1. Department of Renewable Resources, University of Alberta, Edmonton, AB T6G 2R3, Canada
2. Department of Soil Science, University of Saskatchewan, Saskatoon, SK S7N 2H1, Canada
3. Present address, Department of Terrestrial Ecology, Netherlands Institute of Ecology (NIOO-KNAW), Droevendaalsesteeg 10, Wageningen, 6708 PB, the Netherlands