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James Cook Univ N Queensland, Cooperat Res Ctr Trop Rainforest Ecol, Sch Trop Biol, Townsville, Qld 4811, Australia James Cook Univ N Queensland, Cooperat Res Ctr Trop Rainforest Ecol, Sch Trop Biol, Townsville, Qld 4811, Australia