Supplementary analysis of the model.

To see how the flow of energy across ecosystems can derive evolution, I introduce a framework in which individuals interact with their peers and environment to accumulate resources, and use the resources to pay for their metabolic costs, grow and reproduce. I show that two conservation principles determine the system’s equilibrium state: conservation of resources- a physical principle stating that in the equilibrium, resource production and consumption should balance, and payoff equality- an economic principle, stating that the payoffs of different types in equilibrium should equal. Besides the equilibrium state, the system shows non-equilibrium fluctuations derived by the exponential growth of the individuals in which the payoff equality principle does not hold. A simple gradient-ascend dynamical mean-field equation predicts the onset of non-equilibrium fluctuations. As an example, I study the evolution of cooperation in public goods games. In both mixed and structured populations, cooperation evolves naturally in resource-poor environments but not in resource-rich environments. Population viscosity facilitates cooperation in poor environments but can be detrimental to cooperation in rich environments. In addition, cooperators and defectors show different life-history strategies: Cooperators live shorter lives and reproduce more than defectors. Both population structure and, more significantly, population viscosity reduce lifespan and life history differences between cooperators and defectors.