1. Centre for Evolutionary Biology, School of Biological Sciences, University of Western Australia, Perth, Western Australia, Australia 6009
2. School of Natural Sciences, Macquarie University, Sydney, New South Wales, Australia 2109
3. FitzPatrick Institute of African Ornithology, University of Cape Town, Cape Town, South Africa, 7701
4. Centre for Ecology and Conservation, University of Exeter, Penryn, TR10 9FE, UK