The three faces of ecological fitness

This paper argues that fitness is most usefully understood as those properties of organisms that are explanatory of survival in the broadest sense, not merely descriptive of reproductive success. Borrowing from Rosenberg and Bouchard (2009), fitness in this sense is ecological in that it is defined by the interactions between organisms and environments. There are three sorts of ecological fitness: the well-documented ability to compete, the ability to cooperate (as in mutualistic symbiosis), and a third sense of fitness that has received insufficient attention in evolutionary theory, the ability to construct. Following Lotka, it can be understood thermodynamically as the ability to maintain or enlarge the energy-circulating capacity of an ecosystem. An organism that does this could end with its gene frequency unchanged but its probability of survival enhanced since it would sustain or increase the total carrying capacity of its ecosystem. Photosynthesizers and other autotrophs are obvious candidates for organisms that are fit in the constructive sense, but any organisms, including heterotrophs, can exhibit constructive fitness if they have some mechanism for channeling external flows of free energy into their ecosystems. I will briefly examine the prospects for the human species in the light of these considerations.