In a decade-long soil warming experiment in a mid-latitude hardwood forest, we documented changes in soil carbon and nitrogen cycling in order to investigate the consequences of these changes for the climate system. Here we show that whereas soil warming accelerates soil organic matter decay and carbon dioxide fluxes to the atmosphere, this response is small and short-lived for a mid-latitude forest, because of the limited size of the labile soil carbon pool. We also show that warming increases the availability of mineral nitrogen to plants. Because plant growth in many mid-latitude forests is nitrogen-limited, warming has the potential to indirectly stimulate enough carbon storage in plants to at least compensate for the carbon losses from soils. Our results challenge assumptions made in some climate models that lead to projections of large long-term releases of soil carbon in response to warming of forest ecosystems.

Fine roots ( 1 mm diameter) are critical in plant water and nutrient absorption, and it is important to understand how rising atmospheric CO 2 will affect them as part of terrestrial ecosystem responses to global change. This study's objective was to determine the effects of elevated CO 2 on production, mortality, and standing crops of fine root length over 2 years in a free-air CO 2 enrichment (FACE) facility in the Mojave Desert of southern Nevada, USA. Three replicate 25 m diameter FACE rings were maintained at ambient ( $ 370 lmol mol À1 ) and elevated CO 2 ( $ 550 lmol mol À1 ) atmospheric concentrations. Twenty-eight minirhizotron tubes were placed in each ring to sample three microsite locations: evergreen Larrea shrubs, drought-deciduous Ambrosia shrubs, and along systematic community transects (primarily in shrub interspaces which account for $ 85% of the area). Seasonal dynamics were similar for ambient and elevated CO 2 : fine root production peaked in April-June, with peak standing crop occurring about 1 month later, and peak mortality occurring during the hot summer months, with higher values for all three measures in a wet year compared with a dry year. Fine root standing crop, production, and mortality were not significantly different between treatments except standing crop along community transects, where fine root length was significantly lower in elevated CO 2 . Fine root turnover (annual cumulative mortality/mean standing crop) ranged from 2.33 to 3.17 year À1 , and was not significantly different among CO 2 treatments, except for community transect tubes where it was significantly lower for elevated CO 2 . There were no differences in fine root responses to CO 2 between evergreen (Larrea) and drought-deciduous (Ambrosia) shrubs. Combined with observations of increased leaf-level water-use efficiency and lack of soil moisture differences, these results suggest that under elevated CO 2 conditions, reduced root systems (compared with ambient CO 2 ) appear sufficient to provide resources for modest aboveground production increases across the community, but in more fertile shrub microsites, fine root systems of comparable size with those in ambient CO 2 were required to support the greater aboveground production increases. For community transects, development of the difference in fine root standing crops occurred primarily through lower stimulation of fine root production in the elevated CO 2 treatment during periods of high water availability.

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High nitrogen (N) accumulation in terrestrial ecosystems can shift patterns of nutrient limitation and deficiency beyond N toward other nutrients, most notably phosphorus (P) and base cations (calcium [Ca], magnesium [Mg], and potassium [K]). We examined how naturally high N accumulation from a legacy of symbiotic N fixation shaped P and base cation cycling across a gradient of nine temperate conifer forests in the Oregon Coast Range. We were particularly interested in whether long-term legacies of symbiotic N fixation promoted coupled N and organic P accumulation in soils, and whether biotic demands by non-fixing vegetation could conserve ecosystem base cations as N accumulated. Total soil N (0-100 cm) pools increased nearly threefold across the N gradient, leading to increased nitrate leaching, declines in soil pH from 5.8 to 4.2, 10-fold declines in soil exchangeable Ca, Mg, and K, and increased mobilization of aluminum. These results suggest that long-term N enrichment had acidified soils and depleted much of the readily weatherable base cation pool. Soil organic P increased with both soil N and C across the gradient, but soil inorganic P, biomass P, and P leaching loss did not vary with N, implying that historic symbiotic N fixation promoted soil organic P accumulation and P sufficiency for non-fixers. Even though soil pools of Ca, Mg, and K all declined as soil N increased, only Ca declined in biomass pools, suggesting the emergence of Ca deficiency at high N. Biotic conservation and tight recycling of Ca increased in response to whole-ecosystem Ca depletion, as indicated by preferential accumulation of Ca in biomass and surface soil. Our findings support a hierarchical model of coupled N-Ca cycling under long-term soil N enrichment, whereby ecosystem-level N saturation and nitrate leaching deplete readily available soil Ca, stimulating biotic Ca conservation as overall supply diminishes. We conclude that a legacy of biological N fixation can increase N and P accumulation in soil organic matter to the point that neither nutrient is limiting to subsequent non-fixers, while also resulting in natural N saturation that intensifies base cation depletion and deficiency.

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