[1] US Geol Survey, Pacific Isl Ecosyst Res Ctr, Honolulu, HI 96818 USA
[2] Univ Hawaii Manoa, Dept Plant & Environm Protect Sci, Honolulu, HI 96822 USA
[3] Univ Hawaii Manoa, Sch Life Sci, Honolulu, HI USA
[4] Univ Hawaii Manoa, Pacific Cooperat Studies Unit, Honolulu, HI 96822 USA