From Empedocles to Symbiogenetics: Lynn Margulis's revolutionary influence on evolutionary biology

As a primary expositor of the work of Lynn Margulis collaborating with her over thirty years on over thirty books and forty articles, scientific and popular, I attempt here to summarize her unique and lasting influence on evolutionary biology. Describing life on Earth as the multi-billion-year evolution of microbial communities, from prokaryotes maintaining Earth's atmosphere away from thermodynamic equilibrium to all eukaryotes as polygenomic beings, Margulis's interdisciplinary work has deeply influenced multiple fields including systematics, theories of the evolution of metabolism, paleobiology, and biogeochemistry. Overturning the neo-Darwinist narrative that speciation almost always occurs by the gradual accumulation of random mutations, Margulis's work revives a discarded philosophical speculation of the pre-Socratic Empedocles, who suggested that Earth's early beings both merged and differentially reproduced (were naturally selected); a speculation that was rejected by Aristotle probably because it smacked of mythological chimeras that had no place in observational biology, and later by Charles Darwin, who mentioned Aristotle's rejection of Empedocles to show that he knew of but did not accept natural selection, thus helping lay his own claim to its own proper scientific presentation in a Victorian culture whose thinking of origins was dominated not by Greek mythology but Christian special creation. Margulis's curiosity-driven science, collaborative work ethic, status as a woman, embrace of novelty, philosophical stance, current status of her theories, and the proposal for a new science of symbiogenetics are among the topics examined.