Postfire carbon pools and fluxes in semiarid ponderosa pine in Central Oregon

Forest fire dramatically affects the carbon storage and underlying mechanisms that control the carbon balance of recovering ecosystems. In western North America where fire extent has increased in recent years, we measured carbon pools and fluxes in moderately and severely burned forest stands 2 years after a fire to determine the controls on net ecosystem productivity (NEP) and make comparisons with unburned stands in the same region. Total ecosystem carbon in soil and live and dead pools in the burned stands was on average 66% that of unburned stands (11.0 and 16.5 kg C m⁻², respectively, P<0.01). Soil carbon accounted for 56% and 43% of the carbon pools in burned and unburned stands. NEP was significantly lower in severely burned compared with unburned stands (P<0.01) with an increasing trend from −125±44 g C m⁻² yr⁻¹ (±1 SD) in severely burned stands (stand replacing fire), to −38±96 and +50±47 g C m⁻² yr⁻¹ in moderately burned and unburned stands, respectively. Fire of moderate severity killed 82% of trees <20 cm in diameter (diameter at 1.3 m height, DBH); however, this size class only contributed 22% of prefire estimates of bole wood production. Larger trees (> 20 cm DBH) suffered only 34% mortality under moderate severity fire and contributed to 91% of postfire bole wood production. Growth rates of trees that survived the fire were comparable with their prefire rates. Net primary production NPP (g C m⁻² yr⁻¹, ±1 SD) of severely burned stands was 47% of unburned stands (167±76, 346±148, respectively, P<0.05), with forb and grass aboveground NPP accounting for 74% and 4% of total aboveground NPP, respectively. Based on continuous seasonal measurements of soil respiration in a severely burned stand, in areas kept free of ground vegetation, soil heterotrophic respiration accounted for 56% of total soil CO₂ efflux, comparable with the values of 54% and 49% previously reported for two of the unburned forest stands. Estimates of total ecosystem heterotrophic respiration (R_h) were not significantly different between stand types 2 years after fire. The ratio NPP/R_h averaged 0.55, 0.85 and 1.21 in the severely burned, moderately burned and unburned stands, respectively. Annual soil CO₂ efflux was linearly related to aboveground net primary productivity (ANPP) with an increase in soil CO₂ efflux of 1.48 g C yr⁻¹ for every 1 g increase in ANPP (P<0.01, r²= 0.76). There was no significant difference in this relationship between the recently burned and unburned stands. Contrary to expectations that the magnitude of NEP 2 years postfire would be principally driven by the sudden increase in detrital pools and increased rates of R_h, the data suggest NPP was more important in determining postfire NEP.     

Supply-side controls on soil respiration among Oregon forests

To test the hypothesis that variation in soil respiration is related to plant production across a diverse forested landscape, we compared annual soil respiration rates with net primary production and the subsequent allocation of carbon to various ecosystem pools, including leaves, fine roots, forests floor, and mineral soil for 36 independent plots arranged as three replicates of four age classes in three climatically distinct forest types. Across all plots, annual soil respiration was not correlated with aboveground net primary production (R²=0.06, P>0.1) but it was moderately correlated with belowground net primary production (R²=0.46, P<0.001). Despite the wide range in temperature and precipitation regimes experienced by these forests, all exhibited similar soil respiration per unit live fine root biomass, with about 5 g of carbon respired each year per 1 g of fine root carbon (R²=0.45, P<0.001). Annual soil respiration was only weakly correlated with dead carbon pools such as forest floor and mineral soil carbon (R²=0.14 and 0.12, respectively). Trends between soil respiration, production, and root mass among age classes within forest type were inconsistent and do not always reflect cross‐site trends. These results are consistent with a growing appreciation that soil respiration is strongly influenced by the supply of carbohydrates to roots and the rhizosphere, and that some regional patterns of soil respiration may depend more on belowground carbon allocation than the abiotic constraints imposed on subsequent metabolism.     

Estimating carbon carrying capacity in natural forest ecosystems across heterogeneous landscapes: addressing sources of error

Evaluating contributions of forest ecosystems to climate change mitigation requires well‐calibrated carbon cycle models with quantified baseline carbon stocks. An appropriate baseline for carbon accounting of natural forests at landscape scales is carbon carrying capacity (CCC); defined as the mass of carbon stored in an ecosystem under prevailing environmental conditions and natural disturbance regimes but excluding anthropogenic disturbance. Carbon models require empirical measurements for input and calibration, such as net primary production (NPP) and total ecosystem carbon stock (equivalent to CCC at equilibrium). We sought to improve model calibration by addressing three sources of errors that cause uncertainty in carbon accounting across heterogeneous landscapes: (1) data‐model representation, (2) data‐object representation, (3) up‐scaling. We derived spatially explicit empirical models based on environmental variables across landscape scales to estimate NPP (based on a synthesis of global site data of NPP and gross primary productivity, n=27), and CCC (based on site data of carbon stocks in natural eucalypt forests of southeast Australia, n=284). The models significantly improved predictions, each accounting for 51% of the variance. Our methods to reduce uncertainty in baseline carbon stocks, such as using appropriate calibration data from sites with minimal human disturbance, measurements of large trees and incorporating environmental variability across the landscape, have generic application to other regions and ecosystem types. These analyses resulted in forest CCC in southeast Australia (mean total biomass of 360 t C ha^?1, with cool moist temperate forests up to 1000 t C ha^?1) that are larger than estimates from other national and international (average biome 202 t C ha^?1) carbon accounting systems. Reducing uncertainty in estimates of carbon stocks in natural forests is important to allow accurate accounting for losses of carbon due to human activities and sequestration of carbon by forest growth.     

Biometric Based Carbon Flux Measurements and Net Ecosystem Production (NEP) in a Temperate Deciduous Broad-Leaved Forest Beneath a Flux Tower

Biometric based carbon flux measurements were conducted over 5 years (1999–2003) in a temperate deciduous broad-leaved forest of the AsiaFlux network to estimate net ecosystem production (NEP). Biometric based NEP, as measured by the balance between net primary production (including NPP of canopy trees and of forest floor dwarf bamboo) and heterotrophic respiration (RH), clarified the contribution of various biological processes to the ecosystem carbon budget, and also showed where and how the forest is storing C. The mean NPP of the trees was 5.4 ± 1.07 t C ha⁻¹ y⁻¹, including biomass increment (0.3 ± 0.82 t C ha⁻¹ y⁻¹), tree mortality (1.0 ± 0.61 t C ha⁻¹ y⁻¹), aboveground detritus production (2.3 ± 0.39 t C ha⁻¹ y⁻¹) and belowground fine root production (1.8 ± 0.31 t C ha⁻¹ y⁻¹). Annual biomass increment was rather small because of high tree mortality during the 5 years. Total NPP at the site was 6.5 ± 1.07 t C ha⁻¹ y⁻¹, including the NPP of the forest floor community (1.1 ± 0.06 t C ha⁻¹ y⁻¹). The soil surface CO₂ efflux (RS) was averaged across the 5 years of record using open-flow chambers. The mean estimated annual RS amounted to 7.1 ± 0.44 t C ha⁻¹, and the decomposition of soil organic matter (SOM) was estimated at 3.9 ± 0.24 t C ha⁻¹. RH was estimated at 4.4 ± 0.32 t C ha⁻¹ y⁻¹, which included decomposition of coarse woody debris. Biometric NEP in the forest was estimated at 2.1 ± 1.15 t C ha⁻¹ y⁻¹, which agreed well with the eddy-covariance based net ecosystem exchange (NEE). The contribution of woody increment (Δbiomass + mortality) of the canopy trees to NEP was rather small, and thus the SOM pool played an important role in carbon storage in the temperate forest. These results suggested that the dense forest floor of dwarf bamboo might have a critical role in soil carbon sequestration in temperate East Asian deciduous forests.     

Reconciling Carbon-cycle Concepts, Terminology, and Methods

Recent projections of climatic change have focused a great deal of scientific and public attention on patterns of carbon (C) cycling as well as its controls, particularly the factors that determine whether an ecosystem is a net source or sink of atmospheric carbon dioxide (CO₂). Net ecosystem production (NEP), a central concept in C-cycling research, has been used by scientists to represent two different concepts. We propose that NEP be restricted to just one of its two original definitions—the imbalance between gross primary production (GPP) and ecosystem respiration (ER). We further propose that a new term—net ecosystem carbon balance (NECB)—be applied to the net rate of C accumulation in (or loss from [negative sign]) ecosystems. Net ecosystem carbon balance differs from NEP when C fluxes other than C fixation and respiration occur, or when inorganic C enters or leaves in dissolved form. These fluxes include the leaching loss or lateral transfer of C from the ecosystem; the emission of volatile organic C, methane, and carbon monoxide; and the release of soot and CO₂ from fire. Carbon fluxes in addition to NEP are particularly important determinants of NECB over long time scales. However, even over short time scales, they are important in ecosystems such as streams, estuaries, wetlands, and cities. Recent technological advances have led to a diversity of approaches to the measurement of C fluxes at different temporal and spatial scales. These approaches frequently capture different components of NEP or NECB and can therefore be compared across scales only by carefully specifying the fluxes included in the measurements. By explicitly identifying the fluxes that comprise NECB and other components of the C cycle, such as net ecosystem exchange (NEE) and net biome production (NBP), we can provide a less ambiguous framework for understanding and communicating recent changes in the global C cycle.     

Impact of afforestation-associated management changes on the carbon balance of grassland

Afforestations can be considerable carbon (C) sources due to C losses from the soil after site preparation for tree planting and decreased primary production. In this study, the transition from grassland to afforestation was investigated using two eddy flux towers, which were operated in parallel for 3 years, one on a young afforestation and one on an adjacent grassland. Differences between the fluxes at the two sites were attributable to the management of the sites, without confounding influences of meteorological variability. Site preparation with deep ploughing of the planting rows destroyed 30% of the grassland vegetation at the afforestation site and reduced gross primary productivity by 41% in the first year. At the afforestation site 38 g m^?2 less C was sequestered compared with the nonafforested grassland during the first year. In the following years, the C sink at the afforestation site was higher than at the grassland indicating that soil C loss due to site preparation and land use change on the afforestation occurred only during the first year. Metrological conditions, especially summer drought, caused a high interannual variability of the C balance: both sites were small C sources in 2005 (67 g C m^?2 a^?1 at the grassland and 19 g C g^?1 a^?1 at the afforestation site) and small C sinks in 2004 and 2006 (?72.5 and ?16 g C m^?2 a^?1 at the grassland and ?34 and ?61 g C g^?1 a^?1 at the afforestation). Sheep grazing and mowing affected the short‐term dynamics of the C balance and sheep grazing accelerated the C turnover on the grassland site. The investigated afforestation site did not provide any short‐term way of sequestering additional C even though soil C losses during the first 3 years were relatively small.     

Modeling the effects of snowpack on heterotrophic respiration across northern temperate and high latitude regions: Comparison with measurements of atmospheric carbon dioxide in high latitudes

Simulations by global terrestrial biogeochemical models (TBMs) consistently underestimate the concentration of atmospheric carbon dioxide (CO₂ at high latitude monitoring stations during the non-growing season. We hypothesized that heterotrophic respiration is underestimated during the nongrowing season primarily because TBMs do not generally consider the insulative effects of snowpack on soil temperature. To evaluate this hypothesis, we compared the performance of baseline and modified versions of three TBMs in simulating the seasonal cycle of atmospheric CO₂ at high latitude CO₂ monitoring stations; the modified version maintained soil temperature at 0 °C when modeled snowpack was present. The three TBMs include the Carnegie-Ames-Stanford Approach (CASA), Century, and the Terrestrial Ecosystem Model (TEM). In comparison with the baseline simulation of each model, the snowpack simulations caused higher releases of CO₂ between November and March and greater uptake of CO₂ between June and August for latitudes north of 30° N. We coupled the monthly estimates of CO₂ exchange, the seasonal carbon dioxide flux fields generated by the HAMOCC3 seasonal ocean carbon cycle model, and fossil fuel source fields derived from standard sources to the three-dimensional atmospheric transport model TM2 forced by observed winds to simulate the seasonal cycle of atmospheric CO₂ at each of seven high latitude monitoring stations. In comparison to the CO₂ concentrations simulated with the baseline fluxes of each TBM, concentrations simulated using the snowpack fluxes are generally in better agreement with observed concentrations between August and March at each of the monitoring stations. Thus, representation of the insulative effects of snowpack in TBMs generally improves simulation of atmospheric CO₂ concentrations in high latitudes during both the late growing season and nongrowing season. These simulations highlight the global importance of biogeochemical processes during the nongrowing season in estimating carbon balance of ecosystems in northern high and temperate latitudes.     

Carbon balance in the tundra, boreal forest and humid tropical forest during climate change: scaling up from leaf physiology and soil carbon dynamics

Carbon exchange by the terrestrial biosphere is thought to have changed since pre-industrial times in response to increasing concentrations of atmospheric CO₂ and variations (anomalies) in inter-annual air temperatures. However, the magnitude of this response, particularly that of various ecosystem types (biomes), is uncertain. Terrestrial carbon models can be used to estimate the direction and size of the terrestrial responses expected, providing that these models have a reasonable theoretical base. We formulated a general model of ecosystem carbon fluxes by linking a process-based canopy photosynthesis model to the Rothamsted soil carbon model for biomes that are not significantly affected by water limitation. The difference between net primary production (NPP) and heterotrophic soil respiration (R_h) represents net ecosystem production (NEP). The model includes (i) multiple compartments for carbon storage in vegetation and soil organic matter, (ii) the effects of seasonal changes in environmental parameters on annual NEP, and (iii) the effects of inter-annual temperature variations on annual NEP. Past, present and projected changes in atmospheric CO₂ concentration and surface air temperature (at different latitudes) were analysed for their effects on annual NEP in tundra, boreal forest and humid tropical forest biomes. In all three biomes, annual NEP was predicted to increase with CO₂ concentration but to decrease with warming. As CO₂ concentrations and temperatures rise, the positive carbon gains through increased NPP are often outweighed by losses through increased R_h, particularly at high latitudes where global warming has been (and is expected to be) most severe. We calculated that, several times during the past 140 years, both the tundra and boreal forest biomes have switched between being carbon sources (annual NEP negative) and being carbon sinks (annual NEP positive). Most recently, significant warming at high latitudes during 1988 and 1990 caused the tundra and boreal forests to be net carbon sources. Humid tropical forests generally have been a carbon sink since 1960. These modelled responses of the various biomes are in agreement with other estimates from either field measurements or geochemical models. Under projected CO₂ and temperature increases, the tundra and boreal forests will emit increasingly more carbon to the atmosphere while the humid tropical forest will continue to store carbon. Our analyses also indicate that the relative increase in the seasonal amplitude of the accumulated NEP within a year is about 0–14% year^?1 for boreal forests and 0–23% year^?1 in the tundra between 1960 and 1990.     

A paired study of prairie carbon stocks, fluxes, and phenology: comparing the world's oldest prairie restoration with an adjacent remnant

We measured carbon (C) stocks and fluxes and vegetation phenology in the world's oldest prairie restoration (∼65 years) and an adjacent prairie remnant in southern Wisconsin from 2001–2004 to quantify structural and functional differences. While the species distributions and frequency differed, the number of species measured per 1 m² quadrat were not significantly different (15.8±4.4 and 14.1±2.1 for remnant and planted [order for all reported values in abstract]; P=0.29), and the annual average aboveground net primary productivity (271±51 and 330±55 g C m⁻²) and peak leaf area index (2.9–4.9 m² m⁻²) were comparable under similar fire management. Total root biomass was not significantly different in 2002 (1736±1062 and 1690±459 g dry matter m⁻²) or 2003 (3029±2081 and 2146±898 g m⁻²), but annual average soil respiration (1229±77 and 1428±24 g C m⁻² yr⁻¹) was significantly higher in the restoration (P<0.0001). However, the prairie remnant contained 37% greater soil C (P<0.0001) in the top 25 cm. Soil respiration response to 10 cm soil temperature (Q₁₀) varied with respect to prairie and soil moisture conditions as annual Q₁₀ values ranged from 2.5 to 3.6. We calculated a range of net ecosystem production (NEP) values using estimated heterotrophic respiration and three root turnover values. Average NEP varied from −1.4 to 1.9 and −2.3 to 1.3 Mg C ha⁻¹ yr⁻¹ for the remnant and planted prairies, respectively. While these two prairies share similar structural components and functional attributes, the large uncertainty in NEP casts doubt as to whether we can verify these prairies as C sources or sinks without direct measures of heterotrophic respiration and root turnover. We argue that quantitative studies of C exchange in prairies, which differ in restoration methodology, management intensity, and fire frequency, are needed to solidify the relationship between prairie structure and potentially desired functions such as C sequestration.     

The Carbon Balance of an Old-growth Forest: Building Across Approaches

The carbon budget of the Wind River old-growth forest is being addressed from a variety of perspectives and with a range of approaches. The goal of this comprehensive analysis is developing a thorough, general, and validated understanding of the carbon balance, as well as the processes controlling it. The initial results from studies addressing annual carbon (C) balance with ground-based methods, eddy flux, leaf-based models, and ecosystem models are consistent in some, but not all, respects. Net primary production is 500–600 g C m^–2 y^–1 (5–6 Mg C ha^–1 y^–1), consistent with estimates based on climate alone. The site appears to be close to carbon equilibrium, as a multiyear average, using ground-based methods but a sink of approximately 150–190 g C m^–2 y^–1 from eddy flux for a single year. An overview of the mechanisms that can drive forest carbon sinks illustrates why methods emphasizing different temporal and spatial scales, as well as different processes, can come to different conclusions, and it highlights opportunities in moving toward a truly integrated approach.     

Changes in carbon storage and fluxes in a chronosequence of ponderosa pine

Forest development following stand‐replacing disturbance influences a variety of ecosystem processes including carbon exchange with the atmosphere. On a series of ponderosa pine (Pinius ponderosa var. Laws.) stands ranging from 9 to> 300 years in central Oregon, USA, we used biological measurements to estimate carbon storage in vegetation and soil pools, net primary productivity (NPP) and net ecosystem productivity (NEP) to examine variation with stand age. Measurements were made on plots representing four age classes with three replications: initiation (I, 9–23 years), young (Y, 56–89 years), mature (M, 95–106 years), and old (O, 190–316 years) stands typical of the forest type in the region. Net ecosystem productivity was lowest in the I stands (?124 g C m^?2 yr^?1), moderate in Y stands (118 g C m^?2 yr^?1), highest in M stands (170 g C m^?2 yr^?1), and low in the O stands (35 g C m^?2 yr^?1). Net primary productivity followed similar trends, but did not decline as much in the O stands. The ratio of fine root to foliage carbon was highest in the I stands, which is likely necessary for establishment in the semiarid environment, where forests are subject to drought during the growing season (300–800 mm precipitation per year). Carbon storage in live mass was the highest in the O stands (mean 17.6 kg C m^?2). Total ecosystem carbon storage and the fraction of ecosystem carbon in aboveground wood mass increased rapidly until 150–200 years, and did not decline in older stands. Forest inventory data on 950 ponderosa pine plots in Oregon show that the greatest proportion of plots exist in stands ～ 100 years old, indicating that a majority of stands are approaching maximum carbon storage and net carbon uptake. Our data suggests that NEP averages ～ 70 g C m^?2 year^?1 for ponderosa pine forests in Oregon. About 85% of the total carbon storage in biomass on the survey plots exists in stands greater than 100 years, which has implications for managing forests for carbon sequestration. To investigate variation in carbon storage and fluxes with disturbance, simulation with process models requires a dynamic parameterization for biomass allocation that depends on stand age, and should include a representation of competition between multiple plant functional types for space, water, and nutrients.     

Soil carbon cycling in a temperate forest: radiocarbon-based estimates of residence times, sequestration rates and partitioning of fluxes

Temperate forests of North America are thought to besignificant sinks of atmospheric CO₂. Wedeveloped a below-ground carbon (C) budget forwell-drained soils in Harvard Forest Massachusetts, anecosystem that is storing C. Measurements of carbonand radiocarbon (¹⁴C) inventory were used todetermine the turnover time and maximum rate ofCO₂ production from heterotrophic respiration ofthree fractions of soil organic matter (SOM):recognizable litter fragments (L), humified lowdensity material (H), and high density ormineral-associated organic matter (M). Turnover timesin all fractions increased with soil depth and were2–5 years for recognizable leaf litter, 5–10 years forroot litter, 40–100+ years for low density humifiedmaterial and >100 years for carbon associated withminerals. These turnover times represent the timecarbon resides in the plant + soil system, and mayunderestimate actual decomposition rates if carbonresides for several years in living root, plant orwoody material.Soil respiration was partitioned into two componentsusing ¹⁴C: recent photosynthate which ismetabolized by roots and microorganisms within a yearof initial fixation (Recent-C), and C that is respiredduring microbial decomposition of SOM that resides inthe soil for several years or longer (Reservoir-C).For the whole soil, we calculate that decomposition ofReservoir-C contributes approximately 41% of thetotal annual soil respiration. Of this 41%,recognizable leaf or root detritus accounts for 80%of the flux, and 20% is from the more humifiedfractions that dominate the soil carbon stocks.Measurements of CO₂ and ¹⁴CO₂ in thesoil atmosphere and in total soil respiration werecombined with surface CO₂ fluxes and a soil gasdiffusion model to determine the flux and isotopicsignature of C produced as a function of soil depth. 63% of soil respiration takes place in the top 15 cmof the soil (O + A + Ap horizons). The average residencetime of Reservoir-C in the plant + soil system is8±1 years and the average age of carbon in totalsoil respiration (Recent-C + Reservoir-C) is 4±1years.The O and A horizons have accumulated 4.4 kgC m^–2above the plow layer since abandonment by settlers inthe late-1800's. C pools contributing the most to soilrespiration have short enough turnover times that theyare likely in steady state. However, most C is storedas humified organic matter within both the O and Ahorizons and has turnover times from 40 to 100+ yearsrespectively. These reservoirs continue to accumulatecarbon at a combined rate of 10–30 gC m^{minus 2}yr^–1. This rate of accumulation is only 5–15% of the total ecosystem C sink measured in this stand using eddy covariance methods.