[1] Univ Melbourne, Inst Bio21, Dept Zool, Melbourne, Vic 3010, Australia
[2] Univ Melbourne, Inst Bio21, Dept Genet, Melbourne, Vic 3010, Australia
[3] Monash Univ, Sch Biol Sci, Clayton, Vic 3800, Australia
[4] Monash Univ, Australian Ctr Biodivers, Clayton, Vic 3800, Australia