1. Department of Zoology, University of Cambridge, Cambridge CB2 3EJ, UK
2. Department of Physiology, Development and Neuroscience, University of Cambridge, Cambridge CB2 3EL, UK
3. School of Life Sciences, University of Lincoln, Lincoln LN6 7TS, UK
4. Department of Ecology, Evolution and Behaviour, University of Minnesota, Saint Paul, MN 55108, USA