1. Université de Lyon, F-69000, Lyon; Université Lyon 1; CNRS, UMR5558; Laboratoire de Biométrie et Biologie Evolutive; F-69622; Villeurbanne France
2. MRC Centre for Outbreak Analysis and Modelling; Department of Infectious Disease Epidemiology; Imperial College - School of Public Health; London UK
3. Office National de la Chasse et de la Faune Sauvage; Montfort Birieux France