1. Department of Biology, McGill University, 1205 Avenue Dr Penfield, H3A 1B1, Montréal, Québec, Canada
2. These authors contributed equally to this work
3. Present address: The Jackson Laboratory, 600 Main Street, Bar Harbor, Maine 04609, USA
4. Present address: Genzentrum, Ludwig-Maximilians Universitaet, Feodor-Lynen-Str. 25, D-81377, Munich, Germany