Seed regeneration of Scots pine in boreal forest stands dominated by lichen and feather moss

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.