[1] Mahidol Univ, Fac Sci, Dept Microbiol, Thailand Hornbill Project, Bangkok 10400, Thailand
[2] Kyoto Univ, Ctr Ecol Res, Otsu, Shiga 5202113, Japan
[3] Univ Shiga Prefecture, Sch Environm Sci, Hikone 5228533, Japan
[4] Res Inst Human & Nat, Kyoto 6020878, Japan
[5] Musashi Univ, Dept Human & Culture, Tokyo 1768534, Japan
[6] Dept Natl Pk Wildlife & Plant Conservat, Natl Pk Div, Bangkok 10900, Thailand