Origins and basic logic
Hutchinson (1961) argued that if the environment changes before reaching equilibrium, competitive exclusion is never completed and many species can coexist. Grime (1973) proposed a hump-shaped relationship in grasslands, where intermediate stress such as mowing or grazing suppresses dominant species and raises diversity, and Horn (1975) treated forest succession with probabilistic models. Explaining why tropical rainforests and coral reefs are so diverse, Joseph Connell (1978) drew these ideas together as the intermediate disturbance hypothesis. Right after a disturbance only fast-colonizing pioneer species arrive, and after long periods without disturbance strong late-successional competitors dominate through competitive exclusion; at intermediate levels pioneers and late species mix, so diversity peaks.
Experimental tests and theoretical extensions
Sousa (1979) studied boulders of different sizes in the California intertidal zone that waves overturn at different rates. Small, frequently overturned boulders held only pioneer green algae, large boulders that rarely moved were dominated by competitively superior red algae, and intermediate boulders held the most species. Huston (1979) proposed a dynamic equilibrium model in which disturbance frequency and population growth rates jointly determine diversity, and Pickett and White (1985) established disturbance and patch dynamics as central topics in ecology. Petraitis, Latham and Niesenbaum (1989) systematized the mechanisms by which disturbance maintains diversity, and Roxburgh, Shea and Wilson (2004) and Shea, Roxburgh and Rauschert (2004) modeled the conditions under which hump-shaped patterns reflect genuine coexistence mechanisms.
Evidence and the debate over abandonment
Reviewing more than 100 studies, Mackey and Currie (2001) reported that hump-shaped disturbance–diversity relationships appear only in a minority of cases, with monotonic increases, decreases or no relationship also common. Fox (2013) argued that the hypothesis should be abandoned, because the logic that disturbance interrupts competition and thereby promotes coexistence is theoretically flawed and the empirical support is weak. Sheil and Burslem (2013) countered that Connell's original hypothesis concerned the interaction of disturbance and succession and remains useful in tropical forests and elsewhere. The debate has led to the understanding that disturbance affects diversity not through a single pattern but in ways that vary with ecosystem, scale and type of disturbance. The hypothesis is treated as a classic theory of community diversity alongside keystone species and island biogeography.
FAQ
- What is the intermediate disturbance hypothesis in simple terms?
- If disturbances like fires or storms are too frequent, only tough species survive; if too rare, strong competitors take over. At moderate levels many species live together, so diversity is highest.
- Is the hypothesis still accepted?
- It is widely taught as a classic theory, but empirical studies do not consistently find the hump-shaped pattern and there are theoretical criticisms, so whether to abandon it is debated.
Sources reviewed: 2026-10-26
References
- Connell, J. H. (1978). Diversity in tropical rain forests and coral reefs. Science, 199(4335), 1302–1310.
- Grime, J. P. (1973). Competitive exclusion in herbaceous vegetation. Nature, 242(5396), 344–347.
- Horn, H. S. (1975). Markovian properties of forest succession. In M. L. Cody & J. M. Diamond (Eds.), Ecology and Evolution of Communities (pp. 196–211). Cambridge, MA: Harvard University Press.
- Hutchinson, G. E. (1961). The paradox of the plankton. American Naturalist, 95(882), 137–145.
- Sousa, W. P. (1979). Disturbance in marine intertidal boulder fields: The nonequilibrium maintenance of species diversity. Ecology, 60(6), 1225–1239.
- Huston, M. (1979). A general hypothesis of species diversity. American Naturalist, 113(1), 81–101.
- Pickett, S. T. A., & White, P. S. (Eds.). (1985). The Ecology of Natural Disturbance and Patch Dynamics. Orlando: Academic Press.
- Petraitis, P. S., Latham, R. E., & Niesenbaum, R. A. (1989). The maintenance of species diversity by disturbance. Quarterly Review of Biology, 64(4), 393–418.
- Roxburgh, S. H., Shea, K., & Wilson, J. B. (2004). The intermediate disturbance hypothesis: Patch dynamics and mechanisms of species coexistence. Ecology, 85(2), 359–371.
- Mackey, R. L., & Currie, D. J. (2001). The diversity–disturbance relationship: Is it generally strong and peaked? Ecology, 82(12), 3479–3492.
- Shea, K., Roxburgh, S. H., & Rauschert, E. S. J. (2004). Moving from pattern to process: Coexistence mechanisms under intermediate disturbance regimes. Ecology Letters, 7(6), 491–508.
- Fox, J. W. (2013). The intermediate disturbance hypothesis should be abandoned. Trends in Ecology & Evolution, 28(2), 86–92.
- Sheil, D., & Burslem, D. F. R. P. (2013). Defining and defending Connell's intermediate disturbance hypothesis: A response to Fox. Trends in Ecology & Evolution, 28(10), 571–572.
