1. Department of Zoology and Biodiversity Research Centre; University of British Columbia; 6270 University Boulevard Vancouver BC V6T 1Z4 Canada
2. Centre for Ecology & Conservation; University of Exeter; Penryn Campus Penryn TR10 9FE United Kingdom
3. Current address : Center for Interdisciplinary Research in Biology, CNRS UMR 7241; Equipe Stochastic Models for the Inference of Life Evolution, Collège de France; 11 place Marcelin Berthelot 75231 Paris Cedex 5 France