The influence of stand density on bilberry (Vaccinium myrtillus L.) cover depends on stand age, solar irradiation, and tree species composition

The ericaceous shrub bilberry (Vaccinium myrtillus L.) is a keystone species of the Eurasian boreal forest. The most optimal light condition for this plant is partial shading. Shade from the forest canopy depends on the stand density, a forest attribute that can be manipulated by forest managers. Most previous studies of the relationship between bilberry abundance and forest density have not explored the potentially modifying impacts of factors like stand age, tree species composition, and the solar irradiation at the site, as determined by location and topography. Using data from the Norwegian National Forest Inventory, we developed a generalized linear model applicable to estimate local bilberry cover across a wide range of environmental conditions in Norway. The explanatory terms in the final model were stand density (basal area per ha), solar irradiation, stand age, percentages of deciduous, pine, and spruce trees, summer (June-August) mean temperature and precipitation sum, mean temperature in January, site index, and soil category, in addition to the two-way interactions between stand density and the following: solar irradiation, stand age, percentage of deciduous trees, and percentage of Norway spruce (Picea abies). The final model explained ca. 21% of the total variation in bilberry cover. We conclude that a stand density of c. 30 m2 ha−1 in general will create favourable conditions for bilberry. If the forest is younger than 80 years old, or dominated by Norway spruce or deciduous trees, the optimal stand density is reduced to around 20 m2 ha−1. In a forest dominated by Scots pine (Pinus sylvestris), basal areas up to 40 m2 ha−1 would be beneficial to bilberry abundance. Our results demonstrate the importance of considering interactions between stand density and other stand and site characteristics.