Habitat restoration affects immature stages of a wetland butterfly through indirect effects on predation

Habitat loss worldwide has led to the widespread use of restoration practices for the recovery of imperiled species. However, recovery success may be hampered by focusing on plant communities, rather than the complex suite of direct and indirect interactions among trophic levels that occur in natural systems. Through a factorial field experiment, we tested the effects of wetland restoration on egg and juvenile survival of a locally rare butterfly, Satyrodes appalachia, via tree removal and damming. Tree removal more than tripled S. appalachia host plant abundance, but neither restoration action directly affected S. appalachia egg and juvenile survival. Instead, we found strong indirect effects of habitat manipulation on S. appalachia egg and juvenile survival that were mediated through predation. The interaction of tree removal and damming significantly decreased predation of S. appalachia eggs relative to each treatment alone. Damming alone had a significant positive indirect effect on the survival of S. appalachia juveniles, likely because increases in standing water reduced predator access. Our results emphasize the need for experiments that evaluate the demographic responses of imperiled species to habitat restoration prior to management action and quantify potential indirect effects mediated through higher trophic levels.