Leaf litter patches in stream create overwintering habitats for Ezo brown frog (Rana pirica)

Only fragments are known about the functions of leaf litter patches in streams for terrestrial organisms. We investigated the movement patterns of Ezo brown frogs (Rana pirica) on two occasions, in autumn 2014 and summer 2015, and the occurrence of frogs in rivers and along riverbanks on three occasions from summer to autumn 2014 along a 1.5-km stream segment that flows through the Bekanbeushi wetland, northern Japan. Ezo brown frog movement was biased toward the river channel in late autumn and this movement pattern coincided with relatively abundant Ezo brown frogs on the riverbed from autumn to winter 2014. The distribution shift was also supported by decreasing abundance of Ezo brown frogs on riverbanks from autumn to winter (no sighting in winter) during daytime observations. The abundance of overwintering Ezo brown frogs in the channel was associated most positively with leaf litter dry mass and depth, with flow velocity being relatively less important at the quadrat scale (0.063 m(2)). Our results demonstrated that Ezo brown frogs overwinter in the stream and suggest that leaf litter patches positively affect the quality of the overwintering habitat.