All Sizes Fit the Red Queen

Abstract
The Red Queen's hypothesis portrays evolution as a never-ending competition for expansive energy, where one species’ gain is another species’ loss. The Red Queen is neutral with respect to body size, implying that neither small nor large species have a universal competitive advantage. Here we ask whether, and if so how, the Red Queen's hypothesis really can accommodate differences in body size. The maximum population growth in ecology clearly depends on body size—the smaller the species, the shorter the generation length, and the faster it can expand given sufficient opportunity. On the other hand, large species are more efficient in energy use due to metabolic scaling and can maintain more biomass with the same energy. The advantage of shorter generation makes a wide range of body sizes competitive, yet large species do not take over. We analytically show that individuals consume energy and reproduce in physiological time, but need to compete for energy in real time. The Red Queen, through adaptive evolution of populations, balances the pressures of real and physiological time. Modeling competition for energy as a proportional prize contest from economics, we further show that Red Queen's zero-sum game can generate unimodal hat-like patterns of species rise and decline that can be neutral in relation to body size.

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