Drought degrades riparian subsidy quality and constrains aquatic ecosystem functioning

The exchange of energy and organisms across habitat boundaries links aquatic and terrestrial ecosystems and sustains ecosystem functioning. Although disturbance may disrupt these linkages, the mechanisms at play remain poorly understood. Here, we investigated the extent to which flow intermittency may disrupt riparian-aquatic ecosystem linkages by altering consumer communities in the recipient ecosystem or by altering resource quality in the donor ecosystem. We ran an experiment in an intermittent river network in California, focusing on a critical forest-to-river subsidy (organic matter in the form of leaf litter), its transformation, and its reciprocal benefit (aquatic insect production). Using three riparian species (willow, cottonwood, and oak) at sites spanning a gradient of flow permanence, we quantified intraspecific plasticity in leaf traits (specific leaf area, nitrogen and phosphorus concentrations, and {delta}13C), measured decomposition rates, and estimated the secondary production of aquatic shredders (Plecoptera). Across all leaf species, decomposition rates were 16-36% lower at intermittent than perennial sites, an effect largely driven by intraspecific leaf trait plasticity rather than changes in consumer abundance. At high flow intermittency, willow experienced water stress (enriched {delta}13C) and reduced specific leaf area, while cottonwood showed primarily stoichiometric responses (reduced leaf nitrogen and phosphorus). Despite these divergent strategies, all species produced lower-quality litter at intermittent sites. Variance partitioning confirmed that initial litter quality uniquely explained 51.5% of variation in decomposition rates, more than double the contribution of invertebrate community metrics; and structural equation modeling revealed that both leaf traits and stonefly (Plecoptera) secondary production significantly predicted decomposition rates, with leaf traits exerting the stronger effect. Notably, stonefly secondary production was 37-98% lower at intermittent sites across leaf species. Because these insects later emerge as terrestrial adults, they provide a significant energy flux to riparian predators, and, thus, impoverished litter quality suppresses the reciprocal transfer of energy back to terrestrial food webs. As drought intensifies globally, the decoupling of terrestrial-aquatic linkages may begin in the riparian canopy.