1. School of Biology, University of St Andrews, St Andrews, Fife KY16 9TH, UK
2. Institute of Evolutionary Biology, School of Biological Sciences, King's Buildings, University of Edinburgh, Edinburgh EH9 3JT, UK
3. Institute of Biomedical and Life Sciences, Division of Environmental and Evolutionary Biology, Graham Kerr Building, University of Glasgow, Glasgow G12 8QQ, UK
4. PO Box 603, Gaborone, Botswana