Abstract. We manipulated light, temperature, and nutrients in moist tussock tundra near Toolik Lake, Alaska to determine how global changes in these parameters might affect community and ecosystem processes. Some of these manipulations altered nutrient availability, growth-form composition, net primary production, and species richness in less than a decade, indicating that arctic vegetation at this site is sensitive to climatic change. In general, short-term (3-yr) responses were poor predictors of longer term (9-yr) changes in community composition. The longer term responses showed closer correspondence to patterns of vegetation distribution along environmental gradients. Nitrogen and phosphorus availability tended to increase in response to elevated temperature, reflecting increased mineralization, and in response to light attenuation, reflecting reduced nutrient uptake by vegetation. Nutrient addition increased biomass and production of deciduous shrubs but reduced growth of evergreen shrubs and nonvascular plants. Light attenuation reduced biomass of all growth forms. Elevated temperature enhanced shrub production but reduced production of nonvascular plants. These contrasting responses to temperature increase and to nutrient addition by different growth forms "canceled out" at the ecosystem level, buffering changes in ecosystem characteristics such as biomass, production, and nutrient uptake. The major effect of elevated temperature was to speed plant response to changes in soil resources and, in the long term (9 yr), to increase nutrient availability through changes in N mineralization. Species within a growth form were similar to one another in their responses to changes in resources (light or nutrients) but showed no consistent response to elevated temperature. Species richness was reduced 30-50% by temperature and nutrient treatments, due to loss of less abundant species. Declines in diversity occurred disproportionately in forbs, which are important for animal nutrition, and in mosses, which maintain soil thermal regime. There was no increased abundance of initially rare species in response to any treatment.During our 9-yr study (the warmest decade on record in the region), biomass of one dominant tundra species unexpectedly changed in control plots in the direction predicted by our experiments and by Holocene pollen records. This suggests that regional climatic warming may already be altering the species composition of Alaskan arctic tundra.
Recent observations of changes in some tundra ecosystems appear to be responses to a warming climate. Several experimental studies have shown that tundra plants and ecosystems can respond strongly to environmental change, including warming; however, most studies were limited to a single location and were of short duration and based on a variety of experimental designs. In addition, comparisons among studies are difficult because a variety of techniques have been used to achieve experimental warming and different measurements have been used to assess responses. We used metaanalysis on plant community measurements from standardized warming experiments at 11 locations across the tundra biome involved in the International Tundra Experiment. The passive warming treatment increased plant-level air temperature by 1-3°C, which is in the range of predicted and observed warming for tundra regions. Responses were rapid and detected in whole plant communities after only two growing seasons. Overall, warming increased height and cover of deciduous shrubs and graminoids, decreased cover of mosses and lichens, and decreased species diversity and evenness. These results predict that warming will cause a decline in biodiversity across a wide variety of tundra, at least in the short term. They also provide rigorous experimental evidence that recently observed increases in shrub cover in many tundra regions are in response to climate warming. These changes have important implications for processes and interactions within tundra ecosystems and between tundra and the atmosphere.arctic and alpine ecosystems ͉ biodiversity ͉ climate change ͉ vegetation change
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