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Foundations of biodiversity in managed Douglas-fir forests.

The controversy over old-growth Douglas-fir ( Pseudotsuga menziesii) forests and late-seral species has resulted in substantial information about biodiversity (Carey 1989, 1995, Carey and Johnson 1995, Carey et al. 1990, 1992, Forsmann et al. 1984). Major compilations of old-growth related research and its management implications include Gutierrez and Carey (1985), Ruggiero et al. (1991), Thomas et al. ( 1990, 1993), and Thomas and Raphael (1993). The need to restore and create late-seral ecosystems in the future was recognized (Carey 1985, Thomas et al. 1990), and Thomas et al. (1990) called for experiments to determine if northern spotted owl (Strix occidentalis) habitat could be created silviculturally. Our study addresses the management of second-growth forests to develop late-seral "composition, structure, function, ecologic products, and economic products" (Carey 1994a). Our Forest Ecosystem Study (PES) was designed to test whether late-seral forest attributes (specific targets include high architectural diversity, large diameter trees, high mycorrhizal and non-mycorrhizal fungal diversity, high vascular plant diversity, large prey base) can be developed from stem-exclusion phase ( Oliver and Larson 1990) forests through silviculture, and whether providing dens (cavities or nestboxes) can increase the density of northern flying squirrels ( Glaucomys sabrinus) in second-growth. A new silvicultural technique, variable-density thinning ( Carey and Miller 1991, Carey 1994b) was devised to create spatial heterogeneity in composition and structure that mimics conditions found in old-growth forests. We adopted a trophic pathway approach to the evaluation of our treatments. The basis for the forest ecosystem is the soil; forest soil foodwebs are characterized by a dominance of fungal activity over bacterial activity (Kromack et al. 1988). Mycorrhizal fungi assimilate large amounts of net primary productivity in Douglas-fir forests (Fogel and Hunt 1983) and enhance primary productivity by augmenting plant uptake of water and nutrients. Fungi are food for a diversity of consumers including arthropods, nematodes, and mammals (Perry et al.