The Disturbing History of Intermediate DisturbanceDavid M. WilkinsonOikos, Vol. 84, No. 1. (Jan., 1999), pp. 145-147.Stable URL:http://links.jstor.org/sici?sici=0030-1299%28199901%2984%3A1%3C145%3ATDHOID%3E2.0.CO%3B2-ROikos is currently published by Nordic Society Oikos.Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available athttp://www.jstor.org/about/terms.html. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtainedprior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content inthe JSTOR archive only for your personal, non-commercial use.Please contact the publisher regarding any further use of this work. 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Discussion and debate about important issues in ecology, e.g. theory or terminology, may I I also be included. Contributions should be as precise as possible and references should be P 0 kept to a minimum. A summary is not required. 0 N The disturbing history of intermediate disturbance David M. Wilkinson, Biology and Earth Sciences, Livevpool John Mooves Univ., Byvom Street, Livevpool, UK L3 3AF ([email protected]). 'A person who is not interested in the growth and flux of ideas is probably not interested in the life of the mind. A ... scientist working in an advancing field of research should certainly try to identify the origin and growth of current opinion.' Medawar (1979) A recent review introduced the idea of intermediate disturbance thus: 'One of the best-known consequences of disturbance is reduction in the proportionate abun- dance of competitively dominant species. as in Con- nell's (1978) 'intermediate disturbance hypothesis'. Connell proposed that too little disturbance leads to low diversity through competitive exclusion. and too much disturbance eliminates species incapable of rapid re-colonization' (Hoopes and Harrison 1998: 146). They go on to claim that 'Intermediate disturbance has become perhaps one of the best-accepted principles in ecology'. This is a significant complement, given the trouble ecology has experienced in identifying general principles of potentially predictive use (McIntosh 1995). This identification of the intermediate disturbance hy- pothesis with Connell's 1978 paper is shared by nearly all ecology textbooks (e.g. Ricklefs 1990, Colinvaux 1993, Brewer 1994 and Krebs 1994), the partial excep- tion to this being Begon et al. (1996) who credit Con- nell but then write 'see also the earlier account by Horn 1975'. If the idea is indeed 'one of the best-accepted principles in ecology' then surely its real history is of some interest and should replace the pseudo-history given in the textbooks. Many current textbooks when describing the idea of intermediate disturbance make use of a graph similar to Fig. 1. Rudwick (1992) has stressed the importance of illustration in the history of science rather than just using textual analysis. In this respect this 'hump-backed' graph is very useful as it makes explicit the authors' ideas on the relationship between diversity and disturbance. Connell (1978) presents such a graph (his Fig. 1) as does Horn (1975: 209). In his legend for this figure Horn wrote 'Note that intermediate disturbances produce higher diversity than either very high or very low levels' (Horn 1975: 209). A very clear statement of the intermediate disturbance hypothesis three years be- fore the paper the textbooks cite as its origin. This graph has an even longer history. In his paper on 'Competitive exclusion in herbaceous vegetation' Grime (1973a) produced a graph (his Fig. 2) showing the characteristic hump-backed relationship between 'spe-cies density' (i.e. species richness per unit area) and both 'environmental stress' and 'intensity of manage- ment' (defined as 'intensity of grazing, mowing etc.' i.e. disturbance). Grime (1973a: 345) pointed out that Odum (1963) had previously observed that 'the greatest diversity occurs in the moderate or middle range of a Q Disturbanceto Fig. 1. A generalised version of the hump-backed graphical model of the intermediate disturbance hypothesis.physical gradient' but cites no prior source for the similar relationship with disturbance. Grime (1973a) provides the earliest graph linking disturbance and species richness that I am aware off, five years before Connell. He reprinted this graphical model in a paper in the Journal of Encironmental Management published in the same year as the Nature paper (Grime 1973b). In a later book Grime (1979: 163) provided a more gener- alised version of this graph calling it the 'hump-backed model'. If this characteristically shaped graph is taken as central to the intermediate disturbance hypothesis then it should be Grime rather than Connell who is cited as its originator. It will come as no surprise to anyone with an interest in the history of science that the intermediate distur- bance hypothesis does not emerge fully formed during the 1970's without any prior history. Scientific ideas seldom appear 'from nowhere' without a history of earlier related ideas. A classic illustration of this are the attempts at theories of evolution and selection prior to Darwin