[1] Arizona State Univ, Sch Life Sci, Tempe, AZ 85287 USA
[2] Univ British Columbia, Dept Zool, Vancouver, BC V6T 1Z4, Canada
[3] Univ British Columbia, Biodivers Res Ctr, Vancouver, BC V6T 1Z4, Canada
[4] Conservat Int, Betty & Gordon Moore Ctr Sci & Oceans, Arlington, VA 22202 USA