Spite (in the biological or evolutionary sense) is behavior that harms others at a cost to the actor. The presence of spite in human and animal populations presents an evolutionary puzzle. Recent work has suggested small populations and pre-play signaling can have a significant effect on the evolution of spite. Here, we use computational methods to explore these factors in fluctuating populations that may go extinct. We find that the presence of spite can make a population significantly more

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... ely to go extinct, but that this does not preclude the possibility of spite reliably evolving. Additionally, we find that the stochastic effects of small fluctuating populations allow for the evolution and predominance of signal-mediated conditional spite across a wide range of conditions. These results suggest that directed harm, even if costly, can play a significant early role in the evolution of social behaviors and this provides a possible origin for punishment.