[1] Temple Univ, Dept Biol, Philadelphia, PA 19122 USA
[2] Senckenberg Meer, German Ctr Marine Biodivers Res DZMB, Bioctr Grindel, Martin Luther King Pl 3, D-20146 Hamburg, Germany
[3] Univ Hamburg, Ctr Nat Hist CeNak, Zool Museum, Martin Luther King Pl 3, D-20146 Hamburg, Germany
[4] Univ Iceland, Fac Life & Environm Sci, Askja Nat Sci Bldg,Sturlugata 7, IS-101 Reykjavik, Iceland
[5] Smithsonian Inst, Natl Museum Nat Hist, Labs Analyt Biol, Washington, DC 20560 USA