This work benefited and matured from discussions with the late D. Grant, who was invited to be a co-author on an earlier (1996, never submitted) version of this manuscript, but who disagreed with our findings which according to him lacked an adequate means of explaining the distribution of disjunct species of beetles and mosses in the highest weathering zones. B. Lawn completed most of the chemistry for the 1992 samples. The manuscript improved with discussions with I. Brookes and A. Murphy. Field support from C. Gallagher and S. Tubb, in 2002, and field and helicopter support from R.A. Klassen, Geological Survey of Canada, in 1992, were greatly appreciated. Grants ACOA-AIF-1005052 and NSF-0PP-9906280 to JCG and NSERC Discovery Grants to TB and JTG supported this research.

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