1. Department of Ecology and Evolutionary Biology, Princeton University, Princeton, NJ 08544, USA
2. Centre for Ecology and Conservation, University of Exeter, Penryn Campus, Penryn, Cornwall TR10 9FE, UK
3. Department of Zoology, University of Cambridge, Cambridge CB2 3EJ, UK
4. FitzPatrick Institute of African Ornithology, DST-NRF Centre of Excellence, University of Cape Town, Rondebosch 7701, Cape Town, South Africa