[1] Khon Kaen Univ, Fac Sci, Dept Biol, Khon Kaen 40002, Thailand
[2] Kasetsart Univ, Fac Sci, Dept Zool, Anim Systemat & Ecol Special Res Unit ASESRU, Bangkok 10900, Thailand
[3] Museum Zool, Pl Riponne 6, CH-1005 Lausanne, Switzerland
[4] Univ Lausanne, Dept Ecol & Evolut, CH-1015 Lausanne, Switzerland
[5] Khon Kaen Univ, Fac Sci, Appl Taxon Res Ctr ATRC, Khon Kaen 40002, Thailand