1. Research Institute for the Environment and Livelihoods; Charles Darwin University; Darwin Northern Territory 0909 Australia
2. Department of Biological Sciences; California State University; Fullerton California 92834-6850 USA
3. Philip L. Boyd Deep Canyon Research Center; 54-900 Desert Research Trail Indian Wells California 92210 USA
4. Department of Biology; University of Nevada; Reno Nevada 89557 USA