The multi-scale approach to conserving forest biodiversity has been used in Sweden since the 1980s, a period defined by increased reserve area and conservation actions within production forests. However, two thousand forest-associated species remain on Sweden's red-list, and Sweden's 2020 goals for sustainable forests are not being met. We argue that ongoing changes in the production forest matrix require more consideration, and that multi-scale conservation must be adapted to, and integrated with, production forest development. To make this case, we summarize trends in habitat provision by Sweden's protected and production forests, and the variety of ways silviculture can affect biodiversity. We discuss how different forestry trajectories affect the type and extent of conservation approaches needed to secure biodiversity, and suggest leverage points for aiding the adoption of diversified silviculture. Sweden's long-term experience with multi-scale conservation and intensive forestry provides insights for other countries trying to conserve species within production landscapes.
Mires (bogs and fens) are nutrient-limited peatland ecosystems, the vegetation of which is especially sensitive to nitrogen deposition and climate change. The role of mires in the global carbon cycle, and the delivery of different ecosystem services can be considerably altered by changes in the vegetation, which has a strong impact on peat-formation and hydrology. Mire ecosystems are commonly open with limited canopy cover but both nitrogen deposition and increased temperatures may increase the woody vegetation component. It has been predicted that such an increase in tree cover and the associated effects on light and water regimes would cause a positive feed-back loop with respect to the ground vegetation. None of these effects, however, have so far been confirmed in large-scale spatiotemporal studies. Here we analyzed data pertaining to mire vegetation from the Swedish National Forest Inventory collected from permanent sample plots over a period of 20 yr along a latitudinal gradient covering 14°. We hypothesized that the changes would be larger in the southern parts as a result of higher nitrogen deposition and warmer climate. Our results showed an increase in woody vegetation with increases in most ericaceous dwarf-shrubs and in the basal area of trees. These changes were, in contrast to our expectations, evenly distributed over most of the latitudinal gradient. While nitrogen deposition is elevated in the south, the increase in temperatures during recent decades has been larger in the north. Hence, we suggest that different processes in the north and south have produced similar vegetation changes along the latitudinal gradient. There was, however, a sharp increase in compositional change at high deposition, indicating a threshold effect in the response. Instead of a positive feed-back loop caused by the tree layer, an increase in canopy cover reduced the changes in composition of the ground vegetation, whereas a decrease in canopy cover lead to larger changes. Increased natural disturbances of the tree layer due to, for example, pathogens or climate is a predicted outcome of climate change. Hence, these results may have important implications for predictions of long-term effects of increased temperature on peatland vegetation.
Plant growth in northern forest ecosystems is considered to be primarily nitrogen limited. Nitrogen deposition is predicted to change this towards co-limitation/limitation by other nutrients (e.g., phosphorus), although evidence of such stoichiometric effects is scarce. We utilized two forest fertilization experiments in southern Sweden to analyze single and combined effects of nitrogen and phosphorus on the productivity, composition, and diversity of the ground vegetation. Our results indicate that the productivity of forest ground vegetation in southern Sweden is co-limited by nitrogen and phosphorus. Additionally, the combined effect of nitrogen and phosphorus on the productivity was larger than when applied solely. No effects on species richness of any of these two nutrients were observed when applied separately, while applied in combination, they increased species richness and changed species composition, mainly by promoting more mesotrophic species. All these effects, however, occurred only for the vascular plants and not for bryophytes. The tree layer in a forest has a profound impact on the productivity and diversity of the ground vegetation by competing for both light and nutrients. This was confirmed in our study where a combination of nitrogen and high tree basal area reduced cover of the ground vegetation compared to all the other treatments where basal area was lower after stand thinning. During the past decades, nitrogen deposition may have further increased this competition from the trees for phosphorus and gradually reduced ground vegetation diversity. Phosphorus limitation induced by nitrogen deposition may, thus, contribute to ongoing changes in forest ground vegetation.
Northern forest ecosystems are exposed to a range of anthropogenic processes including global warming, atmospheric deposition, and changing land-use. The vegetation of northern forests is composed of species with several functional traits related to these processes, whose effects may be difficult to disentangle. Here, we combined analyses of spatio-temporal dynamics and functional traits of ground flora species, including morphological characteristics, responses to macro- and microclimate, soil conditions, and disturbance. Based on data from the Swedish National Forest Inventory, we compared changes in occurrence of a large number of ground flora species during a 20-year period (1994-2013) in boreal and temperate Sweden respectively. Our results show that a majority of the common ground flora species have changed their overall frequency. Comparisons of functional traits between increasing and declining species, and of trends in mean trait values of sample plots, indicate that current floristic changes are caused by combined effects of climate warming, nitrogen deposition and changing land-use. Changes and their relations with plant traits were generally larger in temperate southern Sweden. Nutrient-demanding species with mesotrophic morphology were favored by ongoing eutrophication due to nitrogen deposition in the temperate zone, while dwarf shrubs with low demands on nitrogen decreased in frequency. An increase of species with less northern and less eastern distribution limits was also restricted to temperate Sweden, and indicates effects of a moister and milder macroclimate. A trend toward dense plantation forests is mirrored by a decrease of light-demanding species in both vegetation zones, and a decrease of grassland species in the temperate zone. Although denser tree canopies may buffer effects of a warmer climate and of nitrogen deposition to some extent, traits related to these processes were weakly correlated in the group of species with changing frequency. Hence, our results indicate specific effects of these often confounded anthropogenic processes.