[1] Univ Pretoria, Dept Zool & Entomol, Ctr Invas Biol, Private Bag X 20, ZA-0028 Hatfield, South Africa
[2] South African Inst Aquat Biodivers, ZA-6140 Grahamstown, South Africa
[3] Imperial Coll London, Dept Life Sci, Silwood Pk Campus,Buckhurst Rd, Ascot SL5 7PY, Berks, England
[4] Univ Witwatersrand, Sch Anim Plant & Environm Sci, Ctr Invas Biol, Private Bag 3, ZA-2050 Johannesburg, South Africa
[5] South African Inst Aquat Biodivers, Ctr Invas Biol, ZA-6140 Grahamstown, South Africa