1. Department of Biology; University of Massachusetts; 221 Morrill Science Center Amherst MA 01003 USA
2. Organismic and Evolutionary Biology Program; University of Massachusetts; 319 Morrill Science Center Amherst MA 01003 USA
3. Department of Biological Sciences; North Carolina State University; Raleigh NC 27695 USA
4. Institute of Biodiversity, Animal Health & Comparative Medicine; University of Glasgow; Glasgow G12 8QQ UK