Trade-offs in biodiversity and ecosystem services between edges and interiors in European forests.

Forest biodiversity and ecosystem services are hitherto predominantly quantified in forest interiors, well away from edges. However, these edges also represent a substantial proportion of the global forest cover. Here we quantified plant biodiversity and ecosystem service indicators in 225 plots along forest edge-to-interior transects across Europe. We found strong trade-offs: phylogenetic diversity (evolutionary measure of biodiversity), proportion of forest specialists, decomposition and heatwave buffering increased towards the interior, whereas species richness, nectar production potential, stemwood biomass and tree regeneration decreased. These trade-offs were mainly driven by edge-to-interior structural differences. As fragmentation continues, recognizing the role of forest edges is crucial for integrating biodiversity and ecosystem service considerations into sustainable forest management and policy.

Using warming tolerances to predict understory plant responses to climate change.

Climate change is pushing species towards and potentially beyond their critical thermal limits. The extent to which species can cope with temperatures exceeding their critical thermal limits is still uncertain. To better assess species' responses to warming, we compute the warming tolerance (ΔTniche ) as a thermal vulnerability index, using species' upper thermal limits (the temperature at the warm limit of their distribution range) minus the local habitat temperature actually experienced at a given location. This metric is useful to predict how much more warming species can tolerate before negative impacts are expected to occur. Here we set up a cross-continental transplant experiment involving five regions distributed along a latitudinal gradient across Europe (43° N-61° N). Transplant sites were located in dense and open forests stands, and at forest edges and in interiors. We estimated the warming tolerance for 12 understory plant species common in European temperate forests. During 3 years, we examined the effects of the warming tolerance of each species across all transplanted locations on local plant performance, in terms of survival, height, ground cover, flowering probabilities and flower number. We found that the warming tolerance (ΔTniche ) of the 12 studied understory species was significantly different across Europe and varied by up to 8°C. In general, ΔTniche were smaller (less positive) towards the forest edge and in open stands. Plant performance (growth and reproduction) increased with increasing ΔTniche across all 12 species. Our study demonstrated that ΔTniche of understory plant species varied with macroclimatic differences among regions across Europe, as well as in response to forest microclimates, albeit to a lesser extent. Our findings support the hypothesis that plant performance across species decreases in terms of growth and reproduction as local temperature conditions reach or exceed the warm limit of the focal species.

Soil seed bank responses to edge effects in temperate European forests.

Aim: The amount of forest edges is increasing globally due to forest fragmentation and land-use changes. However, edge effects on the soil seed bank of temperate forests are still poorly understood. Here, we assessed edge effects at contrasting spatial scales across Europe and quantified the extent to which edges can preserve the seeds of forest specialist plants. Location: Temperate European deciduous forests along a 2,300-km latitudinal gradient. Time period: 2018-2021. Major taxa studied: Vascular plants. Methods: Through a greenhouse germination experiment, we studied how edge effects alter the density, diversity, composition and functionality of forest soil seed banks in 90 plots along different latitudes, elevations and forest management types. We also assessed which environmental conditions drive the seed bank responses at the forest edge versus interior and looked at the relationship between the seed bank and the herb layer species richness. Results: Overall, 10,108 seedlings of 250 species emerged from the soil seed bank. Seed density and species richness of generalists (species not only associated with forests) were higher at edges compared to interiors, with a negative influence of C : N ratio and litter quality. Conversely, forest specialist species richness did not decline from the interior to the edge. Also, edges were compositionally, but not functionally, different from interiors. The correlation between the seed bank and the herb layer species richness was positive and affected by microclimate. Main conclusions: Our results underpin how edge effects shape species diversity and composition of soil seed banks in ancient forests, especially increasing the proportion of generalist species and thus potentially favouring a shift in community composition. However, the presence of many forest specialists suggests that soil seed banks still play a key role in understorey species persistence and could support the resilience of our fragmented forests.

Context matters: the landscape matrix determines the population genetic structure of temperate forest herbs across Europe.

Context: Plant populations in agricultural landscapes are mostly fragmented and their functional connectivity often depends on seed and pollen dispersal by animals. However, little is known about how the interactions of seed and pollen dispersers with the agricultural matrix translate into gene flow among plant populations. Objectives: We aimed to identify effects of the landscape structure on the genetic diversity within, and the genetic differentiation among, spatially isolated populations of three temperate forest herbs. We asked, whether different arable crops have different effects, and whether the orientation of linear landscape elements relative to the gene dispersal direction matters. Methods: We analysed the species' population genetic structures in seven agricultural landscapes across temperate Europe using microsatellite markers. These were modelled as a function of landscape composition and configuration, which we quantified in buffer zones around, and in rectangular landscape strips between, plant populations. Results: Landscape effects were diverse and often contrasting between species, reflecting their association with different pollen- or seed dispersal vectors. Differentiating crop types rather than lumping them together yielded higher proportions of explained variation. Some linear landscape elements had both a channelling and hampering effect on gene flow, depending on their orientation. Conclusions: Landscape structure is a more important determinant of the species' population genetic structure than habitat loss and fragmentation per se. Landscape planning with the aim to enhance the functional connectivity among spatially isolated plant populations should consider that even species of the same ecological guild might show distinct responses to the landscape structure. Supplementary Information: The online version contains supplementary material available at 10.1007/s10980-021-01376-7.

Forest understorey communities respond strongly to light in interaction with forest structure, but not to microclimate warming.

Forests harbour large spatiotemporal heterogeneity in canopy structure. This variation drives the microclimate and light availability at the forest floor. So far, we do not know how light availability and sub-canopy temperature interactively mediate the impact of macroclimate warming on understorey communities. We therefore assessed the functional response of understorey plant communities to warming and light addition in a full factorial experiment installed in temperate deciduous forests across Europe along natural microclimate, light and macroclimate gradients. Furthermore, we related these functional responses to the species' life-history syndromes and thermal niches. We found no significant community responses to the warming treatment. The light treatment, however, had a stronger impact on communities, mainly due to responses by fast-colonizing generalists and not by slow-colonizing forest specialists. The forest structure strongly mediated the response to light addition and also had a clear impact on functional traits and total plant cover. The effects of short-term experimental warming were small and suggest a time-lag in the response of understorey species to climate change. Canopy disturbance, for instance due to drought, pests or logging, has a strong and immediate impact and particularly favours generalists in the understorey in structurally complex forests.

Macronutrient balancing in free-ranging populations of moose.

At northern latitudes, large spatial and temporal variation in the nutritional composition of available foods poses challenges to wild herbivores trying to satisfy their nutrient requirements. Studies conducted in mostly captive settings have shown that animals from a variety of taxonomic groups deal with this challenge by adjusting the amounts and proportions of available food combinations to achieve a target nutrient balance. In this study, we used proportions-based nutritional geometry to analyze the nutritional composition of rumen samples collected in winter from 481 moose (Alces alces) in southern Sweden and examine whether free-ranging moose show comparable patterns of nutrient balancing. Our main hypothesis was that wild moose actively regulate their rumen nutrient composition to offset ecologically imposed variation in the nutritional composition of available foods. To test this, we assessed the macronutritional composition (protein, carbohydrates, and lipids) of rumen contents and commonly eaten foods, including supplementary feed, across populations with contrasting winter diets, spanning an area of approximately 10,000 km2. Our results suggest that moose balanced the macronutrient composition of their rumen, with the rumen contents having consistently similar proportional relationship between protein and nonstructural carbohydrates, despite differences in available (and eaten) foods. Furthermore, we found that rumen macronutrient balance was tightly related to ingested levels of dietary fiber (cellulose and hemicellulose), such that the greater the fiber content, the less protein was present in the rumen compared with nonstructural carbohydrates. Our results also suggest that moose benefit from access to a greater variety of trees, shrubs, herbs, and grasses, which provides them with a larger nutritional space to maneuver within. Our findings provide novel theoretical insights into a model species for ungulate nutritional ecology, while also generating data of direct relevance to wildlife and forest management, such as silvicultural or supplementary feeding practices.

Temperature effects on forest understorey plants in hedgerows: a combined warming and transplant experiment.

BACKGROUND AND AIMS: Hedgerows have been shown to improve forest connectivity, leading to an increased probability of species tracking the shifting bioclimatic envelopes. However, it is still unknown how species in hedgerows respond to temperature changes, and whether effects differ compared with those in nearby forests. We aimed to elucidate how ongoing changes in the climate system will affect the efficiency of hedgerows in supporting forest plant persistence and migration in agricultural landscapes. METHODS: Here we report results from the first warming experiment in hedgerows. We combined reciprocal transplantation of plants along an 860-km latitudinal transect with experimental warming to assess the effects of temperature on vegetative growth and reproduction of two common forest herbs (Anemone nemorosa and Geum urbanum) in hedgerows versus forests. KEY RESULTS: Both species grew taller and produced more biomass in forests than in hedgerows, most likely due to higher competition with ruderals and graminoids in hedgerows. Adult plant performance of both species generally benefitted from experimental warming, despite lower survival of A. nemorosa in heated plots. Transplantation affected the species differently: A. nemorosa plants grew taller, produced more biomass and showed higher survival when transplanted at their home site, indicating local adaptation, while individuals of G. urbanum showed greater height, biomass, reproductive output and survival when transplanted northwards, likely owing to the higher light availability associated with increasing photoperiod during the growing season. CONCLUSIONS: These findings demonstrate that some forest herbs can show phenotypic plasticity to warming temperatures, potentially increasing their ability to benefit from hedgerows as ecological corridors. Our study thus provides novel insights into the impacts of climate change on understorey plant community dynamics in hedgerows, and how rising temperature can influence the efficiency of these corridors to assist forest species' persistence and colonization within and beyond their current distribution range.

Half a century of multiple anthropogenic stressors has altered northern forest understory plant communities.

Boreal forests form the largest and least disturbed forest biome in the northern hemisphere. However, anthropogenic pressure from intensified forest management, eutrophication, and climate change may alter the ecosystem functions of understory vegetation and services boreal forests provide. Swedish forests span long gradients of climate, nitrogen deposition, and management intensity. This makes them ideal to study how the species composition and functions of other, more pristine, boreal forests might change under increased anthropogenic pressure. Moreover, the National Forest Inventory (NFI) has collected systematic data on Swedish forest vegetation since the mid-20th century. We use this data to quantify changes in vegetation types between two periods, 1953-1962 and 2003-2012. The results show changes in forest understory vegetation since the 1950s at scales not previously documented in the boreal biome. The spatial extent of most vegetation types changed significantly. Shade-adapted and nutrient-demanding species (those with high specific leaf area) have become more common at the expense of light-demanding and nutrient-conservative (low specific leaf area) species. The cover of ericaceous dwarf shrubs decreased dramatically. These effects were strongest where anthropogenic impacts were greatest, suggesting links to drivers such as nitrogen deposition and land-use change. These changes may impact ecosystem functions and services via effects on higher trophic levels and faster plant litter decomposition in the expanding vegetation types. This, in turn, may influence nutrient dynamics, and consequently ecosystem productivity and carbon sequestration.

Understanding context dependency in the response of forest understorey plant communities to nitrogen deposition.

Understorey communities can dominate forest plant diversity and strongly affect forest ecosystem structure and function. Understoreys often respond sensitively but inconsistently to drivers of ecological change, including nitrogen (N) deposition. Nitrogen deposition effects, reflected in the concept of critical loads, vary greatly not only among species and guilds, but also among forest types. Here, we characterize such context dependency as driven by differences in the amounts and forms of deposited N, cumulative deposition, the filtering of N by overstoreys, and available plant species pools. Nitrogen effects on understorey trajectories can also vary due to differences in surrounding landscape conditions; ambient browsing pressure; soils and geology; other environmental factors controlling plant growth; and, historical and current disturbance/management regimes. The number of these factors and their potentially complex interactions complicate our efforts to make simple predictions about how N deposition affects forest understoreys. We review the literature to examine evidence for context dependency in N deposition effects on forest understoreys. We also use data from 1814 European temperate forest plots to test the ability of multi-level models to characterize context-dependent understorey responses across sites that differ in levels of N deposition, community composition, local conditions and management history. This analysis demonstrated that historical management, and plot location on light and pH-fertility gradients, significantly affect how understorey communities respond to N deposition. We conclude that species' and communities' responses to N deposition, and thus the determination of critical loads, vary greatly depending on environmental contexts. This complicates our efforts to predict how N deposition will affect forest understoreys and thus how best to conserve and restore understorey biodiversity. To reduce uncertainty and incorporate context dependency in critical load setting, we should assemble data on underlying environmental conditions, conduct globally distributed field experiments, and analyse a wider range of habitat types.

Phosphorus and nitrogen co-limitation of forest ground vegetation under elevated anthropogenic nitrogen deposition.

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.

Peatland plant communities under global change: negative feedback loops counteract shifts in species composition.

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.

Trait variations of ground flora species disentangle the effects of global change and altered land-use in Swedish forests during 20 years.

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.

The Nutritional Balancing Act of a Large Herbivore: An Experiment with Captive Moose (Alces alces L).

The nutrient balancing hypothesis proposes that, when sufficient food is available, the primary goal of animal diet selection is to obtain a nutritionally balanced diet. This hypothesis can be tested using the Geometric Framework for nutrition (GF). The GF enables researchers to study patterns of nutrient intake (e.g. macronutrients; protein, carbohydrates, fat), interactions between the different nutrients, and how an animal resolves the potential conflict between over-eating one or more nutrients and under-eating others during periods of dietary imbalance. Using the moose (Alces alces L.), a model species in the development of herbivore foraging theory, we conducted a feeding experiment guided by the GF, combining continuous observations of six captive moose with analysis of the macronutritional composition of foods. We identified the moose's self-selected macronutrient target by allowing them to compose a diet by mixing two nutritionally complementary pellet types plus limited access to Salix browse. Such periods of free choice were intermixed with periods when they were restricted to one of the two pellet types plus Salix browse. Our observations of food intake by moose given free choice lend support to the nutrient balancing hypothesis, as the moose combined the foods in specific proportions that provided a particular ratio and amount of macronutrients. When restricted to either of two diets comprising a single pellet type, the moose i) maintained a relatively stable intake of non-protein energy while allowing protein intakes to vary with food composition, and ii) increased their intake of the food item that most closely resembled the self-selected macronutrient intake from the free choice periods, namely Salix browse. We place our results in the context of the nutritional strategy of the moose, ruminant physiology and the categorization of food quality.

Interactions with successional stage and nutrient status determines the life-form-specific effects of increased soil temperature on boreal forest floor vegetation.

The boreal forest is one of the largest terrestrial biomes and plays a key role for the global carbon balance and climate. The forest floor vegetation has a strong influence on the carbon and nitrogen cycles of the forests and is sensitive to changes in temperature conditions and nutrient availability. Additionally, the effects of climate warming on forest floor vegetation have been suggested to be moderated by the tree layer. Data on the effects of soil warming on forest floor vegetation from the boreal forest are, however, very scarce. We studied the effects on the forest floor vegetation in a long-term (18 years) soil warming and fertilization experiment in a Norway spruce stand in northern Sweden. During the first 9 years, warming favored early successional species such as grasses and forbs at the expense of dwarf shrubs and bryophytes in unfertilized stands, while the effects were smaller after fertilization. Hence, warming led to significant changes in species composition and an increase in species richness in the open canopy nutrient limited forest. After another 9 years of warming and increasing tree canopy closure, most of the initial effects had ceased, indicating an interaction between forest succession and warming. The only remaining effect of warming was on the abundance of bryophytes, which contrary to the initial phase was strongly favored by warming. We propose that the suggested moderating effects of the tree layer are specific to plant life-form and conclude that the successional phase of the forest may have a considerable impact on the effects of climate change on forest floor vegetation and its feedback effects on the carbon and nitrogen cycles, and thus on the climate.