[1] James Cook Univ, Australian Trop Herbarium, Cairns, Qld 4870, Australia
[2] Natl Museum Nat Hist, Dept Bot, Smithsonian Inst, Washington, DC 20013 USA
[3] James Cook Univ, Ctr Trop Biodivers & Climate Change, Cairns, Qld 4870, Australia
[4] CSIRO Ecosyst Sci, Cairns, Qld 4870, Australia
[5] James Cook Univ, Coll Marine & Environm Sci, Cairns, Qld 4870, Australia
[6] James Cook Univ, Ctr Trop Environm Sustainabil Sci, Cairns, Qld 4870, Australia