1. University of Hawai`i at Hilo, Tropical Conservation Biology and Environmental Science Program, 200 West Kawili Street, Hilo, Hawai`i 97620, USA
2. U.S. Geological Survey, Pacific Island Ecosystems Research Center, PO Box 44, Hawai`i National Park, Hawai`i 96718, USA
3. University of Hawai`i at Hilo, Department of Biology, 200 West Kawili Street, Hilo, Hawai`i 96720, USA