1. Departments of Zoology and Botany and Beaty Biodiversity Museum, University of British Columbia, 6270 University Boulevard, Vancouver, British Columbia, Canada V6T 1Z4; 2Departments of Zoology and Biodiversity Research Centre, University of British Columbia, 6270 University Boulevard, Vancouver, British Columbia, Canada V6T 1Z4 and 3Department of Biological Sciences, Macquarie University, Sydney NSW 2109, Australia