Recovery of arctic tundra from thermal erosion disturbance is constrained by nutrient accumulation: a modeling analysis.

Abstract. We calibrated the Multiple Element Limitation (MEL) model to Alaskan arctic tundra to simulate recovery of thermal erosion features (TEFs) caused by permafrost thaw and mass wasting. TEFs could significantly alter regional carbon (C) and nutrient budgets because permafrost soils contain large stocks of soil organic matter (SOM) and TEFs are expected to become more frequent as the climate warms. We simulated recovery following TEF stabilization and did not address initial, short-term losses of C and nutrients during TEF formation. To capture the variability among and within TEFs, we modeled a range of post-stabilization conditions by varying the initial size of SOM stocks and nutrient supply rates. Simulations indicate that nitrogen (N) losses after the TEF stabilizes are small, but phosphorus (P) losses continue. Vegetation biomass recovered 90% of its undisturbed C, N, and P stocks in 100 years using nutrients mineralized from SOM. Because of low litter inputs but continued decomposition, younger SOM continued to be lost for 10 years after the TEF began to recover, but recovered to about 84% of its undisturbed amount in 100 years. The older recalcitrant SOM in mineral soil continued to be lost throughout the 100-year simulation. Simulations suggest that biomass recovery depended on the amount of SOM remaining after disturbance. Recovery was initially limited by the photosynthetic capacity of vegetation but became co-limited by N and P once a plant canopy developed. Biomass and SOM recovery was enhanced by increasing nutrient supplies, but the magnitude, source, and controls on these supplies are poorly understood. Faster mineralization of nutrients from SOM (e.g., by warming) enhanced vegetation recovery but delayed recovery of SOM. Taken together, these results suggest that although vegetation and surface SOM on TEFs recovered quickly (25 and 100 years, respectively), the recovery of deep, mineral soil SOM took centuries and represented a major ecosystem C loss.

Pan-Arctic modelling of net ecosystem exchange of CO2.

Net ecosystem exchange (NEE) of C varies greatly among Arctic ecosystems. Here, we show that approximately 75 per cent of this variation can be accounted for in a single regression model that predicts NEE as a function of leaf area index (LAI), air temperature and photosynthetically active radiation (PAR). The model was developed in concert with a survey of the light response of NEE in Arctic and subarctic tundras in Alaska, Greenland, Svalbard and Sweden. Model parametrizations based on data collected in one part of the Arctic can be used to predict NEE in other parts of the Arctic with accuracy similar to that of predictions based on data collected in the same site where NEE is predicted. The principal requirement for the dataset is that it should contain a sufficiently wide range of measurements of NEE at both high and low values of LAI, air temperature and PAR, to properly constrain the estimates of model parameters. Canopy N content can also be substituted for leaf area in predicting NEE, with equal or greater accuracy, but substitution of soil temperature for air temperature does not improve predictions. Overall, the results suggest a remarkable convergence in regulation of NEE in diverse ecosystem types throughout the Arctic.

Recovery from disturbance requires resynchronization of ecosystem nutrient cycles.

Nitrogen (N) and phosphorus (P) are tightly cycled in most terrestrial ecosystems, with plant uptake more than 10 times higher than the rate of supply from deposition and weathering. This near-total dependence on recycled nutrients and the stoichiometric constraints on resource use by plants and microbes mean that the two cycles have to be synchronized such that the ratio of N:P in plant uptake, litterfall, and net mineralization are nearly the same. Disturbance can disrupt this synchronization if there is a disproportionate loss of one nutrient relative to the other. We model the resynchronization of N and P cycles following harvest of a northern hardwood forest. In our simulations, nutrient loss in the harvest is small relative to postharvest losses. The low N:P ratio of harvest residue results in a preferential release of P and retention of N. The P release is in excess of plant requirements and P is lost from the active ecosystem cycle through secondary mineral formation and leaching early in succession. Because external P inputs are small, the resynchronization of the N and P cycles later in succession is achieved by a commensurate loss of N. Through succession, the ecosystem undergoes alternating periods of N limitation, then P limitation, and eventually co-limitation as the two cycles resynchronize. However, our simulations indicate that the overall rate and extent of recovery is limited by P unless a mechanism exists either to prevent the P loss early in succession (e.g., P sequestration not stoichiometrically constrained by N) or to increase the P supply to the ecosystem later in succession (e.g., biologically enhanced weathering). Our model provides a heuristic perspective from which to assess the resynchronization among tightly cycled nutrients and the effect of that resynchronization on recovery of ecosystems from disturbance.

Modelling carbon responses of tundra ecosystems to historical and projected climate: a comparison of a plot- and a global-scale ecosystem model to identify process-based uncertainties.

We are developing a process-based modelling approach to investigate how carbon (C) storage of tundra across the entire Arctic will respond to projected climate change. To implement the approach, the processes that are least understood, and thus have the most uncertainty, need to be identified and studied. In this paper, we identified a key uncertainty by comparing the responses of C storage in tussock tundra at one site between the simulations of two models - one a global-scale ecosystem model (Terrestrial Ecosystem Model, TEM) and one a plot-scale ecosystem model (General Ecosystem Model, GEM). The simulations spanned the historical period (1921-94) and the projected period (1995-2100). In the historical period, the model simulations of net primary production (NPP) differed in their sensitivity to variability in climate. However, the long-term changes in C storage were similar in both simulations, because the dynamics of heterotrophic respiration (RH ) were similar in both models. In contrast, the responses of C storage in the two model simulations diverged during the projected period. In the GEM simulation for this period, increases in RH tracked increases in NPP, whereas in the TEM simulation increases in RH lagged increases in NPP. We were able to make the long-term C dynamics of the two simulations agree by parameterizing TEM to the fast soil C pools of GEM. We concluded that the differences between the long-term C dynamics of the two simulations lay in modelling the role of the recalcitrant soil C. These differences, which reflect an incomplete understanding of soil processes, lead to quite different projections of the response of pan-Arctic C storage to global change. For example, the reference parameterization of TEM resulted in an estimate of cumulative C storage of 2032 g C m-2 for moist tundra north of 50°N, which was substantially higher than the 463 g C m-2 estimated for a parameterization of fast soil C dynamics. This uncertainty in the depiction of the role of recalcitrant soil C in long-term ecosystem C dynamics resulted from our incomplete understanding of controls over C and N transformations in Arctic soils. Mechanistic studies of these issues are needed to improve our ability to model the response of Arctic ecosystems to global change.

Arctic and boreal ecosystems of western North America as components of the climate system.

Synthesis of results from several Arctic and boreal research programmes provides evidence for the strong role of high-latitude ecosystems in the climate system. Average surface air temperature has increased 0.3 °C per decade during the twentieth century in the western North American Arctic and boreal forest zones. Precipitation has also increased, but changes in soil moisture are uncertain. Disturbance rates have increased in the boreal forest; for example, there has been a doubling of the area burned in North America in the past 20 years. The disturbance regime in tundra may not have changed. Tundra has a 3-6-fold higher winter albedo than boreal forest, but summer albedo and energy partitioning differ more strongly among ecosystems within either tundra or boreal forest than between these two biomes. This indicates a need to improve our understanding of vegetation dynamics within, as well as between, biomes. If regional surface warming were to continue, changes in albedo and energy absorption would likely act as a positive feedback to regional warming due to earlier melting of snow and, over the long term, the northward movement of treeline. Surface drying and a change in dominance from mosses to vascular plants would also enhance sensible heat flux and regional warming in tundra. In the boreal forest of western North America, deciduous forests have twice the albedo of conifer forests in both winter and summer, 50-80% higher evapotranspiration, and therefore only 30-50% of the sensible heat flux of conifers in summer. Therefore, a warming-induced increase in fire frequency that increased the proportion of deciduous forests in the landscape, would act as a negative feedback to regional warming. Changes in thermokarst and the aerial extent of wetlands, lakes, and ponds would alter high-latitude methane flux. There is currently a wide discrepancy among estimates of the size and direction of CO2 flux between high-latitude ecosystems and the atmosphere. These discrepancies relate more strongly to the approach and assumptions for extrapolation than to inconsistencies in the underlying data. Inverse modelling from atmospheric CO2 concentrations suggests that high latitudes are neutral or net sinks for atmospheric CO2 , whereas field measurements suggest that high latitudes are neutral or a net CO2 source. Both approaches rely on assumptions that are difficult to verify. The most parsimonious explanation of the available data is that drying in tundra and disturbance in boreal forest enhance CO2 efflux. Nevertheless, many areas of both tundra and boreal forests remain net sinks due to regional variation in climate and local variation in topographically determined soil moisture. Improved understanding of the role of high-latitude ecosystems in the climate system requires a concerted research effort that focuses on geographical variation in the processes controlling land-atmosphere exchange, species composition, and ecosystem structure. Future studies must be conducted over a long enough time-period to detect and quantify ecosystem feedbacks.

Effects of drainage and temperature on carbon balance of tussock tundra micrososms.

We examined the importance of temperature (7°C or 15°C) and soil moisture regime (saturated or field capacity) on the carbon (C) balance of arctic tussock tundra microcosms (intact blocks of soil and vegetation) in growth chambers over an 81-day simulated growing season. We measured gaseous CO2 exchanges, methane (CH4) emissions, and dissolved C losses on intact blocks of tussock (Eriophorum vaginatum) and intertussock (moss-dominated). We hypothesized that under increased temperature and/or enhanced drainage, C losses from ecosystem respiration (CO2 respired by plants and heterotrophs) would exceed gains from gross photosynthesis causing tussock tundra to become a net source of C to the atmosphere. The field capacity moisture regime caused a decrease in net CO2 storage (NEP) in tussock tundra micrososms. This resulted from a stimulation of ecosystem respiration (probably mostly microbial) with enhanced drainage, rather than a decrease in gross photosynthesis. Elevated temperature alone had no effect on NEP because CO2 losses from increased ecosystem respiration at elevated temperature were compensated by increased CO2 uptake (gross photosynthesis). Although CO2 losses from ecosystem respiration were primarily limited by drainage, CH4 emissions, in contrast, were dependent on temperature. Furthermore, substantial dissolved C losses, especially organic C, and important microhabitat differences must be considered in estimating C balance for the tussock tundra system. As much as ∼ 20% of total C fixed in photosynthesis was lost as dissolved organic C. Tussocks stored ∼ 2x more C and emitted 5x more methane than intertussocks. In spite of the limitations of this microcosm experiment, this study has further elucidated the critical role of soil moisture regime and dissolved C losses in regulating net C balance of arctic tussock tundra.

Potential Net Primary Productivity in South America: Application of a Global Model.

We use a mechanistically based ecosystem simulation model to describe and analyze the spatial and temporal patterns of terrestrial net primary productivity (NPP) in South America. The Terrestrial Ecosystem Model (TEM) is designed to predict major carbon and nitrogen fluxes and pool sizes in terrestrial ecosystems at continental to global scales. Information from intensively studies field sites is used in combination with continental-scale information on climate, soils, and vegetation to estimate NPP in each of 5888 non-wetland, 0.5° latitude °0.5° longitude grid cells in South America, at monthly time steps. Preliminary analyses are presented for the scenario of natural vegetation throughout the continent, as a prelude to evaluating human impacts on terrestrial NPP. The potential annual NPP of South America is estimated to be 12.5 Pg/yr of carbon (26.3 Pg/yr of organic matter) in a non-wetland area of 17.0 ° 106 km2 . More than 50% of this production occurs in the tropical and subtropical evergreen forest region. Six independent model runs, each based on an independently derived set of model parameters, generated mean annual NPP estimates for the tropical evergreen forest region ranging from 900 to 1510 g°m-2 °yr-1 of carbon, with an overall mean of 1170 g°m-2 °yr-1 . Coefficients of variation in estimated annual NPP averaged 20% for any specific location in the evergreen forests, which is probably within the confidence limits of extant NPP measurements. Predicted rates of mean annual NPP in other types of vegetation ranged from 95 g°m-2 °yr-1 in arid shrublands to 930 g°m@ ?yr-1 in savannas, and were within the ranges measured in empirical studies. The spatial distribution of predicted NPP was directly compared with estimates made using the Miami mode of Lieth (1975). Overall, TEM predictions were °10% lower than those of the Miami model, but the two models agreed closely on the spatial patterns of NPP in south America. Unlike previous models, however, TEM estimates NPP monthly, allowing for the evaluation of seasonal phenomena. This is an important step toward integration of ecosystem models with remotely sensed information, global climate models, and atmospheric transport models, all of which are evaluated at comparable spatial and temporal scales. Seasonal patterns of NPP in South America are correlated with moisture availability in most vegetation types, but are strongly influenced by seasonal differences in cloudiness in the tropical evergreen forests. On an annual basis, moisture availability was the factor that was correlated most strongly with annual NPP in South America, but differences were again observed among vegetation types. These results allow for the investigation and analysis of climatic controls over NPP at continental scales, within and among vegetation types, and within years. Further model validation is needed. Nevertheless, the ability to investigate NPP-environment interactions with a high spatial and temporal resolution at continental scales should prove useful if not essential for rigorous analysis of the potential effects of global climate changes on terrestrial ecosystems.