Inverse reconstructions of ecosystem flows in investigating regime shifts: impact of the choice of objective function

Inverse analysis is increasingly used in ecosystem modelling to objectively reconstruct a large number of unknown flows or interactions from a small number of observations. This type of analysis may be useful in relating observed regime shifts in ecosystem structure to underlying processes. Inversions of ecosystem flow networks currently use a constrained least-squares solution which at the same time minimizes the squared norm (the sum of squares) of the reconstructed flows. This minimum norm (MN) inversion is thought to be a parsimonious solution to the ecosystem flow inverse problem, but it may well not reflect how ecosystems are organised. It has been proposed instead that ecosystems evolve to maximize energy/mass flows or that they maximize the information content of the network weighted by ecosystem flows (ascendancy). We used simulated inverse experiments, where inverse analyses are applied to simulations of flow networks, to explore objective functions different than the MN generally used. We could not compute inverse solutions that maximize ascendancy because the objective function is unbounded. We could calculate inversions that maximize flows; however, these generally overestimated the simulated flows, even though the simulations were designed to maximize flows. It appears that the ecosystem flow inverse problem is too under-determined (too few data relative to the number of unknowns) to allow the use of these maximizing goal functions. We introduce a new minimization that simultaneously minimizes the squared flows and the squared differences between flows. This smoothing minimization makes the inverse flows as even as possible and it helps with some technical issues with MN inversions. The simulated inverse experiments indicated that this smoothed norm (SM) is the most robust in comparative analyses of contrasting ecosystem states, such as those that can be associated with regime shifts. Like the MN inversion, the SM inversion has no ecological basis. However, it is a conservative norm that is less likely to produce false differences between the dynamics of regimes.