The role of the hyporheic zone for a benthic fish in an intermittent river: a refuge, not a graveyard

In lotic systems, the hyporheic zone has been suggested as a potential refuge for aquatic organisms during disturbances (hyporheic refuge hypothesis). However, the supporting evidence is unclear, especially regarding the survival of hyporheic refugees and their contribution to the recovery of post-disturbance populations. Moreover, few studies have focused on the importance of the hyporheic refuge for aquatic vertebrates such as fish. In this study, we present evidence that the hyporheic zone acts as a refuge for a small benthic fish (Cobitis shikokuensis) following surface drying in an intermittent river. We examined its survival during and recolonization after dry periods by direct hyporheic sampling and mark-and-recapture surveys. When the streambed dried, hyporheic sampling was conducted 58 times across 33 locations in the intermittent reach and 31 individuals of C. shikokuensis were captured from extracted hyporheic water. Mark-and-recapture surveys revealed that recolonizers after re-wetting included C. shikokuensis individuals that had survived dry periods in the hyporheic refuge. The condition factor of C. shikokuensis significantly declined after dry periods, suggesting that most recolonizers suffered from physiological stress, probably within the hyporheic refuge. These results clearly support the long-debated, hyporheic refuge hypothesis, and provide a striking example of the critical role of the hyporheic zone in population maintenance of lotic organisms.