CAUSES AND ECOSYSTEM CONSEQUENCES OF MULTIPLE PATHWAYS OF PRIMARY SUCCESSION AT GLACIER BAY, ALASKA

The classic account of primary succession inferred from a 220-yr glacial retreat chronosequence at Glacier Bay National Park, Alaska was compared to reconstructions of stand development based on tree-ring records from 850 trees at 10 sites of different age. The three oldest sites (deglaciated prior to 1840) differ from all younger sites in the early recruitment of Sitka spruce (Picea sitchensis), the presence of western hemlock (Tsuga heterophylla), and the inferred importance of early shrub thickets. The nitrogen-fixing shrub Sitka alder (Alnus sinuata) has been an important and long-lived species only at sites deglaciated since 1840. Black cottonwood (Populus trichocarpa) has been an overstory dominant only at sites deglaciated since 1900. These single-species additions or replacements distinguish three pathways of vegetation compositional change which are segregated spatially and temporally. The communities of different age at Glacier Bay do not constitute a single chronosequence and should nut be used uncritically to infer long-term successional trends. Among-site differences in texture and lithology of soil parent, material cannot account for the multiple pathways. However, distance from each study site to the closest seed source of Sitka spruce at the time of deglaciation explains up to 58% of the among-site variance in early spruce recruitment. Multiple pathways of compositional change at Glacier Bay appear to be a function of landscape context, which, in conjunction with species life history trails (dispersal capability and generation time), affects seed rain to newly deglaciated surfaces and thereby alters the arrival sequence of species. Differences among the pathways probably include long-term differences in ecosystem function resulting from substantial accumulation of nitrogen at sites where nitrogen-fixing shrubs are important.