1. Department of Ecology and Evolution, University of Lausanne, 1015 Lausanne, Switzerland
2. Department of Mathematics and Statistics, Bristol Institute of Technology, University of the West of England, Bristol BS16 1QY, UK
3. School of Biological Sciences, University of Bristol, Bristol BS8 1UB, UK
4. Bristol Centre for Complexity Sciences, Department of Engineering Mathematics, University of Bristol, Bristol BS8 1UB, UK