“…Traditional detection-based monitoring is timeconsuming and labor-intensive (Hanya, Yoshihiro, Zamma, Kubo, & Takahata, 2003;Whitesides, Oates, Green, & Kluberdanz, 1988), and the precision of identifying species or age-sex classes may depend on the ability of the observers. Methods based on animal traces, such as feces, nests or footprints, enable researchers to collect more data per unit effort than detection-based methods (Delibes et al, 2012;Hanya et al, 2017;Kanamori, Kuze, Bernard, Malim, & Kohshima, 2017). However, one can only obtain limited information on the individual animal that left the trace compared with direct observation (Hanya et al, 2017).…”