[1] Univ Hong Kong, Sch Biol Sci, Swire Inst Marine Sci, Hong Kong, Hong Kong, Peoples R China
[2] Univ Hong Kong, Dept Earth Sci, Hong Kong, Hong Kong, Peoples R China
[3] US Geol Survey, Reston, VA 20192 USA
[4] Univ Tromso, Dept Geol, N-9037 Tromso, Norway
[5] Smithsonian Inst, Natl Museum Nat Hist, Dept Paleobiol, Washington, DC 20013 USA