The endocranial anatomy of protocetids and its implications for early whale evolution

Extant whales, dolphins, and porpoises result from a major macroevolutionary lifestyle transition that transformed land-dwelling cetaceans into fully aquatic species. This involved significant changes in sensory systems. The increase in brain size relative to body size (encephalization quotient) is an outstanding feature of modern cetaceans, especially toothed whales. Conversely, olfactory capabilities are assumed to have diminished along this transition, with airborne olfaction becoming less relevant. The extent and timing of olfactory reduction remain obscure due to challenges in accessing well-preserved fossil endocranial anatomy. This study shows that early cetaceans had already evolved an increased encephalization quotient, and that their olfactory apparatus was likely not yet under selective pressure leading to its reduction. We demonstrate this through an analysis of the extinct whale, Protocetus atavus, a member of the middle Eocene semiaquatic cetacean group Protocetidae. We provide the first documentation of its endocranial anatomy using high-resolution computed tomography and compare it to other early cetaceans as well as extant mammals. We conclude that cetaceans increased their brain size earlier than previously thought, while relying on a well-developed olfactory system at a time when they were still partly terrestrial.