[1] Tshwane Univ Technol, Dept Nat Conservat, Pretoria, South Africa
[2] South African Environm Observat Network SAEON, Ndlovu Node, Phalaborwa, South Africa
[3] Univ Witwatersrand, Sch Geog Archaeol & Environm Studies, Johannesburg, South Africa
[4] Univ Calif Santa Barbara, Dept Ecol Evolut & Marine Biol, Santa Barbara, CA 93106 USA
[5] South African Natl Pk, Sci Serv, Skukuza, South Africa
[6] South African Natl Biodivers Inst, Pretoria, South Africa