1. Tropical Conservation Biology and Environmental Science, University of Hawai‘i at Hilo, 200 West Kawili Street, Hilo, HI 96720, USA.
2. Department of Biology, University of Hawai‘i at Hilo, 200 West Kawili Street, Hilo, HI 96720, USA.
3. U.S. Geological Survey, Pacific Island Ecosystem Research Center, P.O. Box 44, Hawaii National Park, HI 96718, USA.