1. School of BioSciences, University of Melbourne , Parkville 3010 , Australia
2. Biological, Earth and Environmental Sciences, The University of New South Wales , Kensington, NSW 2052 , Australia
3. Department of Biology, Pennsylvania State University, University Park , PA 16802 , USA
4. Marine Biological Section, Department of Biology, University of Copenhagen , DK- 3000 Helsingør, Denmark