Path-based network unfolding: A solution for the problem of mixed trophic and non-trophic processes in trophic dynamic analysis.

The purpose of this paper is to describe a quantitative method of trophic dynamic analysis derived from a systems ecology theoretical foundation. This method was devised to provide a solution for the problem of how to deal with mixed trophic and non-trophic processes in cyclic ecosystem networks, a problem that has vexed trophic ecology since Lindeman first presented a formal concept of trophic dynamics in 1942. The author's initial attempt to solve this problem was presented in Whipple & Patten (1993, J. theor. Biol. 163, 393-411). The path-based network unfolding method described in this paper provides a quantitative method for conducting trophic dynamic analysis of cyclic ecosystems containing non-living storages and non-trophic flows to produce a true energy-transformation trophic macrochain. This method solves the "trophic-level inflation" problem described in Whipple & Patten (193, J. theor. Biol. 163, 393-411). The results of the analysis of an oyster reef ecosystem model demonstrate that the dual trophic macrochain produced by path-based network unfolding may be used to compare the relative contribution of grazing and detrital sub-webs to the trophic dynamics of ecosystems. It was found that the standing stock and flow contribution of the detrital sub-web was quantitatively dominant in the oyster reef ecosystem model. This method might be used to compare the contribution of grazing and detrital sub-webs for models of different ecosystem types. Because a true energy transformation trophic chain is produced, the progressive efficiency concept of the Lindeman-Hutchinson paradigm may be applied in comparative trophic analyses of ecosystems. In comparing the oyster reef model results of three quantitative trophic analysis methods, the path-based network unfolding method was found to produce a trophic macrochain with progressive efficiencies intermediate between those produced by the original Higashi et al. method and the Burns et al. unfolding analysis of a modified version of the oyster reef model.Copyright 1998 Academic Press Limited