The Paradox of the Plankton

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Abstract
The principle of competitive exclusion would suggest that at equilibrium only one species of fully autotrophic organism could survive in a uniform volume of water. In general the phytoplankton is highly diversified; the reasons for diversification are however not obvious. It is suggested that symbiosis and commensalism, in the sense of an otherwise better adapted species requiring external metabolites pro-ducedby another species, and possibly the occurrence of specific predators may permit the development of some equilibrium diversity. If the phytoplankton assemblage never reaches interspecific equilibrium . much diversity would be possible, but under these circumstances one would expect more random extinction over a long period of time than seems indicated by the fossil record of diatoms in lake sediments. In lakes it is not unlikely that much of the diversity is due to continual recruitment from the littoral benthos. Where there are good regional bodies of comparable data (Finland, Indonesia), the number of phytoplanktonic species in a lake is apparently independent of its area, as Jarnefelt first found. This can be explained qualitatively if it is supposed that increase in the ratio area littoral length decreases diversity, while increase in littoral length, by increasing the chance of the diversification of shoreline, increases the diversity. Apparent failure of the principle of competitive exclusion provides cases whose analysis may be particularly rewarding.

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