Dwarf Shrub Vaccinium Myrtillus Research Articles

Disturbance interval modulates the starting point for vegetation succession.

Increased frequency and new types of disturbances caused by global change calls for deepened insights into possible alterations of successional pathways. Despite current interest in disturbance interactions there is a striking lack of studies focusing on the implication of decreasing times between disturbances. We surveyed forest-floor vegetation (vascular plants and bryophytes) in a Pinus sylvestris-dominated, even-aged production forest landscape, unique because of the presence of stands under a precisely dated disturbance interval gradient, ranging from 0 to 123yr between clearcutting and a subsequent megafire. Despite a dominance of early-successional species in all burned stands 5 yr after fire, progression of succession was linked to time since the preceding clearcutting disturbance. This was most clearly seen in increased frequency with time since clearcutting of the dominant, late-successional dwarf shrub Vaccinium myrtillus, with surviving rhizomes as an important mechanism for postfire recovery. Our results demonstrate the role of legacy species as significant drivers of succession. We conclude that the starting point for succession is modulated by disturbance interval, so that shortened intervals risk reducing development towards late-successional stages. We suggest that a decrease in long successional sequences caused by more frequent disturbances may represent a general pattern, relevant also for other forest types and ecosystems.

Nitrogen deposition is positively correlated to foliar nitrogen content in Vaccinium myrtillus and other understory species in temperate forests on acidic soil

Excessive atmospheric nitrogen deposition is known to alter nutrient cycles and species composition in temperate forest ecosystems. Foliar N concentrations reflect the nutrient status of plants and can be used to assess the impact of atmospheric N deposition on ecosystems. Our objective was to test whether the foliar N content reflects increasing N deposition in the understory vegetation of temperate forests on acidic soil. Therefore, we analyzed the foliar N content of seven plant taxa representing different functional groups of the understory vegetation and correlated them to modeled deposition datasets. Linear mixed models revealed no relationship of N deposition with foliar N concentrations across all species examined but decreasing foliar concentrations with increasing soil C/N ratio. Nevertheless, we found positive correlations of foliar N concentrations with increasing deposition in conifer saplings (Picea abies and Abies alba), the dwarf shrub Vaccinium myrtillus and the graminoid Luzula luzuloides. Such an increase was, however, not found in saplings of broad-leaved trees (Fagus sylvatica and Quercus petraea/robur) and the moss Polytrichum formosum. This suggests marked species-specific differences in nitrogen uptake and allocation. Foliar N concentrations were more strongly correlated to NOy than NHx and to wet than dry deposition. Since foliar N concentrations were correlated to deposition while the species’ cover was not, they may provide a valuable indicator of N deposition.

Разнообразие лиственничных лесов и редколесий западного макросклона Приполярного и Северного Урала

Разнообразие лиственничных лесов и редколесий западного макросклона Приполярного и Северного Урала

Xylem anatomical and growth responses of the dwarf shrub Vaccinium myrtillus to experimental CO2 enrichment and soil warming at treeline

Plant growth responses to environmental changes may be linked to xylem anatomical adjustments. The study of such links is essential for improving our understanding of plant functioning under global change. We investigated the xylem anatomy and above-ground growth of the dwarf shrub Vaccinium myrtillus in the understorey of Larix decidua and Pinus uncinata at the Swiss treeline after 9 years of free-air CO2 enrichment (+200 ppm) and 6 years of soil warming (+4 °C). We aimed to determine the responses of xylem anatomical traits and growth to these treatments, and to analyse xylem anatomy–growth relationships. We quantified anatomical characteristics of vessels and ray parenchyma and measured xylem ring width (RW), above-ground biomass and shoot elongation as growth parameters. Our results showed strong positive correlations between theoretical hydraulic conductivity (Kh) and shoot increment length or total biomass across all treatments. However, while soil warming stimulated shoot elongation and RW, it reduced vessel size (Dh) by 14%. Elevated CO2 had smaller effects than soil warming: it increased Dh (5%) in the last experimental years and only influenced growth by increasing basal stem size. The abundance of ray parenchyma, representing storage capacity, did not change under any treatment. Our results demonstrate a link between growth and stem Kh in V. myrtillus, but its growth responses to warming were not explained by the observed xylem anatomical changes. Smaller Dh under warming may increase resistance to freezing events frequently occurring at treeline and suggests that hydraulic efficiency is not limiting for V. myrtillus growing on moist soils at treeline. Our findings suggest that future higher atmospheric CO2 concentrations will have smaller effects on V. myrtillus growth and functioning than rising temperatures at high elevations; further, growth stimulation of this species under future warmer conditions may not be synchronized with xylem adjustments favouring hydraulic efficiency.

Forest management regulates temporal change in the cover of boreal plant species

We analysed the driving forces of temporal change in the cover of boreal forest plant species in Finland over 21years (1985–2006). Our study focused on the effects of forest cuttings of different intensity, i.e. regeneration cuttings (RC) and intermediate cuttings (IC), on 11 common understory species (including vascular plants, mosses and lichens) and their recovery after cuttings. General linear models (LM) were applied in modelling the change in the cover of plant species. The plant species displayed contrasting responses to RC and IC in accordance with their light demand, which also accounted for their recovery rate after cutting disturbance. After RC of old forest, shade and semi-shade tolerant species (herb Oxalis acetosella, moss Hylocomium splendens, dwarf shrub Vaccinium myrtillus) decreased strongly in cover (>70%) and recovered relatively slowly, but increased (40–100%) after IC. Species adapted to semi-light conditions (mosses Dicranum polysetum and Pleurozium schreberi, dwarf shrub Vaccinium vitis-idaea) decreased rather strongly (20–60%) after RC, but recovered relatively fast, reaching their old-forest cover level at a stand age of ca. 30years. These species also increased after IC. In contrast, light-demanding species (dwarf shrubs Empetrum nigrum and Calluna vulgaris, grass Deschampsia flexuosa) increased after RC and decreased after IC. Cladina lichens decreased across the country. In general, early successional light-demanding species increased and late successional shade-tolerant species decreased after RC. IC favoured all but the most light-demanding species. Species-specific responses to cuttings showed considerable variation within plant functional groups (e.g. dwarf shrubs, herbs, mosses). Cuttings determined the principal change in vegetation, while only slight signals of the effect of climate warming were found in few species. The legacy of forest management deserves special attention when e.g. tracking the effect of global change on vegetation.

Vaccinium myrtillus stands show similar structure and functioning under different scenarios of coexistence at the Pyrenean treeline

Plant–plant interactions are key drivers of treeline dynamics. At the Pyrenean treeline, the dwarf shrub Vaccinium myrtillus grows in pure stands or in mixed stands with Vaccinium uliginosum or Rhododendron ferrugineum. They form sparse shrub patches that colonize subalpine grasslands, having dramatic impacts on their structure and functioning. We investigated the role of the two co-occurring shrubs as possible modulators of the structure and performance of V. myrtillus in the Central Pyrenees. We analysed biomass, growth, functional parameters, age distribution, N and C concentrations and isotope compositions (δ15N and δ 13C) of V. myrtillus ramets in pure stands, and stands mixed with V. uliginosum or R. ferrugineum. Volume, above-ground biomass and age of the ramets did not differ between stand types. We found lower δ13C values, indicative of lower water-use efficiency (WUE), in leaves and shoots of V. myrtillus in stands with R. ferrugineum than in pure stands. The N content and δ15N of V. myrtillus leaves and shoots in pure stands were higher than in mixed stands, pointing to a competition for N in mixed stands. Our results indicate that V. myrtillus competes for nutrients with its neighbours, but neither this competition nor the lower WUE affect its above-ground performance. Therefore, the interaction with co-occurring shrubs does not have a major effect on V. myrtillus structure and functioning at treeline and, consequently, should not be considered as a key driver on the dynamics of this species in the encroachment of subalpine grasslands.

Canopy cover and understory composition determine abundance of Vaccinium myrtillus L., a key plant for capercaillie (Tetrao urogallus), in subalpine forests in the Pyrenees

Background: The dwarf shrub Vaccinium myrtillus – with high cover, height, and fruit production – benefits capercaillie (Tetrao urogallus).Aims: Our aim was to quantify landscape (e.g. elevation, geographic location, and precipitation), site (e.g. overstorey cover and stoniness) and very fine scale factors (e.g. spatial associations in the understorey) that affect cover, height, and fruit production of V. myrtillus in subalpine forests in thePyrenees, with understorey usually dominated by Rhododendron ferrugineum.Methods: We sampled 155 plots (0.5 m × 5 m) in six sites. For each plot, in the understorey layer, we assessed species cover, height for R. ferrugineum and V. myrtillus, number of total fruits in V. myrtillus, and spatial associations among V. myrtillus and the remaining cover types.Results: Overstorey cover negatively influenced V. myrtillus cover, its height, and particularly, the number of fruits, which was also negatively influenced by R. ferrugineum cover. Associations between R. ferrugineum and V. myrtillus were site dependent, while V. myrtillus showed mostly positive associations with grasses and mosses.Conclusions: Reducing overstorey and R. ferrugineum cover has the strongest positive effect on increasing V. myrtillus fruit production, but with additional positive effects on V. myrtillus cover and height. Increases in grass and moss coverage could favour V. myrtillus.

Vegetation shift from deciduous to evergreen dwarf shrubs in response to selective herbivory offsets carbon losses: evidence from 19years of warming and simulated herbivory in the subarctic tundra.

Selective herbivory of palatable plant species provides a competitive advantage for unpalatable plant species, which often have slow growth rates and produce slowly decomposable litter. We hypothesized that through a shift in the vegetation community from palatable, deciduous dwarf shrubs to unpalatable, evergreen dwarf shrubs, selective herbivory may counteract the increased shrub abundance that is otherwise found in tundra ecosystems, in turn interacting with the responses of ecosystem carbon (C) stocks and CO2 balance to climatic warming. We tested this hypothesis in a 19-year field experiment with factorial treatments of warming and simulated herbivory on the dominant deciduous dwarf shrub Vacciniummyrtillus. Warming was associated with a significantly increased vegetation abundance, with the strongest effect on deciduous dwarf shrubs, resulting in greater rates of both gross ecosystem production (GEP) and ecosystem respiration (ER) as well as increased C stocks. Simulated herbivory increased the abundance of evergreen dwarf shrubs, most importantly Empetrum nigrum ssp. hermaphroditum, which led to a recent shift in the dominant vegetation from deciduous to evergreen dwarf shrubs. Simulated herbivory caused no effect on GEP and ER or the total ecosystem C stocks, indicating that the vegetation shift counteracted the herbivore-induced C loss from the system. A larger proportion of the total ecosystem C stock was found aboveground, rather than belowground, in plots treated with simulated herbivory. We conclude that by providing a competitive advantage to unpalatable plant species with slow growth rates and long life spans, selective herbivory may promote aboveground C stocks in a warming tundra ecosystem and, through this mechanism, counteract C losses that result from plant biomass consumption.

Dynamics of understory vegetation after restoration of natural characteristics in the boreal forests in Finland

Restoration of boreal coniferous forests is used in Northern Europe to increase the natural variability of forest structure and to add coarse woody debris for red-listed dead-wood dependent species, mainly insects and fungi. In four protection areas of eastern Finland, we evaluated how three commonly used restoration methods (felling with chainsaw, storm simulation, and felling with burning) with two felling levels (20% and 40% of initial living stand volume) affect the vegetation dynamics. The aim was to assess whether the type and intensity of restoration lead to different stages in the plant succession which would reflect the effectiveness of the restoration methods in terms of simulating natural disturbances. The restoration treatments were conducted in 2006 on 43 experimental stands and 15 were set as controls. Understory vegetation cover was surveyed from 372 permanent 1 m2 plots in 2005, 2007 and 2011. The data were analyzed with mixed effect models and multivariate methods. Burning most efficiently initiated natural vegetation succession. Despite the initial decline of nearly all plant functional types, pioneering species such as Betula spp. and Populus tremula, forbs, graminoids and pioneer mosses regenerated quickly and exceeded their pre-disturbance covers in 2011. Dwarf shrubs Vaccinium myrtillus, V. vitis-idaea and Calluna vulgaris recovered nearly to their initial levels, whereas Empetrum nigrum, forest mosses and lichens did not show any recovery in five years. Storm simulation increased species richness on the stand scale and promoted the regeneration of Pinus sylvestris in soil exposed by tree uprooting. Tree felling slightly decreased the cover of lichens on the lower felling level treatment. We generated a species response table illustrating the responses of dominant species to restoration methods in the course of the seven-year study. We conclude that the dominant restoration method, tree felling, could increasingly be replaced by burning and storm simulation which have faster and greater positive impacts on the forest ecosystem.

Changes in the abundance of keystone forest floor species in response to changes of forest structure

AbstractQuestionDoes the abundance of keystone forest floor species change in response to changes in the forest structure?LocationSwedenMethodsWe used data from theSwedishNationalForest Inventory to investigate changes in the abundance of three common species, as well as the total abundance of all understorey vascular plants (the field layer) in forests in the boreal and temperate parts ofSweden.GLMs andGAMs were used to relate species abundance and temporal changes in abundances to forest structure and forest structural change.ResultsProductivity, measured as the site index, was the most important determinant of individual species' abundance. The volume ofPicea abies, the density of tree stems and forest age were among the most important forest structural variables. We found that the dwarf shrubVaccinium myrtillus, the narrow‐leaved grasses (mainlyAvenella flexuosa) and the total field layer cover decreased in borealSweden from 1994 through 2010 and that these changes coincided with an increase in forest density and with a reduction in forest age.ConclusionsChanges inSwedish forests to higher tree layer density and younger age appear to contribute significantly to current changes in forest floor vegetation. The use of more intensive thinning practices to reduce the total density of the forest and to increase the proportion of broad‐leaved tree species andPinus sylvestriswould favour the forest floor species in this study. Moreover, increasing forest age (i.e. the length of rotation periods) might favourV. myrtillusin particular, for which the time since disturbance is important for the recovery of pre‐disturbance abundance. However, increased thinning intensity and forest age will reduce the potential for wood production, implying a trade‐off between production of wood and maintenance of well‐developed forest floor vegetation.

Evidence of enhanced freezing damage in treeline plants during six years of CO2 enrichment and soil warming

Climate change and elevated atmospheric CO2 levels could increase the vulnerability of plants to freezing. We analyzed tissue damage resulting from naturally occurring freezing events in plants from a long–term in situ CO2 enrichment (+ 200 ppm, 2001–2009) and soil warming (+ 4°C since 2007) experiment at treeline in the Swiss Alps (Stillberg, Davos). Summer freezing events caused damage in several abundant subalpine and alpine plant species in four out of six years between 2005 and 2010. Most freezing damage occurred when temperatures dropped below –1.5°C two to three weeks after snow melt. The tree Larix decidua and the dwarf shrubs Vaccinium myrtillus and Empetrum hermaphroditum showed more freezing damage under experimentally elevated CO2 and/or temperatures than under control conditions. Soil warming induced a 50% die‐back of E. hermaphroditum during a single freezing event due to melting of the protective snow cover. Although we could not identify a clear mechanism, we relate greater freezing susceptibility to a combination of advanced plant phenology in spring and changes in plant physiology. The climate record since 1975 at the treeline site indicated a summer warming by 0.58°C/decade and a 3.5 days/decade earlier snow melt, but no significant decrease in freezing events during the vegetation period. Therefore, in a warmer climate with higher CO2 levels but constant likelihood of extreme weather events, subalpine and alpine plants may be more susceptible to freezing events, which may partially offset expected enhanced growth with global change. Hence, freezing damage should be considered when predicting changes in growth of alpine plants or changes in community composition under future atmospheric and climate conditions.

Absence of snow cover reduces understory plant cover and alters plant community composition in boreal forests

Snow regimes affect biogeochemistry of boreal ecosystems and are altered by climate change. The effects on plant communities, however, are largely unexplored despite their influence on relevant processes. Here, the impact of snow cover on understory community composition and below-ground production in a boreal Picea abies forest was investigated using a long-term (8-year) snow cover manipulation experiment consisting of the treatments: snow removal, increased insulation (styrofoam pellets), and control. The snow removal treatment caused longer (118 vs. 57 days) and deeper soil frost (mean minimum temperature -5.5 vs. -2.2°C) at 10 cm soil depth in comparison to control. Understory species composition was strongly altered by the snow cover manipulations; vegetation cover declined by more than 50% in the snow removal treatment. In particular, the dominant dwarf shrub Vaccinium myrtillus (-82%) and the most abundant mosses Pleurozium schreberi (-74%) and Dicranum scoparium (-60%) declined strongly. The C:N ratio in V. myrtillus leaves and plant available N in the soil indicated no altered nitrogen nutrition. Fine-root biomass in summer, however, was negatively affected by the reduced snow cover (-50%). Observed effects are attributed to direct frost damage of roots and/ or shoots. Besides the obvious relevance of winter processes on plant ecology and distribution, we propose that shifts in the vegetation caused by frost damage may be an important driver of the reported alterations in biogeochemistry in response to altered snow cover. Understory plant performance clearly needs to be considered in the biogeochemistry of boreal systems in the face of climate change.

Drought tolerance of juvenile and mature leaves of a deciduous dwarf shrub Vaccinium myrtillus L. in a boreal environment

The difference between drought tolerance of juvenile and mature leaves of the winter-deciduous dwarf shrub bilberry (Vaccinium myrtillus L.) from a northern boreal environment was investigated. It was hypothesised that mature leaves are more drought sensitive than juvenile leaves. Bilberry plants were allowed to dry out by excluding irrigation when leaves were at juvenile and mature stages. Tissue water content decreased at both phenological stages, but the response was more pronounced in the mature leaves. Anthocyanin concentrations increased as the tissue water content decreased, and again this occurred to a greater extent in the mature leaves. Chlorophyll concentrations decreased only marginally at the juvenile stage, while the decrease was significant in the mature leaves. Chlorophyll degradation was enhanced by drought stress. Soluble proteins decreased and protein oxidation increased in the mature leaves, and degradation of oxidised proteins increased in the drought-stressed plants. The results suggest that leaves of bilberry are more sensitive to drought stress at the mature stage, and that drought stress accelerates senescence at the mature stage. The significance of the results is that dry periods during the juvenility of leaves are not as detrimental as they may be later in summer. In addition, the strategy of a winter-deciduous plant is obviously to protect its perennial parts from severe drought by accelerated leaf senescence at the mature stage. Therefore, the deciduous life form may provide an excellent adaptation against drought also in northern ecosystems. The role of anthocyanins in photoprotection under drought stress is also discussed.

Complex Biotic Interactions Drive Long-Term Vegetation Change in a Nitrogen Enriched Boreal Forest

The effects of experimental nitrogen (N) additions (0, 12.5, and 50 kg N ha−1 y−1) on long-term (12 years) understorey vegetation dynamics were examined in a boreal forest. The results showed that two types of natural enemies of the dominant dwarf-shrub Vaccinium myrtillus (pathogenic fungus of the species Valdensia heterodoxa and herbivorous larvae of the genus Operophtera) influenced the vegetation dynamics. The pathogenic fungus, causing premature leaf-shed of V. myrtillus, showed a strong positive N response during the initial 5-year period. For the larvae, a relatively modest N response was overshadowed by an almost 40-fold population increase during an outbreak event that followed the initial 5-year period. This outbreak occurred irrespective of N addition, resulting in V. myrtillus decline and depriving the pathogenic fungus of its substrate. Hence our study demonstrates that vegetation dynamics in this relatively species poor and seemingly simple ecosystem are driven by complex biotic interactions. Further, we show that an important component of these interactions is the temporal alternation of the two natural enemies and, resultant regulation of the dominant plant’s abundance. Finally, we emphasize that long-term data are essential to capture the complexity of this type of biotic interactions. In our case, a short-term study may have resulted in markedly different conclusions regarding effects of N enrichment and the role of biotic interactions for forest vegetation dynamics.

Responses of vegetation and soil microbial communities to warming and simulated herbivory in a subarctic heath

1. Climate warming increases the cover of deciduous shrubs in arctic ecosystems and herbivory is also known to have a strong influence on the biomass and composition of vegetation. However, research combining herbivory with warming is largely lacking. Our study describes how warming and simulated herbivory affect vegetation, soil nutrient concentrations and soil microbial communities after 10-13 years of exposure. 2. We established a factorial warming and herbivory-simulation experiment at a subarctic tundra heath in Kilpisjarvi, Finland, in 1994. Warming was carried out using the open-top chamber setup of the International Tundra Experiment (ITEX). Wounding of the dominant deciduous dwarf shrub Vaccinium myrtillus L. to simulate herbivory was carried out annually. We measured vegetation cover in 2003 and 2007, soil nutrient concentrations in 2003 and 2006, soil microbial respiration in 2003, and composition and function of soil microbial communities in 2006. 3. Warming increased the cover of V. myrtillus, whereas other plant groups did not show any response. Simulated herbivory of V. myrtillus cancelled out the impact of warming on the species cover, and increased the cover of other dwarf shrubs. 4. The concentrations of NH4+-N, and microbial biomass C and N in the soil were significantly reduced by warming after 10 treatment years but not after 13 treatment years. The reduction in NH4+-N by warming was significant only without simultaneous herbivory treatment, which indicates that simulated herbivory reduced N uptake by vegetation. 5. Soil microbial community composition, based on phospholipid fatty acid (PLFA) analysis, was slightly altered by warming. The activity of cultivable bacterial and fungal communities was significantly increased by warming and the substrate utilization patterns were influenced by warming and herbivory. 6. Synthesis. Our results show that warming increases the cover of V. myrtillus, which seems to enhance the nutrient sink strength of vegetation in the studied ecosystem. However, herbivory partially negates the effect of warming on plant N uptake and interacts with the effect of warming on microbial N immobilization. Our study demonstrates that effects of warming on soil microorganisms are likely to differ in the presence and absence of herbivores. (Less)

Commercial forest fertilization causes long-term residual effects in ground vegetation of boreal forests

Due to an increasing demand of forestry products, nitrogen (N) fertilization of coniferous forests may in the future become a standard silvicultural practice in parts of the boreal region that is not too remotely situated. Beside positive effects on productivity, forest fertilization may induce changes in species composition of the ground vegetation. More information on the magnitude of such side effects is needed to evaluate whether commercial forest fertilization is in accordance with principles of sustainable forestry and preservation of biodiversity. We examined the effects of forest fertilization on ground vegetation in boreal forest by comparing species composition in unfertilized stands and stands fertilized twice (>20 years since the last fertilization) with 150 kg N ha −1 per event. By using stands that had been clear-felled and replanted 8–11 years ago, we addressed whether effects of fertilization persists over more than one forest generation. The ground vegetation in fertilized stands was denser, showed decreased species evenness, lower biodiversity, and higher Ellenberg N values. The abundance of dwarf shrubs was reduced by more than 40%, while grasses and some nitrophilous herbs increased by >100%. For bottom layer species, fertilization favoured litter-dwelling bryophytes (e.g. Brachythecium spp.), while Hylocomium splendens and ground living lichens were disfavoured. For the dwarf shrub Vaccinium myrtillus, fertilization reduced the density of current annual shoots by >60% and berry production by >70%. Fertilization had, however, no effect on berries produced per annual shoot. Leaf tissue N concentrations were higher in V. myrtillus, and Deschampsia flexuosa growing in previously fertilized stands, indicating N enrichment although more than 20 years had past since the last fertilization event. Because residual effects of commercial forest fertilization on forest ground vegetation appear to be substantial and extend over more than one forest generation, we question whether forest fertilization, as performed today in the Nordic countries, can be considered a long-term sustainable silvicultural practice that is in accordance with principles of preservation of biodiversity.

Effects of recreational use and fragmentation on the understorey vegetation and soil microbial communities of urban forests in southern Finland

The impacts of fragmentation and recreational use on the hemiboreal urban forest understorey vegetation and the microbial community of the humus layer (the phospholipid fatty acid (PLFA) pattern, microbial biomass and microbial activity, measured as basal respiration) were examined in the greater Helsinki area, southern Finland. Trampling tolerance of 1) herb-rich OMT, 2) mesic MT, and 3) sub-xeric VT forests (in decreasing order of fertility) was studied by comparing relative understorey vegetation cover (urban/untrampled reference ratio) of the three forest types. The trampling tolerance of forest vegetation increased with the productivity of the site (sub-xeric < mesic < herb-rich). Wear of understorey vegetation correlated positively with the number of residents (i.e., recreational pressure) around the forest patch. An increase of 15000 residents within a radius of 1 km around a forest patch was associated with ca. 30% decrease in the relative understorey vegetation cover. The cover of dwarf shrub Vaccinium myrtillus in particular decreased with increasing levels of wear. The cover of mosses in urban forests was less than half of that in untrampled reference areas. Cover of tree saplings, mainly Sorbus aucuparia, and some resilient herbs was higher than in the reference areas. In small urban forest fragments, broad-leaved trees, grasses and herbs were more abundant and mosses were scarcer than in larger urban forest areas. Thus, due to trampling and edge effects, resilient herb and grass species are replacing sensitive dwarf shrubs, mosses and lichens in urban forests. Differences in the soil microbial community structure were found between paths and untrampled areas and the effects of paths extended more than one meter from the paths. Paths supported approximately 25-30% higher microbial biomass with a transition zone of at least 1 m from the path edge. However, microbial activity per unit of biomass was lower on paths than in untrampled areas. Furthermore, microbial biomass and activity were 30-45% lower at the first 20 m into the forest fragments, due to low moisture content of humus near the edge. The decreased microbial activity detected at forest edges and paths implies decreased litter decomposition rates, and thus, a change in nutrient cycling. Changes in the decomposition and nutrient supply may in turn affect the diversity and function of plant communities in urban forests.

The response of Vaccinium myrtillus to variations in grazing intensity in a Scandinavian pine forest on the island of Svanøy

The impact on the performance of dwarf shrub Vaccinium myrtillus L. (bilberry) subjected to differing natural intensities of grazing by Cervus elaphus L. (red deer) was examined in a mature Scandinavian pine forest, on Svanøy, an island on the western coast of Norway. All the study sites were in forest where bilberry dominated the forest floor and no forestry activities had been carried out for several decades. Pellet group counts were used as measure of grazing intensity. Bilberry size, abundance, and fruit set, and invertebrate activity on bilberry were negatively related to grazing intensity. The responses varied between early and late summer (negative or neutral), but impact on plant size was negative throughout the season. Because of the large variation in grazing intensity represented in our study, we can, in contrast to many experimental studies, show that grazing affects the performance of bilberry even at low grazing intensities. The effect on fruit production and invertebrate activity on bilberry indicates that red deer grazing has a negative impact on the population dynamics of the plant and invertebrate herbivores.

Effects of global change on a sub‐Arctic heath: effects of enhanced UV‐B radiation and increased summer precipitation

Summary The responses of sub‐Arctic heathland vegetation to enhanced UV‐B radiation and increased summer precipitation over 7 years were investigated in a field experiment in northern Sweden. Growth, phenology and reproduction of the dominant dwarf shrubs Vaccinium myrtillus, V. uliginosum, V. vitis‐idaea and Empetrum hermaphroditum were studied after 5–7 years of manipulation and retrospective analyses were used to assess growth responses in earlier years. Leaf tissue N and P and 13C natural abundances were determined for V. myrtillus and E. hermaphroditum. Growth responses were also assessed for the moss Hylocomium splendens. The deciduous V. myrtillus showed reduced growth, increased leaf thickness and increased flowering and berry production under enhanced UV‐B in some years. V. uliginosum, V. vitis‐idaea, E. hermaphroditum and H. splendens were, in general, tolerant of UV‐B. Increased precipitation affected growth only in the evergreen species: stem length and branching were sometimes stimulated in E. hermaphroditum, whereas V. vitis‐idaea showed reduced branching. Precipitation also increased leaf thickness in V. uliginosum and reduced flowering and berry production in V. myrtillus. In the interactions that occurred between enhanced UV‐B radiation and increased summer precipitation, combining the two treatments often negated any effect that either may have had separately. The effect of concurrent increases on this ecosystem is therefore likely to be much less than if either occurred singly. Enhanced UV‐B and increased summer precipitation appeared not to effect dwarf shrub abundances during the first 5 years of the experiment, suggesting that overall this heath may be more tolerant of these environmental changes than previously thought.

Boreal forest plants take up organic nitrogen

Plant growth in the boreal forest, the largest terrestrial biome, is generally limited by the availability of nitrogen. The presumed cause of this limitation is slow mineralization of soil organic nitrogen1,2. Here we demonstrate, to our knowledge for the first time, the uptake of organic nitrogen in the field by the trees Pinus sylvestris and Picea abies, the dwarf shrub Vaccinium myrtillus and the grass Deschampsia flexuosa. These results show that these plants, irrespective of their different types of root–fungal associations (mycorrhiza), bypass nitrogen mineralization. A trace of the amino acid glycine, labelled with the stable isotopes 13C and 15N, was injected into the organic (mor) layer of an old successional boreal coniferous forest. Ratios of 13C:15N in the roots showed that at least 91, 64 and 42% of the nitrogen from the absorbed glycine was taken up in intact glycine by the dwarf shrub, the grass and the trees, respectively. Rates of glycine uptake were similar to those of 15N-ammonium. Our data indicate that organic nitrogen is important for these different plants, even when they are competing with each other and with non-symbiotic microorganisms. This has major implications for our understanding of the effects of nitrogen deposition, global warming and intensified forestry.