[1] Univ Alberta, Dept Biol Sci, Edmonton, AB T6G 2E9, Canada
[2] Carleton Univ, Dept Geog & Environm Studies, Ottawa, ON K1S 5B6, Canada
[3] Univ Alberta, Dept Earth & Atmospher Sci, Edmonton, AB T6G 2E3, Canada
[4] Ohio State Univ, Sch Earth Sci, Marion, OH 43210 USA
[5] Univ Calif Berkeley, Lawrence Berkeley Natl Lab, Computat Res Div, Berkeley, CA 94720 USA