Ecological stability

An ecosystem is said to possess ecological stability (or equilibrium) if it is capable of returning to its equilibrium state after a perturbation (a capacity known as resilience) or does not experience unexpected large changes in its characteristics across time.[1] Although the terms community stability and ecological stability are sometimes used interchangeably,[2] community stability refers only to the characteristics of communities. It is possible for an ecosystem or a community to be stable in some of their properties and unstable in others. For example, a vegetation community in response to a drought might conserve biomass but lose biodiversity.[3]

Stable ecological systems abound in nature, and the scientific literature has documented them to a great extent. Scientific studies mainly describe grassland plant communities and microbial communities.[4] Nevertheless, it is important to mention that not every community or ecosystem in nature is stable (for example, wolves and moose on Isle Royale). Also, noise plays an important role on biological systems and, in some scenarios, it can fully determine their temporal dynamics.

The concept of ecological stability emerged in the first half of the 20th century. With the advancement of theoretical ecology in the 1970s, the usage of the term has expanded to a wide variety of scenarios. This overuse of the term has led to controversy over its definition and implementation.[3]

In 1997, Grimm and Wissel made an inventory of 167 definitions used in the literature and found 70 different stability concepts.[5] One of the strategies that these two authors proposed to clarify the subject is to replace ecological stability with more specific terms, such as constancy, resilience and persistence. In order to fully describe and put meaning to a specific kind of stability, it must be looked at more carefully. Otherwise the statements made about stability will have little to no reliability because they would not have information to back up the claim.[6] Following this strategy, an ecosystem which oscillates cyclically around a fixed point, such as the one delineated by the predator-prey equations, would be described as persistent and resilient, but not as constant. Some authors, however, see good reason for the abundance of definitions, because they reflect the extensive variety of real and mathematical systems.[3]

Stability analysis

When the species abundances of an ecological system are treated with a set of differential equations, it is possible to test for stability by linearizing the system at the equilibrium point.[7] Robert May developed this stability analysis in the 1970s which uses the Jacobian matrix.

Types

Although the characteristics of any ecological system are susceptible to changes, during a defined period of time, some remain constant, oscillate, reach a fixed point or present other type of behavior that can be described as stable.[8] This multitude of trends can be labeled by different types of ecological stability.

Dynamical stability

Dynamical stability refers to stability across time.

Stationary, stable, transient, and cyclic points

A stable point is such that a small perturbation of the system will be diminished and the system will come back to the original point. On the other hand, if a small perturbation is magnified, the stationary point is considered unstable.

Local and global stability

Local stability indicates that a system is stable over small short-lived disturbances, while global stability indicates a system highly resistant to change in species composition and/or food web dynamics.

Constancy

Observational studies of ecosystems use constancy to describe living systems that can remain unchanged.

Resistance and inertia (persistence)

Resistance and inertia deal with a system's inherent response to some perturbation.

A perturbation is any externally imposed change in conditions, usually happening in a short time period. Resistance is a measure of how little the variable of interest changes in response to external pressures. Inertia (or persistence) implies that the living system is able to resist external fluctuations. In the context of changing ecosystems in post-glacial North America, E.C. Pielou remarked at the outset of her overview,

"It obviously takes considerable time for mature vegetation to become established on newly exposed ice scoured rocks or glacial till...it also takes considerable time for whole ecosystems to change, with their numerous interdependent plant species, the habitats these create, and the animals that live in the habitats. Therefore, climatically caused fluctuations in ecological communities are a damped, smoothed-out version of the climatic fluctuations that cause them."[9]

Resilience, elasticity and amplitude

Resilience is the tendency of a system to retain its functional and organizational structure and the ability to recover after a perturbation or disturbance. Resilience also expresses the need for persistence although from a management approach it is expressed to have a broad range of choices and events are to be looked at as uniformly distributed.[10] Elasticity and amplitude are measures of resilience. Elasticity is the speed with which a system returns to its original / previous state. Amplitude is a measure of how far a system can be moved from the previous state and still return. Ecology borrows the idea of neighborhood stability and a domain of attraction from dynamical systems theory.

Lyapunov stability

Researchers applying mathematical models from system dynamics usually use Lyapunov stability.[11][12]

Numerical stability

Focusing on the biotic components of an ecosystem, a population or a community possesses numerical stability if the number of individuals is constant or resilient.[13]

Sign stability

It is possible to determine if a system is stable just by looking at the signs in the interaction matrix. 

Stability and diversity

The relation between diversity and stability has been widely studied.[4] Diversity can operate to enhance the stability of ecosystem functions at various ecological scales.[14] For example, genetic diversity can enhance resistance to environmental perturbations.[15] At the community level, the structure of food webs can affect stability. The effect of diversity on stability in food-web models can be either positive or negative, depending on the trophic coherence of the network.[16] At the level of landscapes, environmental heterogeneity across locations has been shown to increase the stability of ecosystem functions [17]

History of the concept

The term 'oekology' was coined by Ernst Haeckel in 1866. Ecology as a science was developed further during the late 19th and the early 20th century, and increasing attention was directed toward the connection between diversity and stability.[18] Frederic Clements and Henry Gleason contributed knowledge of community structure; among other things, these two scientists introduced the opposing ideas that a community can either reach a stable climax or that it is largely coincidental and variable. Charles Elton argued in 1958 that complex, diverse communities tended to be more stable. Robert MacArthur proposed a mathematical description of stability in the number of individuals in a food web in 1955.[19] After much progress made with experimental studies in the 60's, Robert May advanced the field of theoretical ecology and refuted the idea that diversity begets stability.[20] Many definitions of ecological stability have emerged in the last decades while the concept continues to gain attention.

See also

Notes
  1. A., Levin, Simon; R., Carpenter, Stephen (2012-01-01). The Princeton guide to ecology. Princeton University Press. p. 790. ISBN 9780691156040. OCLC 841495663.
  2. "Ecology/Community succession and stability - Wikibooks, open books for an open world". en.wikibooks.org. Retrieved 2017-05-02.
  3. Robert May & Angela McLean (2007). Theoretical Ecology: Principles and Applications (3rd ed.). pp. 98–110. ISBN 9780199209989.
  4. Ives, Anthony R.; Carpenter, Stephen R. (2007-07-06). "Stability and Diversity of Ecosystems". Science. 317 (5834): 58–62. Bibcode:2007Sci...317...58I. doi:10.1126/science.1133258. ISSN 0036-8075. PMID 17615333. S2CID 11001567.
  5. Grimm, V.; Wissel, Christian (1997-02-01). "Babel, or the ecological stability discussions: an inventory and analysis of terminology and a guide for avoiding confusion". Oecologia. 109 (3): 323–334. Bibcode:1997Oecol.109..323G. doi:10.1007/s004420050090. ISSN 0029-8549. PMID 28307528. S2CID 5140864.
  6. Gigon, Andreas (1983). "Typology and Principles of Ecological Stability and Instability". Mountain Research and Development. 3 (2): 95–102. doi:10.2307/3672989. ISSN 0276-4741. JSTOR 3672989.
  7. Carlos., Castillo-Chávez (2012-01-01). Mathematical Models in Population Biology and Epidemiology. Springer New York. ISBN 9781461416869. OCLC 779197058.
  8. Lewontin, Richard C. (1969). "The Meaning of Stability". Brookhaven Symposia in Biology. 22: 13–23. PMID 5372787.
  9. Pielou, After the Ice Age: The Return of Life to Glaciated North America (Chicago: University of Chicago Press) 1991:13
  10. Holling, C. S. (1973). "Resilience and Stability of Ecological Systems" (PDF). Annual Review of Ecology and Systematics. 4: 1–23. doi:10.1146/annurev.es.04.110173.000245. ISSN 0066-4162. JSTOR 2096802.
  11. Justus, James (2006). "Ecological and Lyanupov Stability" (PDF). Paper presented at the Biennial Meeting of The Philosophy of Science Association, Vancouver, Canada.
  12. Justus, J (2008). "Ecological and Lyanupov Stability". Philosophy of Science. 75 (4): 421–436. CiteSeerX 10.1.1.405.2888. doi:10.1086/595836. S2CID 14194437.(Published version of above paper)
  13. A., Levin, Simon; R., Carpenter, Stephen (2012-01-01). The Princeton guide to ecology. Princeton University Press. p. 65. ISBN 9780691156040. OCLC 841495663.
  14. Oliver, Tom H.; Heard, Matthew S.; Isaac, Nick J.B.; Roy, David B.; Procter, Deborah; Eigenbrod, Felix; Freckleton, Rob; Hector, Andy; Orme, C. David L. (2015). "Biodiversity and Resilience of Ecosystem Functions" (PDF). Trends in Ecology & Evolution. 30 (11): 673–684. doi:10.1016/j.tree.2015.08.009. PMID 26437633.
  15. Forsman, Anders; Wennersten, Lena (2016-07-01). "Inter-individual variation promotes ecological success of populations and species: evidence from experimental and comparative studies". Ecography. 39 (7): 630–648. doi:10.1111/ecog.01357. ISSN 1600-0587.
  16. Johnson S, Domı́nguez-Garcı́a V, Donetti L, Muñoz MA (2014). "Trophic coherence determines food-web stability". Proc Natl Acad Sci USA. 111 (50): 17923–17928. arXiv:1404.7728. Bibcode:2014PNAS..11117923J. doi:10.1073/pnas.1409077111. PMC 4273378. PMID 25468963.CS1 maint: multiple names: authors list (link)
  17. Wang, Shaopeng; Loreau, Michel (2014-08-01). "Ecosystem stability in space: α, β and γ variability". Ecology Letters. 17 (8): 891–901. doi:10.1111/ele.12292. ISSN 1461-0248. PMID 24811401.
  18. Elton, Charles S. (1927-01-01). Animal Ecology. University of Chicago Press. ISBN 9780226206394.
  19. MacArthur, Robert (1955-01-01). "Fluctuations of Animal Populations and a Measure of Community Stability". Ecology. 36 (3): 533–536. doi:10.2307/1929601. JSTOR 1929601.
  20. May, Robert M. (1972-08-18). "Will a Large Complex System be Stable?". Nature. 238 (5364): 413–414. Bibcode:1972Natur.238..413M. doi:10.1038/238413a0. PMID 4559589. S2CID 4262204.

References
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