[1] Univ Washington, Dept Biol, Seattle, WA 98195 USA
[2] Burke Museum Nat Hist & Culture, Seattle, WA 98195 USA
[3] Univ Sheffield, Sch Biosci, Sheffield S10 2TN, England
[4] New York Bot Garden, Inst Systemat Bot, Bronx, NY 10458 USA
[5] Univ Washington, Dept Earth & Space Sci, Seattle, WA 98195 USA