1. Department of Genetics, Evolution and Environment; University College London; Gower Street London WC1E 6BT UK
2. Centre for Mathematics and Physics in the Life Sciences and Experimental Biology; University College London; London UK
3. Centre for Biodiversity and Environment Research; University College London; Gower Street London WC1E 6BT UK
4. Department of Statistical Science; University College London; Gower Street London WC1E 6BT UK