Multimodel simulations of forest harvesting effects on long­term productivity and CN cycling in aspen forests.

The effects of forest management on soil carbon (C) and nitrogen (N) dynamics vary by harvest type and species. We simulated long-term effects of bole-only harvesting of aspen (Populus tremuloides) on stand productivity and interaction of CN cycles with a multiple model approach. Five models, Biome-BGC, CENTURY, FORECAST, LANDIS-II with Century-based soil dynamics, and PnET-CN, were run for 350 yr with seven harvesting events on nutrient-poor, sandy soils representing northwestern Wisconsin, United States. Twenty CN state and flux variables were summarized from the models' outputs and statistically analyzed using ordination and variance analysis methods. The multiple models' averages suggest that bole-only harvest would not significantly affect long-term site productivity of aspen, though declines in soil organic matter and soil N were significant. Along with direct N removal by harvesting, extensive leaching after harvesting before canopy closure was another major cause of N depletion. These five models were notably different in output values of the 20 variables examined, although there were some similarities for certain variables. PnET-CN produced unique results for every variable, and CENTURY showed fewer outliers and similar temporal patterns to the mean of all models. In general, we demonstrated that when there are no site-specific data for fine-scale calibration and evaluation of a single model, the multiple model approach may be a more robust approach for long-term simulations. In addition, multimodeling may also improve the calibration and evaluation of an individual model.

Estimating the carbon budget and maximizing future carbon uptake for a temperate forest region in the U.S.

BACKGROUND: Forests of the Midwest U.S. provide numerous ecosystem services. Two of these, carbon sequestration and wood production, are often portrayed as conflicting. Currently, carbon management and biofuel policies are being developed to reduce atmospheric CO2 and national dependence on foreign oil, and increase carbon storage in ecosystems. However, the biological and industrial forest carbon cycles are rarely studied in a whole-system structure. The forest system carbon balance is the difference between the biological (net ecosystem production) and industrial (net emissions from forest industry) forest carbon cycles, but to date this critical whole system analysis is lacking. This study presents a model of the forest system, uses it to compute the carbon balance, and outlines a methodology to maximize future carbon uptake in a managed forest region. RESULTS: We used a coupled forest ecosystem process and forest products life cycle inventory model for a regional temperate forest in the Midwestern U.S., and found the net system carbon balance for this 615,000 ha forest was positive (2.29 t C ha-1 yr-1). The industrial carbon budget was typically less than 10% of the biological system annually, and averaged averaged 0.082 t C ha-1 yr-1. Net C uptake over the next 100-years increased by 22% or 0.33 t C ha-1 yr-1 relative to the current harvest rate in the study region under the optized harvest regime. CONCLUSIONS: The forest's biological ecosystem current and future carbon uptake capacity is largely determined by forest harvest practices that occurred over a century ago, but we show an optimized harvesting strategy would increase future carbon sequestration, or wood production, by 20-30%, reduce long transportation chain emissions, and maintain many desirable stand structural attributes that are correlated to biodiversity. Our results for this forest region suggest that increasing harvest over the next 100 years increases the strength of the carbon sink, and that carbon sequestration and wood production are not conflicting for this particular forest ecosystem. The optimal harvest strategy found here may not be the same for all forests, but the methodology is applicable anywhere sufficient forest inventory data exist.

Ecosystem warming does not affect photosynthesis or aboveground autotrophic respiration for boreal black spruce.

We measured light-saturated photosynthesis (A(net)), foliage respiration (R(fol)) and stem respiration (R(stem)) of black spruce (Picea mariana (Mill.) B.S.P.) in heated (+5 degrees C) and control plots during 2005, 2006 and 2007 in Thompson, MB, Canada. Large greenhouses and soil-heating cables were used to maintain air and soil temperature 5 degrees C above ambient air and soil temperature. Each greenhouse contained approximately nine black spruce trees and the majority of their fine root mass. Treatments were soil and air warming, soil-only warming, greenhouses maintained at ambient air temperature and control. Gas exchange rates ranged 0.71-4.66, 0.04-0.74 and 0.1-1.0 micromol m(-)(2) s(-)(1) for A(net), R(fol) and R(stem), respectively. Treatment differences for A(net), R(fol) and R(stem) were not significant in any of the 3 years of measurements. The results of this experiment suggest that in a warmer climate, black spruce may not have significant changes in the rate of photosynthesis or respiration.

Fire as the dominant driver of central Canadian boreal forest carbon balance.

Changes in climate, atmospheric carbon dioxide concentration and fire regimes have been occurring for decades in the global boreal forest, with future climate change likely to increase fire frequency--the primary disturbance agent in most boreal forests. Previous attempts to assess quantitatively the effect of changing environmental conditions on the net boreal forest carbon balance have not taken into account the competition between different vegetation types on a large scale. Here we use a process model with three competing vascular and non-vascular vegetation types to examine the effects of climate, carbon dioxide concentrations and fire disturbance on net biome production, net primary production and vegetation dominance in 100 Mha of Canadian boreal forest. We find that the carbon balance of this region was driven by changes in fire disturbance from 1948 to 2005. Climate changes affected the variability, but not the mean, of the landscape carbon balance, with precipitation exerting a more significant effect than temperature. We show that more frequent and larger fires in the late twentieth century resulted in deciduous trees and mosses increasing production at the expense of coniferous trees. Our model did not however exhibit the increases in total forest net primary production that have been inferred from satellite data. We find that poor soil drainage decreased the variability of the landscape carbon balance, which suggests that increased climate and hydrological changes have the potential to affect disproportionately the carbon dynamics of these areas. Overall, we conclude that direct ecophysiological changes resulting from global climate change have not yet been felt in this large boreal region. Variations in the landscape carbon balance and vegetation dominance have so far been driven largely by increases in fire frequency.

Improved simulation of poorly drained forests using Biome-BGC.

Forested wetlands and peatlands are important in boreal and terrestrial biogeochemical cycling, but most general-purpose forest process models are designed and parameterized for upland systems. We describe changes made to Biome-BGC, an ecophysiological process model, that improve its ability to simulate poorly drained forests. Model changes allowed for: (1) lateral water inflow from a surrounding watershed, and variable surface and subsurface drainage; (2) adverse effects of anoxic soil on decomposition and nutrient mineralization; (3) closure of leaf stomata in flooded soils; and (4) growth of nonvascular plants (i.e., bryophytes). Bryophytes were treated as ectohydric broadleaf evergreen plants with zero stomatal conductance, whose cuticular conductance to CO(2) was dependent on plant water content. Individual model changes were parameterized with published data, and ecosystem-level model performance was assessed by comparing simulated output to field data from the northern BOREAS site in Manitoba, Canada. The simulation of the poorly drained forest model exhibited reduced decomposition and vascular plant growth (-90%) compared with that of the well-drained forest model; the integrated bryophyte photosynthetic response accorded well with published data. Simulated net primary production, biomass and soil carbon accumulation broadly agreed with field measurements, although simulated net primary production was higher than observed data in well-drained stands. Simulated net primary production in the poorly drained forest was most sensitive to oxygen restriction on soil processes, and secondarily to stomatal closure in flooded conditions. The modified Biome-BGC remains unable to simulate true wetlands that are subject to prolonged flooding, because it does not track organic soil formation, water table changes, soil redox potential or anaerobic processes.

Estimation of stand-level leaf area for boreal bryophytes.

Bryophytes dominate the carbon and nitrogen cycling of many poorly drained terrestrial ecosystems and are important in the vegetation-atmosphere exchange of carbon and water, yet few studies have estimated their leaf area at the stand scale. This study quantified the bryophyte-specific leaf area (SLA) and leaf area index (LAI) in a group of different-aged boreal forest stands in well and poorly drained soils. Species-specific SLA (for three feather mosses, four Sphagnum spp. and Aulacomnium palustre mixed with Tomentypnum nitens) was assessed by determining the projected area using a flatbed scanner and cross-sectional geometry using a dissecting microscope. The hemisurface leaf area was computed as the product of SLA and live biomass and was scaled by coverage data collected at all stands. Pleurozium schreberi dominated the spatial coverage, biomass and leaf area in the well-drained stands, particularly the oldest, while S. fuscum and A. palustre were important in the poorly drained stands. Live moss biomass ranged from 47 to 230 g m(-2) in the well-drained stands dominated by feather mosses and from 102 to 228 g m(-2) in the poorly drained stands. Bryophyte SLA varied between 135 and 473 cm(2) g(-1), for A. palustre and S. capillifolium, respectively. SLA was strongly and significantly affected by bryophyte species, but did not vary between stands; in general, there was no significant difference between the SLA of non-Sphagnum mosses. Bryophyte LAI increased with stand age, peaking at 3.1 and 3.7 in the well and poorly drained stands, respectively; this represented approximately 40% of the overstory LAI in the well-drained stands and 100-1,000% in the poorly drained stands, underscoring the important role bryophytes play in the water and carbon budgets of these boreal forests.

Reimplementation of the Biome-BGC model to simulate successional change.

Biogeochemical process models are increasingly employed to simulate current and future forest dynamics, but most simulate only a single canopy type. This limitation means that mixed stands, canopy succession and understory dynamics cannot be modeled, severe handicaps in many forests. The goals of this study were to develop a version of Biome-BGC that supported multiple, interacting vegetation types, and to assess its performance and limitations by comparing modeled results to published data from a 150-year boreal black spruce (Picea mariana (Mill.) BSP) chronosequence in northern Manitoba, Canada. Model data structures and logic were modified to support an arbitrary number of interacting vegetation types; an explicit height calculation was necessary to prioritize radiation and precipitation interception. Two vegetation types, evergreen needle-leaf and deciduous broadleaf, were modeled based on site-specific meteorological and physiological data. The new version of Biome-BGC reliably simulated observed changes in leaf area, net primary production and carbon stocks, and should be useful for modeling the dynamics of mixed-species stands and ecological succession. We discuss the strengths and limitations of Biome-BGC for this application, and note areas in which further work is necessary for reliable simulation of boreal biogeochemical cycling at a landscape scale.

Measurement of CO2 exchange between Boreal forest and the atmosphere.

The Boreal forest is the world's second largest forested biome occupying the circumpolar region between 50 degrees N and 70 degrees N. This heterogeneous biome stores about 25% of all terrestrial carbon. We have reviewed EC measurements of CO2 exchange between the atmosphere and Boreal forests, and assessed progress in understanding the controlling processes. We have assessed net ecosystem productivity, the net balance between net primary productivity and heterotrophic respiration, measured using the EC method, for 38 Boreal forest sites. Gross ecosystem productivity has been estimated by adding day-time EC-measured CO2 fluxes to respiration estimated from night-time relationships between respiration and temperature. Maximum midday values of gross ecosystem productivity vary from 33 pmol m(-2) s(-1) for aspen to 6 micromol m(-2) s(-1) for larch stands. Long-term EC flux measurements, ongoing at nine Boreal sites, have shown the strong impact of spring weather and growing season water balance on annual net ecosystem productivity. Estimation of net biome production, incorporating the effects of disturbance resulting from forest fires and logging, has progressed significantly in recent years. After disturbance, summer measurements in Boreal chronosequences suggest that it takes about 10 years before growing season carbon uptake offsets the decomposition emissions. Small-scale exchange rate measurements using chambers and manipulative experiments such as stem girdling and soil heating help to understand the processes and mechanisms playing major roles in the carbon balance of terrestrial ecosystems. Aircraft EC flux measurements, convective boundary layer carbon budgets, and (13)C/12C changes in the atmosphere play an important role in validating estimates of regional carbon exchange based on scaled up EC measurements. Atmospheric inverse models are an important approach to studying regional and global carbon balance but need further improvement to yield reliable quantitative results.

Contribution of root respiration to soil surface CO2 flux in a boreal black spruce chronosequence.

We quantified the contributions of root respiration (RC) and heterotrophic respiration to soil surface CO2 flux (RS) by comparing trenched and untrenched plots in well-drained and poorly drained stands of a black spruce (Picea mariana (Mill.) BSP) fire chronosequence in northern Manitoba, Canada. Our objectives were to: (1) test different equations for modeling RS as a function of soil temperature; and (2) model annual RS and RC for the chronosequence from continuous soil temperature measurements. The choice of equation to model RS strongly affected annual RS and RC, with an Arrhenius-based model giving the best fit to the data, especially at low temperatures. Modeled values of annual RS were positively correlated with soil temperature at 2-cm depth and were affected by year of burn and trenching, but not by soil drainage. During the growing season, measured RC was low in May, peaked in late July and declined to low values by the end of the growing season. Annual RC was < 5% of RS in the recently burned stands, approximately 40% in the 21-year-old stands and 5-15% in the oldest (152-year-old) stands. Evidence suggests that RC may have been underestimated in the oldest stands, with residual root decay from trenching accounting for 5-10% of trenched plot RS at most sites.

Environmental controls on carbon dioxide flux from black spruce coarse woody debris.

Carbon dioxide flux from coarse woody debris (CWD) is an important source of CO2 in forests with moderate to large amounts of CWD. A process-based understanding of environmental controls on CWD CO2 flux (R CWD) is needed to accurately model carbon exchange between forests and the atmosphere. The objectives of this study were to: (1) use a laboratory incubation factorial experiment to quantify the effect of temperature (T CWD), water content (W C), decay status, and their interactions on R CWD for black spruce [Picea mariana (Mill.) BSP] CWD; (2) measure and model spatial and temporal dynamics in T CWD for a boreal black spruce fire chronosequence; and (3) validate the R CWD model with field measurements, and quantify potential errors in estimating annual R CWD from this model on various time steps. The R CWD was positively correlated to T CWD (R 2=0.37, P<0.001) and W C (R 2=0.18, P<0.001), and an empirical R CWD polynomial model that included T CWD and W C interactions explained 74% of the observed variation of R CWD. The R CWD estimates from the R CWD model excellently matched the field measurements. Decay status of CWD significantly (P<0.001) affected R CWD. The temperature coefficient (Q 10) averaged 2.5, but varied by 141% across the 5-42°C temperature range, illustrating the potential shortcomings of using a constant Q 10. The CWD temperature was positively correlated to air temperature (R 2=0.79, P<0.001), with a hysteresis effect that was correlated to CWD decay status and stand leaf area index . Ignoring this temperature hysteresis introduced errors of -1% to +32% in annual R CWD estimates. Increasing T CWD modeling time step from hourly to daily or monthly introduced a 5-11% underestimate in annual R CWD. The annual R CWD values in this study were more than two-fold greater than those in a previous study, illustrating the need to incorporate spatial and temporal responses of R CWD to temperature and water content into models for long-term R CWD estimation in boreal forest ecosystems.

Environmental controls on ground cover species composition and productivity in a boreal black spruce forest.

Boreal black spruce forests typically have a dense ground cover of bryophytes. The two main bryophyte groups in boreal black spruce forests, feathermoss and Sphagnum, have ecophysiological characteristics that influence the biogeochemical cycles of black spruce forests differently. The objective of this study was to examine the environmental controls of ground cover composition and net primary production (NPP) of feathermoss and Sphagnum in a boreal black spruce forest in central Saskatchewan. The fraction of Sphagnum ground cover was positively correlated to canopy photosynthetically active radiation (PAR) transmittance (r 2=0.48, P=0.03), but the fraction of feathermoss ground cover was negatively correlated to canopy PAR transmittance in plots where Sphagnum was present (r 2=0.87, P<0.0001). Sphagnum presence was inversely correlated (P=0.0001) to water table index, defined as water table depth relative to the peat layer, while feathermoss occurred in a wider range of microenvironments. Average NPP for 1998 was more than three times greater for Sphagnum (77 g C m-2 year-1) than feathermoss (24 g C m-2 year-1), but the average bryophyte NPP for 1998 was 25 g C m-2 year-1 because feathermoss was the dominant ground cover. The large, but differing, peat carbon content of Sphagnum- versus feathermoss-dominated boreal forests and peatlands necessitates the need to accurately quantify fraction ground cover. Additional validation of the empirical models between environmental variables and fraction ground cover of bryophytes is necessary, but the reported relationships offer an approach to model carbon dynamics of bryophytes in boreal forests and peatlands.

Characterization of radiation regimes in nonrandom forest canopies: theory, measurements, and a simplified modeling approach.

We used field measurements and Monte Carlo simulations of canopy gap-size distribution and gap fraction to examine how beam radiation interacts with clumped boreal forest canopies of aspen (Populus tremuloides Michx.), black spruce (Picea mariana (Mill.) B.S.P.) and jack pine (Pinus banksiana Lamb.). We demonstrate that the Beer-Lambert law can be modified to accommodate transmission of radiation through a clumped forest canopy as a function of path length or sun zenith angle. Multiband Vegetation Imager (MVI) measurements and Monte Carlo simulations showed that values of the zenith element clumping index (Omega(e)(0)) are typically between 0.4 and 0.5 in jack pine and black spruce and 0.65 in aspen. Estimates of LAI obtained from MVI measurements of the canopy gap fraction and adjusted for canopy clumping and branch architecture yielded LAI values of 3.0 in jack pine, 3.3 in aspen, and about 6.0 in black spruce. These LAI estimates were within 10-25% of direct measurements made at the same sites. Data obtained with the MVI, along with numerical simulations, demonstrated that assumptions of random foliage distributions in boreal forests are invalid and could yield erroneous values of LAI measured by indirect techniques and false characterizations of atmosphere-biosphere interactions. Monte Carlo simulations were used to develop a general equation for beam radiation penetration as a function of zenith angle in clumped canopies. The essential measurements included stem spacing, crown diameter, crown depth, and within-crown gap fraction.

Foliar carbon isotope discrimination in Larix species and sympatric evergreen conifers: a global comparison.

Larches (Larix spp.), deciduous conifers, occur in the northern hemisphere in cold-temperate and boreal climates - an environment normally thought to favor evergreen tree species. We compare foliar carbon isotope discrimination (Δ), instantaneous water use efficiency, total foliar nitrogen concentration, and specific leaf area (for a subset of sites) between Larix spp. and co-occurring evergreen conifers at 20 sites throughout the natural range of larches. Except for Larix occidentalis in the xeric Intermountain West, USA, Δ is significantly (P < 0.05) greater for larches than co-occurring evergreen conifers at 77% of the sites, suggesting that larches use water less efficiently. At elevations greater than 3000 m, the Δ of Larix spp. and co-occurring conifers converge, suggesting that water is not the limiting resource. Foliar nitrogen concentration and specific leaf area are two ecophysiological characteristics that are positively correlated with high photosynthetic capacity. Foliar nitrogen concentration is significantly greater for larches than evergreen conifers at 88% of the sites and specific leaf area is approximately three times greater for larches than co-occurring conifers. Future studies should examine the potential effect that global warming may have on the distribution of larch forests because the water use efficiency of larches is commonly less than co-occurring evergreen conifers and the boreal and high-latitude environments are likely to experience the greatest climate warming.

Root mass, net primary production and turnover in aspen, jack pine and black spruce forests in Saskatchewan and Manitoba, Canada.

Root biomass, net primary production and turnover were studied in aspen, jack pine and black spruce forests in two contrasting climates. The climate of the Southern Study Area (SSA) near Prince Albert, Saskatchewan is warmer and drier in the summer and milder in the winter than the Northern Study Area (NSA) near Thompson, Manitoba, Canada. Ingrowth soil cores and minirhizotrons were used to quantify fine root net primary production (NPPFR). Average daily fine root growth (m m(-2) day(-1)) was positively correlated with soil temperature at 10-cm depth (r(2) = 0.83-0.93) for all three species, with black spruce showing the strongest temperature effect. At both study areas, fine root biomass (measured from soil cores) and fine root length (measured from minirhizotrons) were less for jack pine than for the other two species. Except for the aspen stands, estimates of NPPFR from minirhizotrons were significantly greater than estimates from ingrowth cores. The core method underestimated NPPFR because it does not account for simultaneous fine root growth and mortality. Minirhizotron NPPFR estimates ranged from 59 g m(-2) year(-1) for aspen stands at SSA to 235 g m(-2) year(-1) for black spruce at NSA. The ratio of NPPFR to total detritus production (aboveground litterfall + NPPFR) was greater for evergreen forests than for deciduous forests, suggesting that carbon allocation patterns differ between boreal evergreen and deciduous forests. In all stands, NPPFR consistently exceeded annual fine root turnover and the differences were larger for stands in the NSA than for stands in the SSA, whereas the difference between study areas was only significant for black spruce. The imbalance between NPPFR and fine root turnover is sufficient to explain the net accumulation of carbon in boreal forest soils.

Woody tissue maintenance respiration of four conifers in contrasting climates.

We estimate maintenance respiration for boles of four temperate conifers (ponderosa pine, western hemlock, red pine, and slash pine) from CO2 efflux measurements in autumn, when construction respiration is low or negligible. Maintenance respiration of stems was linearly related to sapwood volume for all species; at 10°C, respiration per unit sapwood volume ranged from 4.8 to 8.3 µmol CO2 m-3 s-1. For all sites combined, respiration increased exponentially with temperature (Q 10 =1.7, r 2=0.78). We estimate that maintenance respiration of aboveground woody tissues of these conifers consumes 52-162 g C m-2 y-1, or 5-13% of net daytime carbon assimilation annually. The fraction of annual net daytime carbon fixation used for stem maintenance respiration increased linearly with the average annual temperature of the site.

FOREST-BGC, A general model of forest ecosystem processes for regional applications. II. Dynamic carbon allocation and nitrogen budgets.

A new version of the ecosystem process model FOREST-BGC is presented that uses stand water and nitrogen limitations to alter the leaf/root/stem carbon allocation fraction dynamically at each annual iteration. Water deficit is defined by integrating a daily soil water deficit fraction annually. Current nitrogen limitation is defined relative to a hypothetical optimum foliar N pool, computed as maximum leaf area index multiplied by maximum leaf nitrogen concentration. Decreasing availability of water or nitrogen, or both, reduces the leaf/root carbon partitioning ratio. Leaf and root N concentrations, and maximum leaf photosynthetic capacity are also redefined annually as functions of nitrogen availability. Test simulations for hypothetical coniferous forests were performed for Madison, WI and Missoula, MT, and showed simulated leaf area index ranging from 4.5 for a control stand at Missoula, to 11 for a fertilized stand at Madison, with Year 50 stem carbon biomasses of 31 and 128 Mg ha(-1), respectively. Total nitrogen incorporated into new tissue ranged from 34 kg ha(-1) year(-1) for the unfertilized Missoula stand, to 109 kg ha(-1) year(-1) for the fertilized Madison stand. The model successfully showed dynamic annual carbon partitioning controlled by water and nitrogen limitations.

Estimation of leaf area index in fourteen southern Wisconsin forest stands using a portable radiometer.

Projected leaf area index (LAI) and Beer-Lambert Law extinction coefficients (K) were estimated for 28-year-old Picea abies (L.) Karst., Larix decidua Mill., Pinus resinosa Ait., and Pinus strobus L. plantations using vertical profile data obtained with a portable integrating radiometer (sunfleck ceptometer). Predicted LAI values were compared with direct measures of LAI. Based on dimensional analysis, LAI ranged from 5.0 for Larix decidua to 10.5 for Picea abies. Significant inverse relationships between cumulative LAI and canopy transmitted radiation were observed for the four species (R(2) = 0.92-0.97). Beer-Lambert extinction coefficients ranged from 0.39 for Picea abies to 0.84 for Pinus strobus. Stand-level predictions of LAI based on the Beer-Lambert Law were compared with measured LAI values for eight conifer and six broadleaf stands. Using local K estimates resulted in predicted LAI values with an average 6% error. Using published K values resulted in an average error of 20%. High LAI and concomitantly low light levels below the canopy of Picea abies stands resulted in large overestimation errors in predicted LAI, rendering the sunfleck ceptometer inappropriate for forests with large LAIs.

Effects of nutrient amendments on fine root biomass in a primary successional forest in Hawai'i.

We determined the effect of fertilization treatments (control (C), complete nutrient amendment without nitrogen (PT), nitrogen only (N) and a complete nutrient amendment (NPT)) on fine root biomass in a tropical montane forest in Hawai'i. Fertilization significantly decreased root biomass; live fine root mass (<2 mm diameter) for the C, PT, N and NPT treatments were 335, 145, 110 and 105 g·m-2, respectively. Nutrient availability appears to control fine root mass in this primary successional forest.