Niche overlap and invasion of competitors in random environments I. Models without demographic stochasticity

Abstract The relationship between persistent, small to moderate levels of random environmental fluctuations and limits to the similarity of competing species is studied. The analytical theory hinges on deriving conditions under which a rare invading species will tend to increase when faced with an array of resident competitors in a fluctuating environment. A general approximation scheme predicts that the effects of low levels of stochasticity will typically be small. The technique is applied explicitly to a class of symmetric, discrete-time stochastic analogs of the Lotka-Volterra equations that incorporate cross-correlation but no autocorrelation. The random environment limits to similarity are always very close to the corresponding constant environment limits. However, stochasticity can either facilitate or hinder invasion. The exact limits to similarity are extremely model-dependent. In addition to the symmetric models, an analytically tractable class of models is presented that incorporates both auto- and cross-correlation and no symmetry assumptions. For all of the models investigated, the analytical theory predicts that small-scale stochasticity does little, if anything, to limit similarity. Extensive Monte Carlo results are presented that confirm the analytical results whenever the dynamics of the discretetime models are biologically reasonable in the sense that trajectories do not exhibit unrealistic crashes. Interestingly, the class of stochastic models that is well behaved in this sense includes models whose deterministic analogs are chaotic. The qualitative conclusion, supported by both the analytical and simulation results, is that for competitive guilds adequately modeled by Lotka-Volterra equations including small to moderate levels of random fluctuations, practical limits to similarity can be obtained by ignoring the stochastic terms and performing a deterministic analysis. The mathematical and biological robustness of this conclusion is discussed.

[1]  P Polansky,et al.  Invariant distributions for multi-population models in random environments. , 1979, Theoretical population biology.

[2]  P. Chesson,et al.  Environmental Variability Promotes Coexistence in Lottery Competitive Systems , 1981, The American Naturalist.

[3]  M W Feldman,et al.  A population's stationary distribution and chance of extinction in a stochastic environment with remarks on the theory of species packing. , 1975, Theoretical population biology.

[4]  Calyampudi R. Rao,et al.  Linear Statistical Inference and Its Applications. , 1975 .

[5]  J. Roughgarden,et al.  Species Packing in Two Dimensions , 1977, The American Naturalist.

[6]  J. Roughgarden Theory of Population Genetics and Evolutionary Ecology: An Introduction , 1995 .

[7]  P. Abrams Niche overlap and environmental variability , 1976 .

[8]  M. Levandowsky,et al.  Randomness, Time Scales, and the Evolution of Biological Communities , 1977 .

[9]  T. Schoener,et al.  Effects of density-restricted food encounter on some single-level competition models. , 1978, Theoretical population biology.

[10]  N Keiding,et al.  Extinction and exponential growth in random environments. , 1975, Theoretical population biology.

[11]  Michael Turelli,et al.  Niche Overlap and Invasion of Competitors in Random Environments II. The Effects of Demographic Stochasticity , 1980 .

[12]  R. Macarthur,et al.  The Limiting Similarity, Convergence, and Divergence of Coexisting Species , 1967, The American Naturalist.

[13]  P. Abrams,et al.  Limiting similarity and the form of the competition coefficient. , 1975, Theoretical population biology.

[14]  Yukio Ogura,et al.  Recurrence properties of Lotka-Volterra models with random fluctuations , 1981 .

[15]  R. May,et al.  Biological populations obeying difference equations: stable points, stable cycles, and chaos. , 1975, Journal of theoretical biology.

[16]  Jonathan Roughgarden,et al.  A Simple Model for Population Dynamics in Stochastic Environments , 1975, The American Naturalist.

[17]  T. Schoener,et al.  Population growth regulated by intraspecific competition for energy or time: some simple representations. , 1973, Theoretical population biology.

[18]  J H Gillespie,et al.  A general model to account for enzyme variation in natural populations. V. The SAS--CFF model. , 1978, Theoretical population biology.

[19]  J. Roughgarden Species packing and the competition function with illustrations from coral reef fish. , 1974, Theoretical population biology.

[20]  Robert M. May,et al.  Stability in Randomly Fluctuating Versus Deterministic Environments , 1973, The American Naturalist.

[21]  T. Schoener Theory of Feeding Strategies , 1971 .

[22]  M E Gilpin,et al.  Competition between species: theoretical models and experimental tests. , 1973, Theoretical population biology.

[23]  M. Bartlett,et al.  A comparison of theoretical and empirical results for some stochastic population models , 1960 .

[24]  M. Frank Norman,et al.  An ergodic theorem for evolution in a random environment , 1975, Journal of Applied Probability.

[25]  M. Turelli,et al.  Does environmental variability limit niche overlap? , 1978, Proceedings of the National Academy of Sciences of the United States of America.

[26]  D. Ludwig Persistence of Dynamical Systems under Random Perturbations , 1975 .

[27]  R M May,et al.  On the theory of niche overlap. , 1974, Theoretical population biology.

[28]  M. Bartlett,et al.  Stochastic Population Models in Ecology and Epidemiology. , 1961 .

[29]  M. Turelli,et al.  Density-dependent selection in a random environment: An evolutionary process that can maintain stable population dynamics. , 1980, Proceedings of the National Academy of Sciences of the United States of America.

[30]  W. D. Evans,et al.  PARTIAL DIFFERENTIAL EQUATIONS , 1941 .

[31]  M. Turelli,et al.  A reexamination of stability in randomly varying versus deterministic environments with comments on the stochastic theory of limiting similarity. , 1978, Theoretical population biology.

[32]  T. Schoener,et al.  Competition and the form of habitat shift. , 1974, Theoretical population biology.

[33]  T. Prout Sufficient Conditions for Multiple Niche Polymorphism , 1968, The American Naturalist.

[34]  R. Lewontin,et al.  On population growth in a randomly varying environment. , 1969, Proceedings of the National Academy of Sciences of the United States of America.

[35]  A. Nold Competitive overlap and coexistence , 1979 .

[36]  Peter Chesson,et al.  Predator-Prey Theory and Variability , 1978 .

[37]  M. Turelli Random environments and stochastic calculus. , 1977, Theoretical population biology.

[38]  R M May,et al.  Niche overlap as a function of environmental variability. , 1972, Proceedings of the National Academy of Sciences of the United States of America.

[39]  R. May,et al.  Stability and Complexity in Model Ecosystems , 1976, IEEE Transactions on Systems, Man, and Cybernetics.

[40]  Floyd B. Hanson,et al.  Persistence in density dependent stochastic populations , 1981 .

[41]  Harold J. Kushner,et al.  Stability and existence of diffusions with discontinuous or rapidly growing drift terms , 1972 .

[42]  H. Levene,et al.  Genetic Equilibrium When More Than One Ecological Niche is Available , 1953, The American Naturalist.

[43]  B. Goh,et al.  Nonvulnerability of ecosystems in unpredictable environments. , 1976, Theoretical population biology.

[44]  H. Tuckwell,et al.  Persistence times of populations with large random fluctuations. , 1978, Theoretical population biology.

[45]  Samuel Karlin,et al.  Random temporal variation in selection intensities acting on infinite diploid populations: diffusion method analysis. , 1975, Theoretical population biology.

[46]  J H Gillespie,et al.  Conditions for the existence of stationary densities for some two-dimensional diffusion processes with applications in population biology. , 1980, Theoretical population biology.

[47]  J. Kiefer,et al.  An Introduction to Stochastic Processes. , 1956 .

[48]  P. Abrams Density-Independent Mortality and Interspecific Competition: A Test of Pianka's Niche Overlap Hypothesis , 1977, The American Naturalist.

[49]  Calyampudi Radhakrishna Rao,et al.  Linear Statistical Inference and its Applications , 1967 .