Abstract
Endosymbiosis, symbiosis in which one symbiont lives inside another, is woven throughout the history of life and the story of its evolution. From the mitochondrion residing in almost every eukaryotic cell to the gut microbiome found in every human, endosymbiosis is a cornerstone of the biological processes that sustain life on Earth. While endosym-biosis is ubiquitous, many questions about its origins remain shrouded in mystery; one question in particular regards the general conditions and possible trajectories for its evolution. Modern science has hypothesized two possible pathways for the evolution of mutualistic endosymbiosis: one where an obligate antagonism is co-opted into an obligate mutualism (Co-Opted Antagonism Hypothesis), and one where a facultative mutualism evolves into an obligate mutualism (Black Queen Hypothesis). We investigated the viability of these pathways under different environmental conditions by expanding on the evolutionary agent-based system Symbulation. Specifically, we considered the impact of ectosymbiosis on de novo evolution of obligate mutualistic endosymbiosis. We found that introducing a facultative ectosymbiotic state allows endosym-biosis to evolve in a more diverse set of environmental conditions, while also decreasing the evolution of endosymbiosis in conditions where it can evolve independently.