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Two features characterize all natural communities. First, they are dynamic systems. The densities and age-structures of populations change with time, as do the relative abundances of species; local extinctions are commonplace (37). For many communities, a self-reproducing climax state may only exist as an average condition on a relatively large spatial scale, and even that has yet to be rigorously demonstrated (36). The idea that equilibrium is rarely achieved on the local scale was expressed decades ago by a number of forest ecologists (e.g. 101, 168). One might even argue that continued application of the concept of climax to natural systems is simply an exercise in metaphysics (41). While this view may seem extreme, major climatic shifts often recur at time intervals shorter than that required for a community to reach competitive equilibrium or alter the geographical distributions of species (6, 21, 43, 76, 92). Climatic variation of this kind influences ecological patterns over large areas, sometimes encompassing entire continents. Other agents of temporal change in natural communities operate over a wide range of smaller spatial scales (47, 242). Second, natural communities are spatially heterogeneous. This statement is true at any scale of resolution (242), but it is
Annual Review of Ecology, Evolution, and Systematics – Annual Reviews
Published: Nov 1, 1984
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