Climax community

The Daintree Rainforest in Queensland, Australia is an example of a climax forest ecosystem.

Warren Woods in Michigan, USA is an example of a Beech-Maple climax forest. Beech (pictured) and Sugar Maple (bottom left) dominate the forest due to their towering height and tolerance to shade.

In ecology, climax community, or climatic climax community, is a historic term that expressed a biological community of plants, animals, and fungi which, through the process of ecological succession the development of vegetation in an area over time, had reached a steady state. This equilibrium was thought to occur because the climax community is composed of species best adapted to average conditions in that area. The term is sometimes also applied in soil development. Nevertheless, it has been found that a "steady state" is more apparent than real, particularly if long-enough periods of time are taken into consideration. Notwithstanding, it remains a useful concept.

The idea of a single climatic climax, which is defined in relation to regional Climate, originated with Frederic Clements in the early 1900s. The first analysis of succession as leading to something like a climax was written by Henry Cowles in 1899, but it was Clements who used the term "climax" to describe the idealized endpoint of succession.^[1]

Frederic Clements's use of "climax"

Clements described the successional development of an ecological communities comparable to the ontogenetic development of individual organisms.^[2] Clements suggested only comparisons to very simple organisms.^[3] Later ecologists developed this idea that the ecological community is a "superorganism" and even sometimes claimed that communities could be homologous to complex organisms and sought to define a single climax-type for each area. The English botanist Arthur Tansley developed this idea with the "polyclimax" — multiple steady-state end-points, determined by edaphic factors, in a given climatic zone. Clements had called these end-points other terms, not climaxes, and had thought they were not stable, because by definition climax vegetation is best-adapted to the climate of a given area. Henry Gleason's early challenges to Clements's organism simile, and other of his strategies for describing vegetation, were largely disregarded for several decades until substantially vindicated by research in the 1950s and 1960s (below). Meanwhile, climax theory was deeply incorporated in both theoretical ecology and in vegetation management. Clements's terms such as pre-climax, post-climax, plagioclimax and disclimax continued to be used to describe the many communities which persist in states that diverge from the climax ideal for a particular area.

Though the views are sometimes attributed to him, Clements never argued that climax communities must always occur, or that the different species in an ecological community are tightly integrated physiologically, or that plant communities have sharp boundaries in time or space. Rather, he employed the idea of a climax community — of the form of vegetation best adapted to some idealized set of environmental conditions — as a conceptual starting point for describing the vegetation in a given area. There are good reasons to believe that the species best adapted to some conditions might appear there, when those conditions occur. But much of Clements's work was devoted to characterizing what happens when those ideal conditions do not occur. In those circumstances, vegetation other than the ideal climax will often occur instead. But those different kinds of vegetation can still be described as deviations from the climax ideal. Therefore, Clements developed a very large vocabulary of theoretical terms describing the various possible causes of vegetation, and various non-climax states vegetation adopts as a consequence. His method of dealing with ecological complexity was to define an ideal form of vegetation — the climax community — and describe other forms of vegetation as deviations from that ideal.^[4]

Discomfort with climax theory

Support among ecologists for the climax theory declined, because they found the theory with its many coined terms difficult to apply, because they were dissatisfied how it compared to observed individual organisms, and because better theories developed.^[5]

Although Clements recognized that vegetation follows gradients rather than being tightly bound, his rhetorical comparisons of ecological communities to organisms fostered the impression that communities, including the climax, have distinct edges in space and time. Yet Robert Whittaker's research demonstrated plant species distribute themselves along nutrient and other environmental gradients.^[6] Many ecologists saw this as a major reason to stop using the climax concept.

More recent palynological studies showed that modern species assemblages are ephemeral; vegetation in eastern North America since the last glacial maximum has consisted of several different species assemblages, many of which have no analogues in modern "climax" communities. That would mean, at least, that the climax types for those areas could not be stable to the degree Clements believed they were.

Ultimately, even if succession tends towards a steady state, the time required to achieve this state is unrealistically long; in most cases, external disturbances and environmental change occur so frequently that the realization of a climax community is unlikely, and therefore it has come to be regarded as a less useful concept. Long-term vegetation dynamics are now more often characterized as resulting from the action of stochastic factors.^[7]

Continuing usage of "climax"

Despite the overall abandonment of climax theory, during the 1990s use of climax concepts again became more popular among some theoretical ecologists.^[8] Many authors and nature-enthusiasts continue to use the term "climax" in a diluted form to refer to what might otherwise be called mature or old-growth communities. The term "climax" has also been adopted as description for a late successional stage for marine macroinvertebrate communities.^[9]

Additionally, some contemporary ecologists still use the term "disclimax" to describe an ecosystem been dominated by invasive species that competitively prevent the re-introduction of once native species. This concept borrows from Clement's earliest interpretation of climax as referring to an ecosystem that is resistant to colonization by outside species. The term disclimax was used in-context by Clements (1936), and despite being an anthropogenic phenomenon which prevents the facilitation and succession to a true climax community, it is one of the only examples of climax that can be observed in nature.^[10][11]

References

1. Cowles, Henry Chandler. 1899. The Ecological Relations of the Vegetation on the Sand Dunes of Lake Michigan. Botanical Gazette 27(2): 95-117; 27(3): 167-202; 27(4): 281-308; 27(5): 361-391.
2. Clements, Frederic E. 1916. Plant Succession: An Analysis of the Development of Vegetation. Washington D.C.: Carnegie Institution of Washington.
3. Hagen, Joel B. 1992. An Entangled Bank: The Origins of Ecosystem Ecology. New Brunswick: Rutgers University Press.
4. Eliot, Christopher. 2007. Method and Metaphysics in Clements’s and Gleason’s Ecological Explanations. Studies in History and Philosophy of Biological and Biomedical Sciences 38(1): 85–109.
5. Tobey, Ronald C. 1981. Saving the prairies: the life cycle of the founding school of American plant ecology, 1895–1955. Berkeley: University of California Press.
6. Whittaker, Robert H. 1953. A consideration of climax theory: the climax as a population and pattern. Ecological Monographs 23: 41–78.
7. Cook, James E. 1996. Implications of Modern Successional Theory for Habitat Typing: A Review. Forest Science 42(1): 67–75.
8. See, for example, Roughgarden, Jonathan, Robert M. May and Simon A. Levin, editors. 1989. Perspectives in Ecological Theory. Princeton: Princeton University Press.
9. Rosenberg R., S. Agrenius, B. Hellman, H. C. Nilsson, and K. Norling. 2002. Recovery of marine benthic habitats and fauna in a Swedish fjord following improved oxygen conditions. Marine Ecology Progress Series 234: 43-53.
10. Clements, Frederic E. 1936. Nature and Structure of the Climax. Journal of Ecology. Vol. 24, No. 1, pp. 252-284
11. Johnson, K. 1984. Prairie and plains disclimax and disappearing butterflies, in the central United States. Atala. Vol. 10-12, pp. 20-30