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.

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Ecologists have long been intrigued by the ways co-occurring species divide limiting resources. Such resource partitioning, or niche differentiation, may promote species diversity by reducing competition. Although resource partitioning is an important determinant of species diversity and composition in animal communities, its importance in structuring plant communities has been difficult to resolve. This is due mainly to difficulties in studying how plants compete for below-ground resources. Here we provide evidence from a 15N-tracer field experiment showing that plant species in a nitrogen-limited, arctic tundra community were differentiated in timing, depth and chemical form of nitrogen uptake, and that species dominance was strongly correlated with uptake of the most available soil nitrogen forms. That is, the most productive species used the most abundant nitrogen forms, and less productive species used less abundant forms. To our knowledge, this is the first documentation that the composition of a plant community is related to partitioning of differentially available forms of a single limiting resource.

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Carbon allocation to roots in forest ecosystems is estimated from published data on soil respiration and litterfall. On a global scale, rates of in situ soil respiration and aboveground litter production are highly and positively correlated, suggesting that above— and belowground production are controlled by the same factors. This relationship also allows us to predict rates of total soil respiration and total carbon allocation to roots in forest ecosystems from litterfall measurements. Over a gradient of litterfall carbon ranging from 70 to 500 g°m—2°yr—1, total belowground carbon allocation increases from 260 to 1100 g°m—2°yr—1. The ratio of belowground carbon allocation to litterfall decreases from 3.8 to 2.5 as litterfall carbon increases from 70 to 200 g°m—2°yr—1, but changes little (2.5 to 2.2) as litterfall carbon increases from 200 to 500 g°m—2°yr—1. Use of this relationship permits the construction of simple carbon budgets that can be used to place upper limits on estimates of fine root production in forest ecosystems. Determining live—root respiration rates in forest ecosystems will further constrain the range of possible root production rates.

Quercus-Acer (oak-maple) st'.111d in c~ntral l\ l~1ssac~1u selts, US/\. The key hy pothesis J!OVe rnmg the b1oloA1cal component of the model is Urnt stomata! conductance (g,.) is var ied s o that d ail y carbon uptake per unit of foliar nit r oj:!en is maximized wit hin the limitations of ca1101>Y water avr1ilability. T he h ydraulic system is modelled as :m a nalo~ue to simple electr ical circuits in parallel. includinJ! a separate soil hydraulic resistance. plant resistance and plant capacitance for each c:m op y layer. Stomata! openin(! is initially controlled to conserve plant water stores and delay the o nset or water stress. Stomata! clos ure at :i threshold minimum l~1f water potential prevents xyle m cavitation and contr ols the maximum r:itc of wate r llux through the h ydraulic system. We show a strong correlat ion between predicted hourly C0 2 exchange rate (r2 = 0·86) and LE (r 2 = 0·87) with independent whole-forest measurements made by the eddy correlation method during the s ummer of 1992. Our theoretical derivation s hows that observed relationships between C0 2 assimilation a nd LE nux can he explained on the basis of s tomata! behaviour 01>timizing c:irhon gain, and provides mt exi>licit link hetwecn canopy structure, soil pro1>erties. atmos pheric conditions and stomatal conductance.Ke\'-111ords: Q11erc11s r11hm: l\cl'r rulnw11: soil-pla11t-a1r~10spherc coniinuum model: photosynlhcsis: plant hydr:1ulic conductance: s1o ma1al conduc tance.