[1] Univ Sains Malaysia, Sch Biol Sci, Usm Penang 11800, Malaysia
[2] Univ Sains Malaysia, Ctr Marine & Coastal Studies, George Town 11800, Malaysia
[3] La Sierra Univ, Dept Biol, Herpetol Lab, 4500 Riverwalk Pkwy, Riverside, CA 92515 USA
[4] San Diego Nat Hist Museum, Dept Herpetol, POB 121390, San Diego, CA 92112 USA
[5] Univ Malaysia Sabah, Inst Trop Biol & Conservat, Jalan UMS, Kota Kinabalu 88400, Sabah, Malaysia
[6] Natl Univ Singapore, Lee Kong Chian Nat Hist Museum, 2 Conservatory Dr, Singapore 117377, Singapore