1 Macrolichens are important for the functioning and biodiversity of cold northern ecosystems and their reindeer-based cultures and economies. 2 We hypothesized that, in climatically milder parts of the Arctic, where ecosystems have relatively dense plant canopies, climate warming and/or increased nutrient availability leads to decline in macrolichen abundance as a function of increased abundance of vascular plants. In more open high-arctic or arctic-alpine plant communities such a relationship should be absent. To test this, we synthesized cross-continental arctic vegetation data from ecosystem manipulation experiments simulating mostly warming and increased nutrient availability, and compared these with similar data from natural environmental gradients. 3 Regressions between abundance or biomass of macrolichens and vascular plants were consistently negative across the subarctic and mid-arctic experimental studies. Such a pattern did not emerge in the coldest high-arctic or arctic-alpine sites. The slopes of the negative regressions increased across 10 sites as the climate became milder (as indicated by a simple climatic index) or the vegetation denser (greater site above-ground biomass). 4 Seven natural vegetation gradients in the lower-altitude sub-and mid-arctic zone confirmed the patterns seen in the experimental studies, showing consistent negative relationships between abundance of macrolichens and vascular plants. 5 We conclude that the data supported the hypothesis. Macrolichens in climatically milder arctic ecosystems may decline if and where global changes cause vascular plants to increase in abundance. 6 However, a refining of our findings is needed, for instance by integrating other abiotic and biotic effects such as reindeer grazing feedback on the balance between vascular plants and lichens.

Summary 1Macrolichens are important for the functioning and biodiversity of cold northern ecosystems and their reindeer-based cultures and economies. 2 We hypothesized that, in climatically milder parts of the Arctic, where ecosystems have relatively dense plant canopies, climate warming and/or increased nutrient availability leads to decline in macrolichen abundance as a function of increased abundance of vascular plants. In more open high-arctic or arctic-alpine plant communities such a relationship should be absent. To test this, we synthesized cross-continental arctic vegetation data from ecosystem manipulation experiments simulating mostly warming and increased nutrient availability, and compared these with similar data from natural environmental gradients. 3 Regressions between abundance or biomass of macrolichens and vascular plants were consistently negative across the subarctic and mid-arctic experimental studies. Such a pattern did not emerge in the coldest high-arctic or arctic-alpine sites. The slopes of the negative regressions increased across 10 sites as the climate became milder (as indicated by a simple climatic index) or the vegetation denser (greater site above-ground biomass). 4 Seven natural vegetation gradients in the lower-altitude sub-and mid-arctic zone confirmed the patterns seen in the experimental studies, showing consistent negative relationships between abundance of macrolichens and vascular plants. 5 We conclude that the data supported the hypothesis. Macrolichens in climatically milder arctic ecosystems may decline if and where global changes cause vascular plants to increase in abundance. 6 However, a refining of our findings is needed, for instance by integrating other abiotic and biotic effects such as reindeer grazing feedback on the balance between vascular plants and lichens.

Microbial immobilization may decrease the inorganic nutrient concentrations of the soil to the extent of affecting plant nutrient uptake and growth. We have hypothesized that graminoids with opportunistic nutrient-acquisition strategies are strongly influenced by nutrient limitation imposed by microbes, whereas growth forms such as dwarf shrubs are less affected by the mobilization-immobilization cycles in microbes. By adding NPK fertilizer, labile C (sugar) and fungicide (benomyl) over a 5 yr period in a fully factorial design, we aimed to manipulate the sink-source potential for nutrients in a non-acidic heath tundra soil. After 2 yr, N and P accumulated in the microbial biomass after fertilization with no change in microbial C, which suggests that nutrients did not limit microbial biomass growth. After 5 yr, microbial C was enhanced by 60 % in plots with addition of labile C, which points to C-limitation of the microbial biomass. Microbial biomass N and P tended to increase following addition of labile C, by 10 and 25 %, respectively. This caused decreased availability of NH % + and P, showing close microbial control of nutrient availability. The most common graminoid, Festuca ovina, responded to fertilizer addition with a strong increase, and to labile C addition with a strong decrease in cover, providing the first direct field evidence that nutrient limitation imposed by immobilizing microbes can affect the growth of tundra plants. Also in support of our hypothesis, following addition of labile C the concentrations of N and K in leaves and that of N in roots of F. ovina decreased, whilst the demand of roots for P increased. In contrast, the most common dwarf shrub, Vaccinium uliginosum, was only slightly sensitive to changes in resource availability, showing no cover change after 4 yr addition of labile C and fertilizer, and little change in leaf nutrient concentrations. We suggest that the differential responses of the two growth forms are due to differences in storage and nutrient uptake pathways, with the dwarf shrub having large nutrient storage capacity and access to organic forms of N through its mycorrhizal association. While the fungicide had no effect on ericoid mycorrhizal colonization of roots or symbiotic function inferred from plant "&N natural abundance, it decreased microbial biomass C and N after 2 yr. Throughout the fifth season, the availability of soil NO $ − and inorganic P was decreased with no change in microbial biomass C, N or P, suggesting a negative impact of benomyl on N and P mineralization.

Analyses of changes in vegetation were carried out after three, seven and ten years of fertilizer addition, warming and light attenuation in two subarctic, alpine dwarf shrub heaths. One site was just above the tree line, at ca 450 m a.s.l., and the other at a much colder fell‐field at ca 1150 m altitude. The aim was to investigate how the treatments affected the abundance of different species and growth forms over time, including examinations of transient changes. Grasses, which increased in abundance by fertilizer addition, and cryptogams, which, by contrast, decreased by fertilizer addition and warming, were the most sensitive functional groups to the treatments at both sites. Nutrient addition exerted a stronger and more consistent effect than both shading and warming. Warming at the fell‐field had slightly greater effect than at the warmer tree line with an increase in deciduous shrubs. The decreased abundance of mosses and lichens to fertilizer addition and/or warming was most likely an indirect treatment effect, caused by competition through increased abundance and overgrowth of grasses. Such changes in species composition are likely to alter decomposition rates and the water and energy exchange at the soil surface. We observed few, if any, transient effects of declining responses during the 10 yr of treatments. Instead, there were many cumulative effects of the treatments for all functional groups and many interactions between time and treatment, suggesting that once a change in community composition is triggered, it will continue with unchanged or accelerated rate for a long period of time.

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