1. Department of Zoology, University of Otago , Dunedin , New Zealand
2. Department of Ecology, Evolution, and Organismal Biology, Iowa State University , Ames, IA , USA
3. Department of Biology, Iowa State University , Ames, IA , USA
4. Department of Plant Pathology, Entomology, and MIcrobiology, Iowa State University , Ames, IA , USA