The natural age and stand structure of Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies (L.) Karst.), and birch (Betula pubescens Ehrh.) was studied in a virgin forest stand in northern Sweden. The stand has been unaffected by fire during the past 500 years. It is suggested that the variations in age structure and invasion pattern are the result of low-frequency climatic fluctuations influencing seed production, germination, and early survival of seedlings. The periods of climatic amelioration that occurred during the 1870s and during this century have been especially favourable to pine, resulting in a pine-dominated regeneration underneath a tree layer where spruce and birch are the most abundant species. This is contrary to generally accepted theories concerning postfire successional trends in this part of the boreal zone. It is concluded that small gap-phase replacement of trees by climatically induced regeneration and mortality events probably is the reason for the general weak correlation between age and size of trees. It is also concluded that this postfire succession will not lead to a total spruce dominance during a continuous succession under prevailing climate regimes.
Six old-growth, late postfire Scots pine (Pinussylvestris L.) dominated forest stands of the Vaccinium–Cladina type were selected along a latitudinal gradient in northern Sweden. In two of the stands, Scots pine seedlings that had naturally regenerated during the last 40 years were surveyed in relation to field- and bottom-layer vegetation. The most abundant forest floor species, viz. Cladina spp., occupied 41% of the ground cover and dominated the microhabitat of Scots pine seedlings <10 years old. The second most common species, Pleuroziumschreberi (Bird.) Mitt., which occupied 10 and 20% of the ground cover at the two sites, was, in contrast, only found to cover less than 3% of the microhabitat of the seedlings < 10 years old. With increasing age of seedlings (>10 years), however, the cover of Cladina spp. significantly decreased and the cover of P. schreberi and various ericaceous species slightly increased. Thus Scots pine seedlings initially establish in lichen-dominated vegetation but as they develop P. schreberi and ericaceous species gradually become more abundant in the ground vegetation. The relationships between natural establishment patterns and microhabitat were further empirically tested by sowing seeds of Scots pine in patches of Cladina spp. and patches of P. schreberi. Seeding was performed in six sites during 3 consecutive years. A significant higher number of established Scots pine seedlings were found in Cladina spp. vegetation than in P. schreberi dominated vegetation, and Scots pine seedlings in P. schreberi experienced a significantly higher mortality. Laboratory studies indicated a negative chemical influence by P. schreberi on Scots pine seed germination but not on radicle growth of pregerminated seeds. The negative influence of P. schreberi on Scots pine seed germination and seedling survival in the field experiments is interpreted as an effect of moisture factors, chemical interference, and barriers of nutrient availability. Age structures of naturally established Scots pine showed evidence of continuous regeneration over the last 40 years. The study emphasizes the importance of biotically mediated regeneration patterns in explaining spatial stand structure and temporal dynamics in northern boreal Scots pine forest ecosystems during prolonged absence of fire. Tree regeneration tends not to be associated with mesoscale characteristics such as gaps and tip-up mounds, but rather with microhabitat conditions. The results have implications for predicting the effect of climatic warming.
Scots pine (Pinussylvestris L.) were selectively seeded in 1990, 1991, 1992, and 1994 in bottom layer vegetation of Pleuroziumschreberi (Bird.) Mitt, in a clearcut created in 1989 and in an adjacent undisturbed Scots pine forest of Vaccinium type, in northern Sweden. Seedlings from the 1990 seeding established considerably better on the clearcut than in the forest. In contrast, seeding done in 1991, 1992, and 1994 gave no significant difference in seedling establishment between the two sites. Seedlings that established in the clearcut in 1990 grew significantly larger and had a higher content of macro- and micro-nutrients than those in the forest. For seedlings that established in 1992, biomass and nutrient content of seedlings in the clearcut did not differ from those in the forest. Seedlings established in the clearcut in 1990 increased their nutrient content in relation to seed supply, while seedlings established in the clearcut in 1992 and those in the forest had in general gained less or even lost nutrients (especially P) in relation to seed reserve. These results indicate that Pleuroziumschreberi-dominated microsites can constitute a favourable microhabitat for pine regeneration following stand disturbance, although the positive effect quickly diminishes with time after disturbance. The higher establishment and growth of emerging seedlings the 1st year after cutting compared with later established cohorts in the clearcut may be caused by changes in nutrient availability. We propose that environmental stress after cutting temporarily diminishes the effect of the nutrient barrier formed by Pleuroziumschreberi, ericoid mycorrhiza, and ericaceous dwarfshrubs. We conclude that successful natural pine regeneration in Pleuroziumschreberi-dominnted vegetation induced by clear-cutting can be severely time restricted. This is an important consideration when dealing with forestry methods that depend on natural regeneration.
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