The term climax community, also described as a climatic climax community is a largely obsolete ecological term for a biological community of plants and animals which, through the process of succession--the development of vegetation in an area over time--has reached an equilibrium or steady state. This equilibrium occurs because the climax community is composed of species best adapted to average conditions in that local area. The term is sometimes also applied in soil development. 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.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.

Frederic Clements's use of "climax"

Clements described the successional development of an ecological communities as comparable to the ontogenetic development of individual organisms.Clements, Frederic E. 1916. Plant Succession: An Analysis of the Development of Vegetation. Washington D.C.: Carnegie Institution of Washington. Though later ecologists developed this idea of the ecological community as a "superorganism" and even sometimes claimed that communities could be homologous to complex organisms, Clements's use of the organism simile suggested only comparisons to very simple organisms.Hagen, Joel B. 1992. An Entangled Bank: The Origins of Ecosystem Ecology. New Brunswick: Rutgers University Press. Clements's theory, which sought to define a single climax-type for each area, was further developed in Arthur Tansley's idea of the "polyclimax," which allowed for multiple steady-state end-points, determined by edaphic factors, in a given climatic zone. Clements had not referred to these other end-points as climaxes, assigning other terms to them, and had not thought them 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 diverging 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 dominant cause of vegetation is climate, 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 from of vegetation--the climax community--and describe other forms of vegetation as deviations from that ideal.

Rejection of climax theory

A combination of the unwieldiness of employing Clements's theory, with its terribly many coined terms, dissatisfaction with comparisons to individual organisms, and later developments in the field of ecology led to a decline in support for climax theory among ecologists. Though Clements recognized that all vegetation follows gradients, rather than being tightly bounded, his rhetorical comparisons of ecological communities to organisms fostered the impression that communities, including the climax, have distinct edges in space and time. Accordingly, Robert Whittaker's research demonstrating that plant species distribute themselves along nutrient and other environmental gradients was a major impetus for many ecologists for discontinuing to use the climax concept. More recently, palynological studies have shown that modern species assemblages are ephemeral entities; 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 being resulting from of the action of stochastic factors.

Continuing usage of "climax"

Despite this overall abandonment of climax theory, during the 1990s use of climax concepts again became more popular among some theoretical ecologists.See, for example, Roughgarden, Jonathan, Robert M. May and Simon A. Levin, editors. 1989. Perspectives in Ecological Theory. Princeton: Princeton University Press. Though climax theory is no longer itself popular in ecology, 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.

Notes

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See also

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• Ecosystem

Ecology

Klimaks | Climax | Climax (ecologie) | 極相