RESUMEN
Watershed resilience is the ability of a watershed to maintain its characteristic system state while concurrently resisting, adapting to, and reorganizing after hydrological (for example, drought, flooding) or biogeochemical (for example, excessive nutrient) disturbances. Vulnerable waters include non-floodplain wetlands and headwater streams, abundant watershed components representing the most distal extent of the freshwater aquatic network. Vulnerable waters are hydrologically dynamic and biogeochemically reactive aquatic systems, storing, processing, and releasing water and entrained (that is, dissolved and particulate) materials along expanding and contracting aquatic networks. The hydrological and biogeochemical functions emerging from these processes affect the magnitude, frequency, timing, duration, storage, and rate of change of material and energy fluxes among watershed components and to downstream waters, thereby maintaining watershed states and imparting watershed resilience. We present here a conceptual framework for understanding how vulnerable waters confer watershed resilience. We demonstrate how individual and cumulative vulnerable-water modifications (for example, reduced extent, altered connectivity) affect watershed-scale hydrological and biogeochemical disturbance response and recovery, which decreases watershed resilience and can trigger transitions across thresholds to alternative watershed states (for example, states conducive to increased flood frequency or nutrient concentrations). We subsequently describe how resilient watersheds require spatial heterogeneity and temporal variability in hydrological and biogeochemical interactions between terrestrial systems and down-gradient waters, which necessitates attention to the conservation and restoration of vulnerable waters and their downstream connectivity gradients. To conclude, we provide actionable principles for resilient watersheds and articulate research needs to further watershed resilience science and vulnerable-water management.