[1] Univ Washington, Sch Aquat & Fishery Sci, Seattle, WA 98195 USA
[2] Burke Museum Nat Hist & Culture, Seattle, WA 98105 USA
[3] Smithsonian Inst, Natl Museum Nat Hist, Dept Vertebrate Zool, Washington, DC 20560 USA
[4] Smithsonian Trop Res Inst, Balboa, Panama
[5] Calif Acad Sci, Dept Ichthyol, San Francisco, CA 94118 USA