1. Theoretical Biology Institute of Integrative Biology ETH Zürich, Universitätstrasse 16 CH‐8092 Zürich Switzerland
2. CoMPLEX University College London Gower Street London WC1E 6BT United Kingdom
3. The Galton Laboratory Department of Genetics Environment, and Evolution University College London Gower Street London WC1E 6BT United Kingdom
4. Department of Biology Faculty of Sciences Kyushu University Fukuoka 812‐8581 Japan