Mono Basin, on the eastern flank of the central Sierra Nevada, is the highest of the large hydrographically closed basins in the Basin and Range province. We use geomorphic features, shoreline deposits, and basalt-filled paleochannels to reconstruct an early to middle Pleistocene record of shorelines and changing spillways of Lake Russell in Mono Basin. During this period of time, Lake Russell repeatedly attained altitudes between 2205 and 2280 m—levels far above the present surface of Mono Lake (∼1950 m) and above its last overflow level (2188 m). The spill point of Lake Russell shifted through time owing to late Tertiary and Quaternary faulting and volcanism. During the early Pleistocene, the lake periodically discharged through the Mount Hicks spillway on the northeastern rim of Mono Basin and flowed northward into the Walker Lake drainage basin via the East Walker River. Paleochannels recording such discharge were incised prior to 1.6 Ma, possibly between 1.6 and 1.3 Ma, and again after 1.3 Ma (ages of basaltic flows that plugged the paleochannels). Faulting in the Adobe Hills on the southeastern margin of the basin eventually lowered the rim in this area to below the altitude of the Mount Hicks spillway. Twice after 0.76 Ma, and possibly as late as after 0.1 Ma, Lake Russell discharged southward through the Adobe Hills spillway into the Owens–Death Valley system of lakes. This study supports a pre- Pleistocene aquatic connection through Mono Basin between the hydrologically distinct Lahontan and Owens–Death Valley systems, as long postulated by biologists, and also confirms a probable link during the Pleistocene for species adapted to travel upstream in fast-flowing water.