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Hutchinson's Heritage: the Diversity-Disturbance Relationship In View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by OceanRep Hydrobiologia 249: 1-7, 1993. J. Padisctk, C.S. Reynolds & U. Sommer (eds), Intermediate Disturbance Hypothesis in Phytoplankton Ecology. 1 0 1993 Kluwev Academic Publishers. Printed in Belgium. Hutchinson’s heritage: the diversity-disturbance relationship in phytoplankton U. Sommer’, J. Padisak2, C. S. Reynolds3 & P. Juhasz-Nagy4 1Institut fir Biologie und Chemie des Meeres, Universitat Oldenburg, PO@ 2503, D-2900 Oldenburg, Germany; 2Botanical Department of the Hungarian Natural History Museum/Balaton Limnological Institute of the Hungarian Academy of Sciences, H-8237 Tihany, Hungary; 3Freshwater Biological Association, NERC Institute of Freshwater Ecology, Windermere Laboratory, Ambleside, LA22 OLP, UK; 4Department of Plant Taxonomy and Ecology, Eiitviis Lorand University, HI083 Budapest, Ludovika te’r 2, Hungary Key words: the paradox of plankton, Intermediate Disturbance Hypothesis, diversity, general ecology, community changes Abstract This paper introduces a collection of contributions presented at the 8th Workshop of the International Association of Phytoplankton Taxonomy and Ecology. It compares the substance of with what to limnologists is the more familiar ‘paradox of the plankton’ posed by G. E. Hutchinson. The utility of Connell‘s Intermediate Disturbance Hypothesis in plankton ecology is, potentially, more instructive but inherent difficulties in relating response to stimulus have to be overcome. A copy of the brief distributed to contributors before the workshop is appended. Preamble ilimnetic phytoplankton coexist in a well mixed environment and compete for a very small num- All modern students of limnology will have ben- ber of common limiting resources (light and a few efited from the influence of the late Professor nutritional elements). Nevertheless, even in small G. Evelyn Hutchinson. The thoroughness and er- samples of water, it is not unusual to find more udition of his monumental Treatise on Limnology than 30 species of phytoplankton present. Equi- (Hutchinson, 1957a, 1967,1975a) imparted librium concepts predict that competition would knowledge, understanding and inspiration to all eventually select the best fit species to the exclu- engaged in the study of limnetic and lotic ecosys- sion of all others (Hardin, 1960) and that only as tems. Those of us who have specialized in pelagic many species can coexist as there are limiting ecology hold a special regard for Hutchinson’s factors. The search for solution to this apparent challenge that the high diversity of species present paradox has occupied plankton biologists ever in limnetic plankton is inconsistent with the as- since. sumptions of competitive exclusion in the sup- Going into a bit deeper into Hutchinson’s her- posed uniformity of pelagic environments. In- itage, it is interesting to observe the thread of deed, the dilemma has become widely known as reasoning running through his masterly book ‘An the ‘paradox of the plankton’ (Hutchinson, 1961). Introduction to Population Ecology’ (Hutchin- Hutchinson was intrigued by the fact that ep- son, 1978). This begins with a consideration of 2 temporal processes, enriched by a very deep feel- In his seminal paper, Hutchinson (1961) sug- ing for science history, and goes on to cover such gested several possible explanations for his exciting topics as ‘Living Together in Theory and palanktonic paradox of diversity. Among them Practice’ (Chapter 4) or ‘How Is Living Nature was the idea that the boundary conditions of Put Together?’ (Chapter 6). The book is full of competition change frequently enough to revert very interesting examples and speculations on the competitive hierarchies before exclusion occurs. possible interconnections between limnic and ter- The proposal was reformulated by Richerson restrial ecology. et al. (1970) as ‘contemporaneous disequilibrium’. In making his perceptive generalizations, it may Later attempts to solve the paradox include the- be noted that, in addition to his own deep inten- oretical analyses (Grenney, 1973; Armstrong & tions, Hutchinson was influenced by a number of McGehee, 1976; Levins, 1979, Ebenhbh, 1988; friends, students, including Charles Elton, David Grover, 1990) and experimental approaches Lack, R. H. MacArthur, R. H. Whittaker & J. L. (Turpin & Harrison, 1980; Robinson & Sandgren, Harper. It is interesting to observe, however, that 1983; Sommer, 1984,1985; Gaedeke & Sommer, his attitude is always critical, as is shown, for 1986; Grover, 1988,1989,1991). In spite of these example, in his questioning comments on some considerable efforts, no universally accepted ex- statements of Lack (1954) or Slobodkin (1961). planation for the paradox has been forthcoming. We believe that at least three closely related Although freshwater biologists associate the topics, have been relevant in motivating Hutch- problem of species diversity with Hutchinson, it inson‘s appreciation: is fair to say that analogous questions have been (i) the nature of ecological patterns, prevalent among terrestrial ecologists. For in- (ii) the nature of niche, stance, Grime (1973) was among the first to re- (iii) and the nature of coexistence. cognize that although competition can be a major Topic (i) can be followed reasonably well from causal factor in the maintenance of low diversi- Hutchinson (1953) through MacArthur (1965) to ties in herbaceous vegetation, there is neverthe- Hutchinson (1975b). Topic (ii), the story of less a low incidence of competitive species in flo- ‘niche-ology’, is also fairly well accepted, started, ristically rich habitats. Wilson (1990) considered in a sense, by Hutchinson (1957b) and continued twelve possible mechanisms to account for spe- by the activities of R. H. MacArthur, R. K. Col- cies coexistence in New Zealand plant commu- well, I. Emlen, I. Futuyama, M. E. Gilpin, I. nities: most of these are equilibrium concepts, Roughgarden, who are, directly or indirectly, stu- including niche diversification and stabilizing co- dents of Hutchinson. evolution; his non-equilibrium explanations as- The third topic (iii) is the most difficult one: it sumed that there had not been enough time for was also the most pertinent problem of Hutchin- competition to completely exclude relevant spe- son. Following his publications carefully, from cies before sudden or gradual alterations inter- the rather early papers (e.g. Hutchinson, 1948, vene to arrest or shift back the development of the 1951, 1953, 1954) to his most famous syntheses association of species. (Hutchinson, 1959, 1961, 1978), we discern a re- Still more perplexing is the astonishing diver- current view that the rebuses of coexistence would sity observed among tropical forests and in coral be solved through the search for potential optima, reef communities in which an equilibrated climax the recognision and definition of moving patterns condition is intuitively anticipated; competitive and where the interpretation of which remained exclusion appears, again paradoxically, not to within the context of evolutionary ecology. have been effective. Connell (1978) argued that, Needless to say, all the components of such a even here, the time taken to recover from damage reasoning have induced several divergent lines of through change of climatic conditions, landslides, further investigations (e. g. Levins, 1968, 1979; fires or violent storms is such for it to be difficult Lewontin & Cohen, 1969, etc.). for competitive exclusion to occur finally. Thus 3 the intervention of factors delaying progress to- storm damage are likely to be recolonized first by wards, or preventing the attainment of, an equi- invasive species. These will generally be those librium condition is supposed to be responsible which can provide the largest number of for the maintenance of the diversity. This has propagules or which can expand their biomass become widely known as ‘Connell’s Intermediate the most rapidly, assuming that the new condi- Disturbance Hypothesis’. tions are adequate to meet their requirements. In time, however, these colonist species will be re- placed by others which are better equipped to live The Intermediate Disturbance Hypothesis and its close to the limits of the available resources applicability to phytoplankton (McArthur & Wilson, 1967). As the community is assembled, the ratio between the production Connell’s (1978) hypothesis states: and the biomass falls simultaneously until, ulti- (1) In the absence of disturbance (eternal mately, the biomass supported is the maximum steady state), competitive exclusion will reduce achievable with the energy available. diversity to minimal levels. The application of successional theory to the (2) Under very intense disturbance only few seasonally changing structure of phytoplankton populations of pioneer species could establish assemblages has been attempted by Reynolds themselves after each disturbance event. This (1988). His approach depended upon the recog- would also lead to minimal diversity. nition of the importance (a) of the temporal scal- (3) If disturbances are of intermediate fre- ing, which relates primarily to the life spans of quency and/or intensity there will be repeated op- consecutive generations of algae (measured in portunities for the re-establishment of pioneer hours to days rather than months to decades), populations which would otherwise be outcom- and (b) that different species of phytoplankton peted and the populations of the successful com- show adaptations
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