1. School of Mathematics and Statistics, Centre for Research into Ecological and Environmental Modelling, University of St Andrews, St Andrews, United Kingdom
2. United States Geological Survey, Pacific Island Ecosystems Research Center, Hawai‘i National Park, Hawai‘i, United States
3. Current Affiliation: Biomathematics and Statistics Scotland, Dundee, Scotland and UK Centre for Ecology & Hydrology (UKCEH), Lancaster Environment Centre, Lancaster, United Kingdom
4. U.S. Fish and Wildlife Service, Hilo, Hawai‘i, United States