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We present a framework for assessing biogeochemical recovery of terrestrial ecosystems from disturbance. We identify three recovery phases. In Phase 1, nitrogen is redistributed from soil organic matter to vegetation, but the ecosystem continues to lose nitrogen because the recovering vegetation cannot take up nitrogen as fast as it is released from soil. In Phase 2, the ecosystem begins re-accumulating nitrogen and converges on a quasi-steady state in which vegetation and soil-microbial processes are in balance. In Phase 3, vegetation and soil-microbial processes remain in balance and the ecosystem slowly re-accumulates the remaining nitrogen. Phase 3 follows a balanced-accumulation trajectory along a continuum of quasi-steady states that approaches the true steady state asymptotically. We examine the effects of three ecosystem properties on recovery: openness of the nitrogen cycle, nitrogen distribution in and turnover between vegetation and soils, and the proportion of nitrogen losses that are in a refractory form. Openness exacerbates Phase 1 nitrogen losses but speeds recovery in Phases 2 and 3. A high fraction of ecosystem nitrogen in vegetation, resulting from nitrogen turnover that is slow in vegetation but fast in soil, exacerbates Phase 1 nitrogen losses but speeds recovery in Phases 2 and 3. A high proportion of nitrogen loss in refractory form mitigates Phase 1 nitrogen losses and speeds recovery in Phases 2 and 3. Application of our conceptual framework requires empirical recognition of the continuum of quasi-steady states constituting the balanced-accumulation trajectory and a distinction between the balanced-accumulation trajectory and the true steady state.

Releases of the greenhouse gases carbon dioxide (CO2) and methane (CH4) from thawing permafrost are expected to be among the largest feedbacks to climate from arctic ecosystems. However, the current net carbon (C) balance of terrestrial arctic ecosystems is unknown. Recent studies suggest that these ecosystems are sources, sinks, or approximately in balance at present. This uncertainty arises because there are few long-term continuous measurements of arctic tundra CO2 fluxes over the full annual cycle. Here, we describe a pattern of CO2 loss based on the longest continuous record of direct measurements of CO2 fluxes in the Alaskan Arctic, from two representative tundra ecosystems, wet sedge and heath tundra. We also report on a shorter time series of continuous measurements from a third ecosystem, tussock tundra. The amount of CO2 loss from both heath and wet sedge ecosystems was related to the timing of freeze-up of the soil active layer in the fall. Wet sedge tundra lost the most CO2 during the anomalously warm autumn periods of September–December 2013–2015, with CH4 emissions contributing little to the overall C budget. Losses of C translated to approximately 4.1 and 1.4% of the total soil C stocks in active layer of the wet sedge and heath tundra, respectively, from 2008 to 2015. Increases in air temperature and soil temperatures at all depths may trigger a new trajectory of CO2 release, which will be a significant feedback to further warming if it is representative of larger areas of the Arctic.

We scale a model of net ecosystem CO2 exchange (NEE) for tundra ecosystems and assess model performance using eddy covariance measurements at three tundra sites. The model, initially developed using instantaneous (seconds–minutes) chamber flux (~m2) observations, independently represents ecosystem respiration (ER) and gross primary production (GPP), and requires only temperature (T), photosynthetic photon flux density (I 0), and leaf area index (L) as inputs. We used a synthetic data set to parameterize the model so that available in situ observations could be used to assess the model. The model was then scaled temporally to daily resolution and spatially to about 1 km2 resolution, and predicted values of NEE, and associated input variables, were compared to observations obtained from eddy covariance measurements at three flux tower sites over several growing seasons. We compared observations to modeled NEE calculated using T and I 0 measured at the towers, and L derived from MODIS data. Cumulative NEE estimates were within 17 and 11% of instrumentation period and growing season observations, respectively. Predictions improved when one site-year experiencing anomalously dry conditions was excluded, indicating the potential importance of stomatal control on GPP and/or soil moisture on ER. Notable differences in model performance resulted from ER model formulations and differences in how L was estimated. Additional work is needed to gain better predictive ability in terms of ER and L. However, our results demonstrate the potential of this model to permit landscape scale estimates of NEE using relatively few and simple driving variables that are easily obtained via satellite remote sensing.

Nutrient availability in the arctic is expected to increase in the next century due to accelerated decomposition associated with warming and, to a lesser extent, increased nitrogen deposition. To explore how changes in nutrient availability affect ecosystem carbon (C) cycling, we used radiocarbon to quantify changes in belowground C dynamics associated with long-term fertilization of graminoid-dominated tussock tundra at Toolik Lake, Alaska. Since 1981, yearly fertilization with nitrogen (N) and phosphorus (P) has resulted in a shift to shrub-dominated vegetation. These combined changes have altered the quantity and quality of litter inputs, the vertical distribution and dynamics of fine roots, and the decomposition rate of soil organic C. The loss of C from the deep organic and mineral soil has more than offset the C accumulation in the litter and upper organic soil horizons. In the litter and upper organic horizons, radiocarbon measurements show that increased inputs resulted in overall C accumulation, despite being offset by increased decomposition in some soil pools. To reconcile radiocarbon observations in the deeper organic and mineral soil layers, where most of the ecosystem C loss occurred, both a decrease in input of new root material and a dramatic increase of decomposition rates in centuries-old soil C pools were required. Therefore, with future increases in nutrient availability, we may expect substantial losses of C which took centuries to accumulate. © 2007 Springer Science+Business Media, LLC.

In temperate forests, symbiotic nitrogen (N) fixation is restricted to the early phases of succession despite the persistence of N limitation on production late in succession. This paradox has yet to be explained adequately. We hypothesized that the restriction of N fixation to early stages of succession results from the optimization of resource allocation in the vegetation. Because of this optimization, N fixation should be restricted to periods when fixation is less costly than N uptake. Our analysis differs from others in the way we calculate the cost of N uptake; we assess the cost of N uptake as the amount of carbon (C) that could be assimilated if the resources necessary to acquire one gram of N from the soil were allocated instead to photosynthesis. We then simulate N fixation as an asymptotic function of the difference in cost between N uptake and N fixation and proportional to the abundance of host tissues for the N-fixing symbionts. The factors that contribute to conditions that favor N fixation are (a) elevated-carbon dioxide (CO2) concentrations, (b) an open canopy, (c) low available N in the soil, and (d) a soil volume already well exploited by roots. Our results indicate that changes in the relative cost of uptake vs fixation can explain most of the pattern in fixation through both primary and secondary succession, but that competitive interactions with nonfixing species play a role in the final exclusion of fixation in later stages of succession.

Understanding terrestrial carbon metabolism is critical because terrestrial ecosystems play a major role in the global carbon cycle. Furthermore, humans have severely disrupted the carbon cycle in ways that will alter the climate system and directly affect terrestrial metabolism. Changes in terrestrial metabolism may well be as important an indicator of global change as the changing temperature signal. Improving our understanding of the carbon cycle at various spatial and temporal scales will require the integration of multiple, complementary and independent methods that are used by different research communities. Tools such as air sampling networks, inverse numerical methods, and satellite data (top-down approaches) allow us to study the strength and location of the global- and continental-scale carbon sources and sinks. Bottom-up studies provide estimates of carbon fluxes at finer spatial scales and examine the mechanisms that control fluxes at the ecosystem, landscape, and regional scales. Bottom-up approaches include comparative and process studies (for example, ecosystem manipulative experiments) that provide the necessary mechanistic information to develop and validate terrestrial biospheric models. An iteration and reiteration of top-down and bottom-up approaches will be necessary to help constrain measurements at various scales. We propose a major international effort to coordinate and lead research programs of global scope of the carbon cycle.

The capture and efficient use of limiting resources influence the competitive success of individual plant species as well as species diversity across resource gradients. In simulations, efficient nutrient acquisition or nutrient retention by species were key predictors of success when nutrients were limiting. Increased nutrient supply favored species with characteristics that improved light interception or light use. Ecological theory suggests that low diversity on fertile sites may be a consequence of competitive exclusion by one or a few species with superior light-interception characteristics. On infertile sites, competitive exclusion may be a function of superior nutrient-acquisition characteristics in species. At intermediate fertility, a shift from single-resource specialization to a balanced effort in the acquisition of multiple resources should allow for greater species diversity. Thus, a unimodal relationship between diversity and nutrient supply, vegetation biomass, or productivity is predicted. However, simulations demonstrated alternate relationships depending on the ecosystem characteristic to which diversity was compared. Diversity was greatest at intermediate total biomass but increased monotonically with net primary production and nitrogen (N) supply. The highest diversity occurred midrange on a scale of community-level leaf area to fine-root length ratios, which in the context of the model indicates that the vegetation as a whole was simultaneously limited by both N and light and that effort toward the acquisition of both resources is distributed in such a way that both resources are equally exploited. Diversity was lowered by the presence of species with a superior ability to sequester resources.