1. Division of Environmental and Evolutionary Biology, Institute of Biomedical and Life Sciences, University of Glasgow, Glasgow G12 8QQ, UK
2. School of Biological, Earth and Environmental Science, The University of New South Wales, Sydney 2052, NSW, Australia
3. School of Mathematics, University of Bristol, University Walk, Bristol BS8 1TW, UK
4. School of Biological Sciences, University of Bristol, Woodland Road, Bristol BS8 1UG, UK