,et al.  On the evolution of random graphs , 1984 .

[2]  Josef Hofbauer,et al.  Evolutionary Games and Population Dynamics , 1998 .

[3]  Johan van de Koppel,et al.  Reconciling complexity with stability in naturally assembling food webs , 2009, Nature.

[4]  R. Macarthur Fluctuations of Animal Populations and a Measure of Community Stability , 1955 .

[5]  H. W. Hunt,et al.  Microcosms and soil Ecology: Critical Linkages between Fields Studies and Modelling Food Webs , 1996 .

[6]  Madan Lal Mehta,et al.  Random Matrices and the Statistical Theory of Energy Levels. , 1970 .

[7]  K. McCann The diversity–stability debate , 2000, Nature.

[8]  W. Jansen,et al.  A permanence theorem for replicator and Lotka-Volterra systems , 1987 .

[9]  S. Carpenter,et al.  Food Webs, Body Size, and Species Abundance in Ecological Community Description , 2005 .

[10]  U. Jacob,et al.  Perturbing a Marine Food Web: Consequences for Food Web Structure and Trivariate Patterns , 2012 .

[11]  Si Tang,et al.  Stability criteria for complex ecosystems , 2011, Nature.

[12]  Samraat Pawar,et al.  Dimensionality of consumer search space drives trophic interaction strengths , 2012, Nature.

[13]  Stefano Allesina,et al.  The ghost of nestedness in ecological networks , 2013, Nature Communications.

[14]  Colin Fontaine,et al.  Stability of Ecological Communities and the Architecture of Mutualistic and Trophic Networks , 2010, Science.

[15]  M. Emmerson,et al.  Predator–prey body size, interaction strength and the stability of a real food web , 2004 .

[16]  J. Lawton,et al.  On feeding on more than one trophic level , 1978, Nature.

[17]  Richard Levins,et al.  Coexistence in a Variable Environment , 1979, The American Naturalist.

[18]  H. William Hunt,et al.  Influence of Productivity on the Stability of Real and Model Ecosystems , 1993, Science.

[19]  Jerry C. Blackford,et al.  Self-assembling food webs : a global viewpoint of coexistence of species in Lotka-Volterra communities , 1992 .

[20]  Christian Wissel,et al.  Babel, or the ecological stability discussions: an inventory and analysis of terminology and a guide for avoiding confusion , 1997, Oecologia.

[21]  Villy Christensen,et al.  ECOPATH II − a software for balancing steady-state ecosystem models and calculating network characteristics , 1992 .

[22]  Mats Gyllenberg,et al.  Competitive exclusion and limiting similarity: a unified theory. , 2006, Theoretical population biology.

[23]  Johan van de Koppel,et al.  Reconciling complexity with stability in naturally assembling food webs , 2007, Nature.

[24]  Dominique Gravel,et al.  Persistence Increases with Diversity and Connectance in Trophic Metacommunities , 2011, PloS one.

[25]  Stefano Allesina,et al.  Correlation between interaction strengths drives stability in large ecological networks. , 2014, Ecology letters.

[26]  H. W. Hunt,et al.  Calculation of nitrogen mineralization in soil food webs , 1993, Plant and Soil.

[27]  Axel G. Rossberg,et al.  Food webs and biodiversity: Foundations, models, data , 2015 .

[28]  C. Elton The Ecology of Invasions by Animals and Plants , 1960, Springer US.

[29]  Jordi Bascompte,et al.  The architecture of mutualistic networks minimizes competition and increases biodiversity , 2009, Nature.

[30]  Anders Nielsen,et al.  Conservation of species interaction networks , 2010 .

[31]  ROBERT M. MAY,et al.  Will a Large Complex System be Stable? , 1972, Nature.

[32]  H. Luh,et al.  Qualitative Stability and Ambiguity in Model Ecosystems , 2003, The American Naturalist.

[33]  S. Wiggins Introduction to Applied Nonlinear Dynamical Systems and Chaos , 1989 .

[34]  S. Opitz,et al.  Trophic interactions in Caribbean coral reefs , 1996 .

[35]  Anje-Margriet Neutel,et al.  Stability in Real Food Webs: Weak Links in Long Loops , 2002, Science.

[36]  Guy Woodward,et al.  Body size in ecological networks. , 2005, Trends in ecology & evolution.

[37]  Charles C. Elton,et al.  The Ecology of Invasions by Animals and Plants. , 1959 .

[38]  M. Emmerson,et al.  Weak interactions, omnivory and emergent food-web properties , 2004, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[39]  Eoin J. O’Gorman,et al.  Manipulating Interaction Strengths and the Consequences for Trivariate Patterns in a Marine Food Web , 2010 .

[40]  Madan Lal Mehta,et al.  Random Matrices and the Statistical Theory of Energy Levels , 2014 .

[41]  N. Gribble Ecosystem Modelling Of The Great Barrier Reef : A Balanced Trophic Biomass Approach , 2005 .

[42]  B. Bollobás The evolution of random graphs , 1984 .

[43]  Ulrich Brose,et al.  Allometric degree distributions facilitate food-web stability , 2007, Nature.

[44]  A. Hastings,et al.  Weak trophic interactions and the balance of nature , 1998, Nature.

[45]  G. Woodward,et al.  Chapter 1 Allometry of Body Size and Abundance in 166 Food Webs , 2009 .

[46]  C. Möllmann,et al.  Marine Ecosystem Regime Shifts Induced by Climate and Overfishing: A Review for the Northern Hemisphere , 2012 .

[47]  Neo D. Martinez,et al.  Allometric scaling enhances stability in complex food webs. , 2006, Ecology letters.

[48]  Stefano Allesina,et al.  Network structure , predator-prey modules , and stability in large food webs : Electronic Supplementary Material ( ESM ) , 2007 .

[49]  E. Berlow,et al.  QUANTIFYING VARIATION IN THE STRENGTHS OF SPECIES INTERACTIONS , 1999 .

[50]  D. Gravel,et al.  No complexity-stability relationship in natural communities , 2013, 1307.5364.

[51]  V. Christensen,et al.  Food web models and data for studying fisheries and environmental impacts on Eastern Pacific ecosystems , 2005 .

[52]  Stuart L. Pimm,et al.  Food web design and the effect of species deletion , 1980 .

[53]  Guy Woodward,et al.  Quantification and Resolution of a Complex, Size-Structured Food Web , 2005 .

[54]  A. Neutel,et al.  Energetics, Patterns of Interaction Strengths, and Stability in Real Ecosystems , 1995, Science.

[55]  R. Law,et al.  A Jump-Growth Model for Predator–Prey Dynamics: Derivation and Application to Marine Ecosystems , 2008, Bulletin of mathematical biology.

[56]  D. L. Angelis,et al.  STABILITY AND CONNECTANCE IN FOOD WEB MODELS , 1975 .

[57]  C. Walters,et al.  An Ecosim model for exploring ecosystem management options for the Gulf of Mexico: implications of including multistanza life history models for policy predictions , 2006 .

[58]  P. Yodzis,et al.  The stability of real ecosystems , 1981, Nature.

[59]  D. Haydon Pivotal Assumptions Determining the Relationship between Stability and Complexity: An Analytical Synthesis of the Stability-Complexity Debate , 1994, The American Naturalist.