1. Department of Laboratory Medicine and Pathology, University of Washington School of Medicine, Seattle, Washington 98195, USA;
2. Department of Biology, University of Washington, Seattle, Washington 98195, USA
3. Department of Biology, University of Fribourg, CH-1700 Fribourg, Switzerland;
4. Evolution and Ecology Research Centre, School of Biological, Earth and Environmental Sciences, University of New South Wales Sydney, New South Wales 2052, Australia;