1. Pacific Cooperative Studies Unit (University of Hawaii at Mānoa), c/o USGS Pacific Island Ecosystems Research Center, Kīlauea Field Station, P.O. Box 44, Hawaii National Park, HI 96718
2. Current address: Department of Biological Sciences, Virginia Polytechnic Institute and State University, 2125 Derring Hall, Blacksburg, Virginia 24061, USA
3. USGS Pacific Island Ecosystems Research Center, Kīlauea Field Station, P.O. Box 44, Hawaii National Park, HI 96718