Climate variability and periodic droughts have complex effects on carbon (C) fluxes, with uncertain implications for ecosystem C balance under a changing climate. Responses to climate change can be modulated by persistent effects of climate history on plant communities, soil microbial activity, and nutrient cycling (i.e., legacies). To assess how legacies of past precipitation regimes influence tallgrass prairie C cycling under new precipitation regimes, we modified a long-term irrigation experiment that simulated a wetter climate for >25 years. We reversed irrigated and control (ambient precipitation) treatments in some plots and imposed an experimental drought in plots with a history of irrigation or ambient precipitation to assess how climate legacies affect aboveground net primary productivity (ANPP), soil respiration, and selected soil C pools. Legacy effects of elevated precipitation (irrigation) included higher C fluxes and altered labile soil C pools, and in some cases altered sensitivity to new climate treatments. Indeed, decades of irrigation reduced the sensitivity of both ANPP and soil respiration to drought compared with controls. Positive legacy effects of irrigation on ANPP persisted for at least 3 years following treatment reversal, were apparent in both wet and dry years, and were associated with altered plant functional composition. In contrast, legacy effects on soil respiration were comparatively short-lived and did not manifest under natural or experimentally-imposed “wet years,” suggesting that legacy effects on CO2 efflux are contingent on current conditions. Although total soil C remained similar across treatments, long-term irrigation increased labile soil C and the sensitivity of microbial biomass C to drought. Importantly, the magnitude of legacy effects for all response variables varied with topography, suggesting that landscape can modulate the strength and direction of climate legacies. Our results demonstrate the role of climate history as an important determinant of terrestrial C cycling responses to future climate changes.