Despite over a century of research, the evolutionary origins of collective animal behavior remain unclear. Dozens of hypotheses explaining the evolution of collective behavior have risen and fallen in the past century, but until recently it has been difficult to perform controlled behavioral evolution experiments to isolate these various hypotheses and test their individual effects. In this dissertation, I outline a relatively new method using digital models of evolution to perform controlled behavioral evolution experiments. In particular, I use these models to directly explore the evolutionary consequence of the selfish herd, predator confusion, and the many eyes hypotheses, and demonstrate how the models can lend key insights useful to behavioral biologists, computer scientists, and robotics researchers. This dissertation lays the groundwork for the experimental study of the hypotheses surrounding the evolution of collective animal behavior, and establishes a path for future experiments to explore and disentangle how the various hypothesized benefits of collective behavior interact over evolutionary time.
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