1. U.S. Geological Survey–Biological Resources Discipline, Pacific Island Ecosystems Research Center, PO Box 44, Hawai‘i National Park, Hawai‘i 96718, USA
2. Current address: Pu‘u Kukui Watershed Preserve, 1000 Kapalua Drive, Lahaina, Hawai‘i 96761, USA
3. Pacific Cooperative Studies Unit, University of Hawai‘i, Mānoa, Honolulu, Hawai‘i 96822, USA
4. Current address: Department of Epidemiology and Public Health, Yale University, New Haven, Connecticut 06520, USA