Distribution of Norway Spruce Undergrowth in Primary Middle Taiga Spruce Forests
摘要
This paper discusses the distribution of Norway spruce (Picea abies (L.) H. Karst.) undergrowth in three types of communities: blueberry spruce forest, wood-sorrel–blueberry spruce forest with silver birch (Betula pendula Roth) and European aspen (Populus tremula L.), and wood-sorrel–blueberry spruce forest codominated by Scots pine (Pinus sylvestris L.). Woody detritus as a microsite is represented not only by lying logs and stumps that account for 37–72% of undergrowth, but also by adjacent spaces (10–27% of undergrowth) and visually indistinguishable deadwood contained in soil (1–11% of undergrowth). In the blueberry spruce forest, intercrown zone occupy one-third of the surveyed area, accounting for 46% of undergrowth; the pericrown zone occupies almost half of the surveyed area (41% of undergrowth). The occurrence of undergrowth is high in all canopy projection zones (84, 73, and 68%); undergrowth occurs in clusters on deadwood (37%) and in adjacent microsites (40%). The density of undergrowth depends on the cumulative effect of multiple factors: the highest values were recorded in intercrown zone on deadwood (1.0 individuals/m2) and medium values were in intercrown zone near deadwood (0.6 individuals/m2), in pericrown zone on deadwood (0.5 individuals/m2) and near deadwood (0.5 individuals/m2), and in undercrown zone without deadwood (0.5 individuals/m2); the lowest values were on deadwood “hidden” in soil (0.1–0.2 individuals/m2). In the wood-sorrel–blueberry spruce forest with aspen and birch, the canopy structure is more complex, and undergrowth is almost evenly distributed between the canopy projection zones whose ratio is 12 : 33 : 55%. The maximum undergrowth occurrence and density values were recorded in intercrown zone on deadwood (39% and 0.7 individuals/m2, respectively), while under the canopy, they are lower (18% and 0.3 individuals/m2, respectively). In the wood-sorrel–blueberry spruce forest codominated by Scots pine, undergrowth is distributed randomly: in all canopy projection zones, its occurrence (31, 33, and 30%) and density (0.6, 0.5, and 0.6 individuals/m2) values are equally low; the highest density values were recorded on deadwood under crowns (0.5 individuals/m2). Overall, the distribution of undergrowth in primary middle taiga spruce forests is determined by a combination of factors: forest growing conditions, stand composition, and diversity of microsites, including those formed by deadwood at different decay stages.