[1] Utah State Univ, Ecol Ctr, 5210 Old Main Hill, Logan, UT 84322 USA
[2] Utah State Univ, Dept Watershed Sci, 5210 Old Main Hill, Logan, UT 84322 USA
[3] Leibniz Inst Freshwater Ecol & Inland Fisheries, Dept Ecohydrol & Biogeochem, Muggelseedamm 301, D-12587 Berlin, Germany
[4] Univ Toronto, Dept Ecol & Evolutionary Biol, 25 Willcocks St, Toronto, ON M5S 3B2, Canada
[5] Royal Ontario Museum, Dept Nat Hist, 100 Queens Pk, Toronto, ON M5S 2C6, Canada