1. FitzPatrick Institute of African Ornithology, DST-NRF Centre of Excellence, University of Cape Town, Private Bag X3, Rondebosch 7701, South Africa
2. Department of Zoology, University of Cambridge, Downing Street, Cambridge CB2 3EJ, UK
3. Centre for Evolutionary Biology, School of Biological Sciences, University of Western Australia, Crawley 6009, Australia