The North American rodent fossil record includes hundreds of species representing both an incredible taxonomic diversity and great ecological disparity. Although it is during the Oligocene that taxonomic diversity first peaks, it is not until the Miocene, almost 10 Myr later, that many ecologies, particularly locomotory ecologies, are recorded. Here, I present a new Oligocene-aged species of beaver from Montana, Microtheriomys articulaquaticus sp. nov., which represents the oldest semi-aquatic rodent in North America and the oldest amphibious beaver in the world, pushing the advent of semi-aquatic ecology in beavers by 7 Myr. I also provide morphological data supporting a terrestrial ecology for the sister taxon to Castoridae. Together with existing data, these findings lead to a new hypothesis for the evolutionary ecology of castorids whereby swimming was exapted from burrowing during the Oligocene. This evolution of semi-aquatic locomotion may have taken place in North America instead of Eurasia. It started in small beavers with gigantism achieved only much later. Indeed, body size evolution in castoroids follows a directional drift. Beavers obey Cope's rule, a selection for larger size over time that appears associated with semi-aquatic ecology and may well explain their low modern diversity.
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