Diversity, variability and persistence elements for a non-equilibrium theory of eco-evolutionary dynamics

Abstract

Natural ecosystems persist in variable environments by virtue of a suite of traits that span from the individual to the community, and from the ecological to the evolutionary scenarios. How these internal characteristics operate to allow living beings to cope with the uncertainty present in their environments is the subject matter of quantitative theoretical ecology. Under the framework of structural realism, the present dissertation project has advocated for the strategy of mathematical modeling as a strategy of abstraction. The goal is to explore if a range of natural ecosystems display the features of complex systems, and evaluate whether these features provide insights into how they persist in their current environments, and how might they cope with changing environments in the future. A suite of inverse, linear and non-linear dynamical mathematical models, including non-equilibrium catastrophe models, and structured demographic approaches is applied to five case studies of natural systems fluctuating in the long-term in diverse scenarios: phytoplankton in the global ocean, a mixotrophic plankton food web in a marine coastal environment, a wintering waterfowl community in a major Mediterranean biodiversity hot-spot, a breeding colony of a keystone avian scavenger in a mountainous environment and the shorebird community inhabiting the coast of UK. In all case studies, there is strong evidence that ecosystems are able to closely track their common environment through several strategies. For example, in global phytoplankton communities, a latitudinal gradient in the positive impact of functional diversity on community stability counteracts the increasing environmental variability with latitude. Mixotrophy, by linking several feeding strategies in a food web, internally drives community dynamics to the edge of instability while maximizing network complexity. In contrast, an externally generated major perturbation, operating through planetary climatic disruptions, induce an abrupt regime shift between alternative stable states in the wintering waterfowl community. Overall, the natural systems studied are shown to posses features of complex systems: connectivity, autonomy, emergence, non-equilibrium, non-linearity, self-organization and coevolution. In rapidly changing environments, these features are hypothesized to allow natural system to robustly respond to stress and disturbances to a large extent. At the same time, future scenarios will be probably characterized by conditions never experienced before by the studied systems. How will they respond to them, is an open question. Based on the results of this dissertation, future research directions in theoretical quantitative ecology will likely benefit from non-autonomous dynamical system approaches, where model parameters are a function of time, and from the deeper exploration of global attractors and the non-equilibriumness of dynamical systems.