1. Division of Evolution, Ecology and Genetics; Research School of Biology; The Australian National University; Canberra ACT 0200 Australia
2. Department of Ecology, Evolution and Behavior; University of Minnesota; St. Paul MN 55108 USA
3. Centre of Excellence in Biological Interactions; Division of Evolution, Ecology and Genetics; Research School of Biology; The Australian National University; Canberra ACT 0200 Australia