[1] Univ Warwick, Sch Life Sci, Coventry CV4 7AL, W Midlands, England
[2] Coral Reef Conservat Project, Mombasa, Kenya
[3] Wildlife Conservat Soc, Marine Program, Bronx, NY 10430 USA
[4] James Cook Univ, ARC Ctr Excellence Coral Reef Studies, Townsville, Qld 4811, Australia