Weather-normalized Evaluation of Substitute Habitats for Kaloula borealis Reveals a Decoupling between Hydroperiod Buffering and Breeding Opportunity
Amphibians are the most threatened vertebrate group worldwide, largely because their biphasic life cycle depends on both aquatic and terrestrial environments and is highly sensitive to fine-scale hydrological variation. Under accelerating habitat loss and climate-driven shifts in rainfall regimes, substitute habitats are increasingly used as a practical conservation measure. Yet their effectiveness cannot be assumed from physical construction alone. When such sites attract individuals but fail to sustain breeding and recruitment, they may function as ecological traps rather than compensatory habitats. This concern is especially acute for the Boreal digging frog, Kaloula borealis, an explosive breeder whose reproductive success depends on the persistence of temporary aquatic habitat during the larval period. We integrated 191 field surveys from four substitute habitats with daily NASA POWER meteorological data and analyzed habitat performance using a weather-normalized framework that accounted for climatic variability among survey periods. The analyses revealed a clear decoupling between structural performance, defined as the capacity to retain breeding-season water, and functional performance, defined as the capacity to translate retained water into aquatic breeding opportunity. Incheon- Geomdan showed the strongest structural buffering, whereas Gwangmyeong-Siheung exhibited the highest functional breeding performance. Cheonan-Jiksan maintained intermediate but balanced performance across both dimensions, while Asan-Tangjeong was consistently poor in both, indicating potential sink-like characteristics. These findings show that restoration success in amphibian substitute habitats cannot be reduced to a single metric or to breeding records from a single year. Habitat evaluation should instead distinguish settlement, hydrological persistence, and breeding function, and interpret them under standardized climatic conditions. This framework provides a stronger basis for adaptive management and for designing substitute habitats that support long-term population viability rather than temporary occupancy.