Environmental Variation, Stochastic Extinction, and Competitive Coexistence

Understanding how environmental fluctuations affect population persistence is essential for predicting the ecological impacts of expected future increases in climate variability. However, two bodies of theory make opposite predictions about the effect of environmental variation on persistence. Single‐species theory, common in conservation biology and population viability analyses, suggests that environmental variation increases the risk of stochastic extinction. By contrast, coexistence theory has shown that environmental variation can buffer inferior competitors against competitive exclusion through a storage effect. We reconcile these two perspectives by showing that in the presence of demographic stochasticity, environmental variation can increase the chance of extinction while simultaneously stabilizing coexistence. Our stochastic simulations of a two‐species storage effect model reveal a unimodal relationship between environmental variation and coexistence time, implying maximum coexistence at intermediate levels of environmental variation. The unimodal pattern reflects the fact that the stabilizing influence of the storage effect accumulates rapidly at low levels of environmental variation, whereas the risk of extinction due to the combined effects of environmental variation and demographic stochasticity increases most rapidly at higher levels of variation. Future increases in environmental variation could either increase or decrease an inferior competitor’s expected persistence time, depending on the distance between the present level of environmental variation and the optimal level anticipated by this theory.

[1]  Bond,et al.  Predicting extinction risks for plants: environmental stochasticity can save declining populations. , 2000, Trends in ecology & evolution.

[2]  Joel s. Brown,et al.  The Selective Interactions of Dispersal, Dormancy, and Seed Size as Adaptations for Reducing Risk in Variable Environments , 1988, The American Naturalist.

[3]  M. Boyce Population Viability Analysis , 1992 .

[4]  D. Ackerly,et al.  A trait-based approach to community assembly: partitioning of species trait values into within- and among-community components. , 2007, Ecology letters.

[5]  H. G. Andrewartha,et al.  The distribution and abundance of animals. , 1954 .

[6]  C. Cáceres,et al.  Temporal variation, dormancy, and coexistence: a field test of the storage effect. , 1997, Proceedings of the National Academy of Sciences of the United States of America.

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

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

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

[10]  E. Menges,et al.  Population viability analyses in plants: challenges and opportunities. , 2000, Trends in ecology & evolution.

[11]  P. Chesson,et al.  Short-term instabilities and long-term community dynamics. , 1989, Trends in ecology & evolution.

[12]  M. J. Salinger Climate Variability and Change: Past, Present and Future – An Overview , 2005 .

[13]  Mark Rees,et al.  Effects of Temporal Variability on Rare Plant Persistence in Annual Systems , 2004, The American Naturalist.

[14]  J. Drake Population effects of increased climate variation , 2005, Proceedings of the Royal Society B: Biological Sciences.

[15]  P. Chesson,et al.  DIFFERENCES IN SEED BIOLOGY OF ANNUAL PLANTS IN ARID LANDS: A KEY INGREDIENT OF THE STORAGE EFFECT , 2005 .

[16]  Jianguo Wu,et al.  Ecosystem stability and compensatory effects in the Inner Mongolia grassland , 2004, Nature.

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

[18]  G. E. Hutchinson,et al.  The Balance of Nature and Human Impact: The paradox of the plankton , 2013 .

[19]  J. Downing,et al.  Biodiversity and stability in grasslands , 1996, Nature.

[20]  Peter B Adler,et al.  A niche for neutrality. , 2007, Ecology letters.

[21]  Luis H. R. Alvarez Does increased stochasticity speed up extinction? , 2001, Journal of mathematical biology.

[22]  Peter Chesson,et al.  Coexistence Mediated by Recruitment Fluctuations: A Field Guide to the Storage Effect , 1985, The American Naturalist.

[23]  John M. Drake,et al.  Population Viability Analysis , 2019, Encyclopedia of Theoretical Ecology.

[24]  R. Lande Risks of Population Extinction from Demographic and Environmental Stochasticity and Random Catastrophes , 1993, The American Naturalist.

[25]  Peter Chesson,et al.  Geometry, heterogeneity and competition in variable environments , 1990 .

[26]  D. Doak,et al.  Book Review: Quantitative Conservation biology: Theory and Practice of Population Viability analysis , 2004, Landscape Ecology.

[27]  J. N. Thompson,et al.  The evolution of species interactions. , 1999, Science.

[28]  D. Cohen Optimizing reproduction in a randomly varying environment. , 1966, Journal of theoretical biology.

[29]  John Sabo,et al.  Morris, W. F., and D. F. Doak. 2003. Quantitative Conservation Biology: Theory and Practice of Population Viability Analysis. Sinauer Associates, Sunderland, Massachusetts, USA , 2003 .

[30]  J. HilleRisLambers,et al.  Climate variability has a stabilizing effect on the coexistence of prairie grasses , 2006, Proceedings of the National Academy of Sciences.

[31]  C. E. Pake,et al.  Is Coexistence of Sonoran Desert Annuals Mediated by Temporal Variability Reproductive Success , 1995 .

[32]  M. Boyce,et al.  Demography in an increasingly variable world. , 2006, Trends in ecology & evolution.

[33]  K. Trenberth,et al.  Modern Global Climate Change , 2003, Science.

[34]  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.

[35]  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.

[36]  D. L. Venable,et al.  DORMANCY AND GERMINATION IN A GUILD OF SONORAN DESERT ANNUALS , 2004 .