1. Flinders University 1 School of Science and Engineering , , Bedford Park, SA 5042 , Australia
2. Research School of Biology, The Australian National University 2 Division of Ecology and Evolution , , Canberra, ACT 2600 , Australia
3. Stellenbosch Institute for Advanced Study (STIAS), Wallenberg Centre 3 , 10 Marais Street, Stellenbosch 7600 , South Africa