Implications of variability on many time scales for scientific advice on sustainable management of living marine resources

The conceptual basis for understanding and management of living marine resources is built on three basic ecological principles developed in the first half of the past century: the law of the minimum, competitive exclusion, and succession. This paper highlights aspects of these principles that make them insufficient as a sound foundation for understanding and managing marine ecosystems, points out dangers of continuing to use approaches built on them, and presents alternatives which might be more appropriate and of lower risk. To do this, the paper considers variability of marine ecosystems on annual, medium and long-term time scales, highlighting that these scales correspond to less than, approximately equal to, and much greater than, the generation times of dominant predators in the systems. It also considers how each interval of variability may affect directly ecosystems which are controlled from the bottom up, top down, and middle outward, and how position and duration of forcing affect five types of responses: growth, maturation, recruitment, predation, and competition. Generally these five processes have manifestations at the scale of individuals, populations, and ecosystems, attention is drawn to which manifestations are the most significant for each duration and position of forcing. Effects of some combinations of duration of forcing and position of forcing can be explained reasonably well by conventional ecological theory. For other combinations, particularly forcing at time scales of predator generations on top-down or middle-out ecosystems, theory based on contest competition and equilibria are likely to be misleading. In these systems the major dynamics are transients, when many ecosystems are far from their carrying capacities, so scramble competition dominates, and the carrying capacity is not helpful in explaining the system dynamics. This review clarifies the sorts of questions that we should be asking, in order to begin to understand the transient behaviour of these non-equilibrium ecosystems. The answers to the new classes of questions may lead to great improvements in how ecosystems are managed, as well as how their variation is explained. (C) 2001 Elsevier Science Ltd. All rights reserved.