Forest transformation from coniferous monocultures to mixed stands is being promoted worldwide, including the introduction of fast-growing broadleaved tree species within native stands. Here, we studied how enrichment of temperate European Scots pine (Pinus sylvestris) forest by North-American northern red oak Quercus rubra impacted macronutrient concentrations in two long-lived and dominant components of the forest understory: bilberry Vaccinium myrtillus and lingonberry V. vitis-idaea. Study sites were located in forest complexes (central Poland) which occupy continuously reforested lands (hereafter ancient forests) as well as post-agricultural lands (recent forests), all suitable for mesic pine forests. Samples of bilberry and lingonberry leaves, stems, and fruits were collected in pine stands and in adjacent Scots pine-red oak stands, in both ancient and recent forests. Concentrations of macronutrients (C, N, P, K, Ca, S, and Mg) in aboveground biomass components were analysed using standardized chemical procedures. The study revealed intra- and interspecific (bilberry vs. lingonberry) differences in concentrations of all nutrients in leaves, stems, and fruits, except for invariable C concentrations. Macronutrient accumulations in plants were decreased by land-use discontinuity and favoured by enrichment of tree stands by Q. rubra. The estimated macronutrient pools were much higher for V. myrtillus than V. vitis-idaea in all forest types studied. They were lower in forests enriched with Q. rubra, both ancient (up to 25.5% for bilberry and 99.9% for lingonberry) and recent (46.9% and 99.9%, respectively), as well as in recent pine forest (46.6% and 81.1%, respectively) than in ancient pine forest. Higher K and S pools (39.3% and 6.5%, respectively) noted for bilberry in an ancient forest with Q. rubra were exceptions. Despite more effective accumulations of elements at the species level, macronutrient pools of Vaccinium myrtillus and V. vitis-idaea decreased significantly in the presence of introduced Q. rubra due to negative impacts of this broadleaved tree on bilberry and lingonberry cover and biomass. Therefore, the limitation of alien Q. rubra planting in sites of mesic pine forest with the abundant occurrence of V. myrtillus and/or V. vitis-idaea is recommended.
Graphic abstract