Soil-surface carbon dioxide efflux and microbial biomass in relation to tree density 13 years after a stand replacing fire in a lodgepole pine ecosystem

The effects of fire on soil‐surface carbon dioxide (CO₂) efflux, F_S, and microbial biomass carbon, C_mic, were studied in a wildland setting by examining 13‐year‐old postfire stands of lodgepole pine differing in tree density (< 500 to > 500 000 trees ha^?1) in Yellowstone National Park (YNP). In addition, young stands were compared to mature lodgepole pine stands (～110‐year‐old) in order to estimate ecosystem recovery 13 years after a stand replacing fire. Growing season F_S increased with tree density in young stands (1.0 µmol CO₂ m^?2 s^?1 in low‐density stands, 1.8 µmol CO₂ m^?2 s^?1 in moderate‐density stands and 2.1 µmol CO₂ m^?2 s^?1 in high‐density stands) and with stand age (2.7 µmol CO₂ m^?2 s^?1 in mature stands). Microbial biomass carbon in young stands did not differ with tree density and ranged from 0.2 to 0.5 mg C g^?1 dry soil over the growing season; C_mic was significantly greater in mature stands (0.5–0.8 mg C g^?1 dry soil). Soil‐surface CO₂ efflux in young stands was correlated with biotic variables (above‐ground, below‐ground and microbial biomass), but not with abiotic variables (litter and mineral soil C and N content, bulk density and soil texture). Microbial biomass carbon was correlated with below‐ground plant biomass and not with soil carbon and nitrogen, indicating that plant activity controls not only root respiration, but C_mic pools and overall F_S rates as well. These findings support recent studies that have demonstrated the prevailing importance of plants in controlling rates of F_S and suggest that decomposition of older, recalcitrant soil C pools in this ecosystem is relatively unimportant 13 years after a stand replacing fire. Our results also indicate that realistic predictions and modeling of terrestrial C cycling must account for the variability in tree density and stand age that exists across the landscape as a result of natural disturbances.     

Decline in Net Ecosystem Productivity Following Canopy Transition to Late-Succession Forests

Boreal forests are critical to the global carbon (C) cycle. Despite recent advances in our understanding of boreal C budgets, C dynamics during compositional transition to late-succession forests remain unclear. Using a carefully replicated 203-year chronosequence, we examined long-term patterns of forest C stocks and net ecosystem productivity (NEP) following stand-replacing fire in the boreal forest of central Canada. We measured all C pools, including understorey vegetation, belowground biomass, and soil C, which are often missing from C budgets. We found a slight decrease in total ecosystem C stocks during early stand initiation, between 1 and 8 years after fire, at ?0.90 Mg C ha^?1 y^?1. As stands regenerated, live vegetation biomass increased rapidly, with total ecosystem C stocks reaching a maximum of 287.72 Mg C ha^?1 92 years after fire. Total ecosystem C mass then decreased in the 140- and 203-year-old stands, losing between ?0.50 and ?0.74 Mg C ha^?1 y^?1, contrasting with views that old-growth forests continue to maintain a positive C balance. The C decline corresponded with canopy transition from dominance of Populus tremuloides, Pinus banksiana, and Picea mariana in the 92-year-old stands to Betula papyrifera, Picea glauca, and Abies balsamea in the 203-year-old stands. Results from this study highlight the role of succession in long-term forest C dynamics and its importance when modeling terrestrial C flux.     

Carbon distribution of a well- and poorly-drained black spruce fire chronosequence

The objective of this study was to quantify carbon (C) distribution for boreal black spruce (Picea mariana (Mill.) BSP) stands comprising a fire chronosequence in northern Manitoba, Canada. The experimental design included seven well‐drained (dry) and seven poorly‐drained (wet) stands that burned between 1998 and 1850. Vegetation C pools (above‐ground + below‐ground) steadily increased from 1.3 to 83.3 t C ha^?1 for the dry chronosequence, and from 0.6 to 37.4 t C ha^?1 for the wet chronosequence. The detritus C pools (woody debris + forest floor) varied from 10.3 to 96.0 t C ha^?1 and from 12.6 to 77.4 t C ha^?1 for the dry and wet chronosequence, respectively. Overstorey biomass, mean annual biomass increment (MAI), woody debris mass, and litterfall were significantly greater (α = 0.05) for the dry stands than for the wet stands, but the bryophyte, understorey, and forest floor C pools were significantly less for the dry than for the wet stands. The root mass ratio decreased with stand age until 37 years after fire, was fairly constant thereafter, and was not significantly affected by soil drainage. The C pools of the overstorey and bryophyte tended to increase with stand age. Foliage biomass, litterfall, and MAI (for the dry stands) peaked at 71 years after fire and declined in the oldest stands. The results from this study illustrate that the effects of disturbance and edaphic conditions must be accounted for in boreal forest C inventories and C models. The appropriateness of using chronosequences to examine effects of wildfire on ecosystem C distribution is discussed.     

Carbon storage and fluxes in ponderosa pine forests at different developmental stages

We compared carbon storage and fluxes in young and old ponderosa pine stands in Oregon, including plant and soil storage, net primary productivity, respiration fluxes, eddy flux estimates of net ecosystem exchange (NEE), and Biome‐BGC simulations of fluxes. The young forest (Y site) was previously an old‐growth ponderosa pine forest that had been clearcut in 1978, and the old forest (O site), which has never been logged, consists of two primary age classes (50 and 250 years old). Total ecosystem carbon content (vegetation, detritus and soil) of the O forest was about twice that of the Y site (21 vs. 10 kg C m^?2 ground), and significantly more of the total is stored in living vegetation at the O site (61% vs. 15%). Ecosystem respiration (R_e) was higher at the O site (1014 vs. 835 g C m^?2 year^?1), and it was largely from soils at both sites (77% of R_e). The biological data show that above‐ground net primary productivity (ANPP), NPP and net ecosystem production (NEP) were greater at the O site than the Y site. Monte Carlo estimates of NEP show that the young site is a source of CO₂ to the atmosphere, and is significantly lower than NEP(O) by c. 100 g C m^?2 year^?1. Eddy covariance measurements also show that the O site was a stronger sink for CO₂ than the Y site. Across a 15‐km swath in the region, ANPP ranged from 76 g C m^?2 year^?1 at the Y site to 236 g C m^?2 year^?1 (overall mean 158 ± 14 g C m^?2 year^?1). The lowest ANPP values were for the youngest and oldest stands, but there was a large range of ANPP for mature stands. Carbon, water and nitrogen cycle simulations with the Biome‐BGC model suggest that disturbance type and frequency, time since disturbance, age‐dependent changes in below‐ground allocation, and increasing atmospheric concentration of CO₂ all exert significant control on the net ecosystem exchange of carbon at the two sites. Model estimates of major carbon flux components agree with budget‐based observations to within ± 20%, with larger differences for NEP and for several storage terms. Simulations showed the period of regrowth required to replace carbon lost during and after a stand‐replacing fire (O) or a clearcut (Y) to be between 50 and 100 years. In both cases, simulations showed a shift from net carbon source to net sink (on an annual basis) 10–20 years after disturbance. These results suggest that the net ecosystem production of young stands may be low because heterotrophic respiration, particularly from soils, is higher than the NPP of the regrowth. The amount of carbon stored in long‐term pools (biomass and soils) in addition to short‐term fluxes has important implications for management of forests in the Pacific North‐west for carbon sequestration.     

Recovery of Ecosystem Carbon Stocks in Young Boreal Forests: A Comparison of Harvesting and Wildfire Disturbance

Corresponding with the increasing global resource demand, harvesting now affects millions of hectares of boreal forest each year, and yet our understanding of harvesting impacts on boreal carbon (C) dynamics relative to wildfire remains unclear. We provide a direct comparison of C stocks following clearcut harvesting and fire over a 27-year chronosequence in the boreal forest of central Canada. Whereas many past studies have lacked measurement of all major C pools, we attempt to provide complete C pool coverage, including live biomass, deadwood, forest floor, and mineral soil C pools. The relative contribution of each C pool to total ecosystem C varied considerably between disturbance types. Live biomass C was significantly higher following harvesting compared with fire because of residual live trees and advanced regeneration. Conversely, most live biomass was killed following fire, and thus post-fire stands contained higher stocks of deadwood C. Snag and stump C mass peaked immediately following fire, but dramatically decreased 8 years after fire as dead trees began to fall over, contributing to the downed woody debris C pool. Forest floor C mass was substantially lower shortly after fire than harvesting, but this pool converged 8 years after fire and harvesting. When total ecosystem C stocks were examined, we found no significant difference during early stand development between harvesting and fire. Maximum total ecosystem C occurred at age 27 years, 185.1 ± 18.2 and 163.6 ± 8.0 Mg C ha^?1 for harvesting and fire, respectively. Our results indicate strong differences in individual C pools, but similar total ecosystem C after fire and clearcutting in boreal forests, and shall help improve modeling terrestrial C flux after stand-replacing disturbances.     

Canopy transpiration in a chronosequence of Central Siberian pine forests

Tree transpiration was measured in 28, 67, 204 and 383‐y‐old uniform stands and in a multicohort stand (140–430 y) of Pinus sylvestris ssp. sibirica Lebed. in Central Siberia during August 1995. In addition transpiration of three codominant trees was monitored for two years in a 130‐y‐old stand. All stands established after fire. Leaf area index (LAI) ranged between 0.6 (28‐y‐old stand) and 1.6 for stands older than 67‐y. Stand xylem area at 1.3 m height increased from 4 cm² m^?2 (28‐y) to 11.5 cm² m^?2 (67‐y) and decreased again to 7 cm² m^?2 in old stands. Above‐ground living biomass increased from 1.5 kg dry weight m^?2 (28‐y) to 14 kg dry weight m^?2 (383‐y). Day‐to‐day variation of tree transpiration in summer was dependent on net radiation, vapour pressure deficit, and soil water stress. Tree‐to‐tree variation of xylem flux was small and increased with heterogeneity in canopy structure. Maximum rates of xylem flux density followed the course of net radiation from mid April when a constant level of maximum rates was reached until mid September when low temperatures and light strongly reduced flux density. Maximum sap flux density (60 g m^?2 s^?1) and canopy transpiration (1.5 mm d^?1) were reached in the 67‐y stand. Average canopy transpiration of all age classes was 0.72 ± 0.3 mm d^?1. Canopy transpiration (E) was not correlated with LAI but related to stand sapwood area SA (E = ? 0.02 + 1.15SA R²) which was determined by stand density and tree sapwood area.     

Carbon balance of different aged Scots pine forests in Southern Finland

We estimated annual net ecosystem exchange (NEE) of a chronosequence of four Scots pine stands in southern Finland during years 2000–2002 using eddy covariance (EC). Net ecosystem productivity (NEP) was estimated using growth measurements and modelled mass losses of woody debris. The stands were 4, 12, 40 and 75 years old. The 4‐year‐old clearcut was a source of carbon throughout the year combining a low gross primary productivity (GPP) with a total ecosystem respiration (TER) similar to the forest stands. The annual NEE of the clearcut, measured by EC, was 386 g C m^?2. Tree growth was negligible and the estimated NEP was ?262 g C m^?2 a^?1. The annual GPPs at the other sites were close to each other (928?1072 g C m^?2 a^?1), but TER differed markedly, being greatest at the 12‐year‐old site (905 g C m^?2 a^?1) and smallest in the 75‐year‐old stand (616 g C m^?2 a^?1). Measurements of soil CO₂ efflux showed that different rates of soil respiration largely explained the differences in TER. The NEE and NEP of the 12‐year‐old stand were close to zero. The forested stands were sinks of carbon. They had similar annual patterns of carbon exchange and half‐hourly eddy fluxes were highly correlated, indicating similar responses to the environment. The NEE in the 40‐year‐old stand varied between ?179 and –192 g C m^?2 a^?1, while NEP was between 214 and 242 g C m^?2 a^?1. The annual NEE of the 75‐year‐old stand was 323 g C m^?2 and NEP was 252 g C m^?2. This indicates that there was no reduction in carbon sink strength with stand age.     

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.     

气候变化和不同强度造林对大兴安岭主要树种林分信息和地上生物量的长期影响

气候变化及相应火干扰在不同尺度上影响着我国大兴安岭地区森林动态,且在未来的影响可能继续加剧。为了提高森林生态功能和应对气候变暖,国家在分类经营基础上全面实施抚育采伐和补植造林,效果较好,但抚育采伐对森林主要树种的长期影响知之甚少,其在未来气候下的可持续性也有待进一步评估,同时,探讨造林措施对未来森林的影响也显得尤为重要。本文运用森林景观模型LANDIS PRO,模拟气候变化及火干扰、采伐和造林对大兴安岭地区主要树种的长期影响。结果表明:1)模型初始化、短期和长期模拟结果均得到了有效验证,模拟结果与森林调查数据之间无显著性差异(P0.05),基于火烧迹地数据的林火干扰验证亦能够反映当前火干扰的效果,模型模拟结果的可信度较高;2)与当前气候相比,气候变暖及火干扰明显改变了树种组成、年龄结构和地上生物量,B1气候下研究区森林基本上以针叶树种为主要树种,A2气候下优势树种向阔叶树转变;3)与无采伐预案相比,当前气候下,抚育采伐使落叶松的林分密度和地上生物量分别降低了(165±94.9)株/hm~2和(8.5±5.1) Mg/hm~2,增加了樟子松、白桦和云杉等树木株数和地上生物量(3.3—753.4株/hm~2和0.2—4.0 Mg/hm~2),而对山杨的影响较小;B1和A2气候下抚育采伐显著改变林分密度,降低景观尺度地上生物量,进而表现为不可持续;4)B1气候下,推荐实施中低强度造林预案(10%和20%强度),在A2气候下,各强度造林均可在模拟后期增加树种地上生物量。     

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.     

Recovery of Aboveground Plant Biomass and Productivity After Fire in Mesic and Dry Black Spruce Forests of Interior Alaska

Plant biomass accumulation and productivity are important determinants of ecosystem carbon (C) balance during post-fire succession. In boreal black spruce (Picea mariana) forests near Delta Junction, Alaska, we quantified aboveground plant biomass and net primary productivity (ANPP) for 4 years after a 1999 wildfire in a well-drained (dry) site, and also across a dry and a moderately well-drained (mesic) chronosequence of sites that varied in time since fire (2 to ∼116 years). Four years after fire, total biomass at the 1999 burn site had increased exponentially to 160 ± 21 g m⁻² (mean ± 1SE) and vascular ANPP had recovered to 138 ± 32 g m⁻² y⁻¹, which was not different than that of a nearby unburned stand (160 ± 48 g m⁻² y⁻¹) that had similar pre-fire stand structure and understory composition. Production in the young site was dominated by re-sprouting graminoids, whereas production in the unburned site was dominated by black spruce. On the dry and mesic chronosequences, total biomass pools, including overstory and understory vascular and non-vascular plants, and lichens, increased logarithmically (dry) or linearly (mesic) with increasing site age, reaching a maximum of 2469 ± 180 (dry) and 4008 ± 233 g m⁻² (mesic) in mature stands. Biomass differences were primarily due to higher tree density in the mesic sites because mass per tree was similar between sites. ANPP of vascular and non-vascular plants increased linearly over time in the mesic chronosequence to 335 ± 68 g m⁻² y⁻¹ in the mature site, but in the dry chronosequence it peaked at 410 ± 43 g m⁻² y⁻¹ in a 15-year-old stand dominated by deciduous trees and shrubs. Key factors regulating biomass accumulation and production in these ecosystems appear to be the abundance and composition of re-sprouting species early in succession, the abundance of deciduous trees and shrubs in intermediate aged stands, and the density of black spruce across all stand ages. A better understanding of the controls over these factors will help predict how changes in climate and fire regime will affect the carbon balance of Interior Alaska. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

An Analysis of Vegetation Restoration on Opencast Oil Shale Mines in Estonia

We compared four types of 30‐year‐old forest stands growing on spoil of opencast oil shale mines in Estonia. The stand types were: (1) natural stands formed by spontaneous succession, and plantations of (2) Pinus sylvestris (Scots pine), (3) Betula pendula (silver birch), and (4) Alnus glutinosa (European black alder). In all stands we measured properties of the tree layer (species richness, stand density, and volume of growing stock), understory (density and species richness of shrubs and tree saplings), and ground vegetation (aboveground biomass, species richness, and species diversity). The tree layer was most diverse though sparse in the natural stands. Understory species richness per 100‐m² plot was highest in the natural stand, but total stand richness was equal in the natural and alder stands, which were higher than the birch and pine stands. The understory sapling density was lower than 50 saplings/100 m² in the plantations, while it varied between 50 and 180 saplings/100 m² in the natural stands. Growing stock volume was the least in natural stands and greatest in birch stands. The aboveground biomass of ground vegetation was highest in alder stands and lowest in the pine stands. We can conclude that spontaneous succession promotes establishment of diverse vegetation. In plantations the establishment of diverse ground vegetation depends on planted tree species.     

Net primary production and net ecosystem production of a boreal black spruce wildfire chronosequence

Net primary production (NPP) was measured in seven black spruce (Picea mariana (Mill.) BSP)‐dominated sites comprising a boreal forest chronosequence near Thompson, Man., Canada. The sites burned between 1998 and 1850, and each contained separate well‐ and poorly drained stands. All components of NPP were measured, most for 3 consecutive years. Total NPP was low (50–100 g C m^?2 yr^?1) immediately after fire, highest 12–20 years after fire (332 and 521 g C m^?2 yr^?1 in the dry and wet stands, respectively) but 50% lower than this in the oldest stands. Tree NPP was highest 37 years after fire but 16–39% lower in older stands, and was dominated by deciduous seedlings in the young stands and by black spruce trees (>85%) in the older stands. The chronosequence was unreplicated but these results were consistent with 14 secondary sites sampled across the landscape. Bryophytes comprised a large percentage of aboveground NPP in the poorly drained stands, while belowground NPP was 0–40% of total NPP. Interannual NPP variability was greater in the youngest stands, the poorly drained stands, and for understory and detritus production. Net ecosystem production (NEP), calculated using heterotrophic soil and woody debris respiration data from previous studies in this chronosequence, implied that the youngest stands were moderate C sources (roughly, 100 g C m^?2 yr^?1), the middle‐aged stands relatively strong sinks (100–300 g C m^?2 yr^?1), and the oldest stands about neutral with respect to the atmosphere. The ecosystem approach employed in this study provided realistic estimates of chronosequence NPP and NEP, demonstrated the profound impact of wildfire on forest–atmosphere C exchange, and emphasized the need to account for soil drainage, bryophyte production, and species succession when modeling boreal forest C fluxes.     

Invasive insects impact forest carbon dynamics

Invasive insects can impact ecosystem functioning by altering carbon, nutrient, and hydrologic cycles. In this study, we used eddy covariance to measure net CO₂ exchange with the atmosphere (NEE), and biometric measurements to characterize net ecosystem productivity (NEP) in oak‐ and pine‐dominated forests that were defoliated by Gypsy moth (Lymantria dispar L.) in the New Jersey Pine Barrens. Three years of data were used to compare C dynamics; 2005 with minimal defoliation, 2006 with partial defoliation of the canopy and understory in a mixed stand, and 2007 with complete defoliation of an oak‐dominated stand, and partial defoliation of the mixed and pine‐dominated stands. Previous to defoliation in 2005, annual net CO₂ exchange (NEE_yr) was estimated at ?187, ?137 and ?204 g C m^?2 yr^?1 at the oak‐, mixed‐, and pine‐dominated stands, respectively. Annual NEP estimated from biometric measurements was 108%, 100%, and 98% of NEE_yr in 2005 for the oak‐, mixed‐, and pine‐dominated stands, respectively. Gypsy moth defoliation strongly reduced fluxes in 2006 and 2007 compared with 2005; NEE_yr was ?122, +103, and ?161 g C m^?2 yr^?1 in 2006, and +293, +129, and ?17 g C m^?2 yr^?1 in 2007 at the oak‐, mixed‐, and pine‐dominated stands, respectively. At the landscape scale, Gypsy moths defoliated 20.2% of upland forests in 2007. We calculated that defoliation in these upland forests reduced NEE_yr by 41%, with a 55% reduction in the heavily impacted oak‐dominated stands. ‘Transient’ disturbances such as insect defoliation, nonstand replacing wildfires, and prescribed burns are major factors controlling NEE across this landscape, and when integrated over time, may explain much of the patterning of aboveground biomass and forest floor mass in these upland forests.     

Patterns of NPP,GPP, respiration,and NEP during boreal forest succession

We combined year‐round eddy covariance with biometry and biomass harvests along a chronosequence of boreal forest stands that were 1, 6, 15, 23, 40, ～74, and ～154 years old to understand how ecosystem production and carbon stocks change during recovery from stand‐replacing crown fire. Live biomass (C_live) was low in the 1‐ and 6‐year‐old stands, and increased following a logistic pattern to high levels in the 74‐ and 154‐year‐old stands. Carbon stocks in the forest floor (C_{forest floor}) and coarse woody debris (C_CWD) were comparatively high in the 1‐year‐old stand, reduced in the 6‐ through 40‐year‐old stands, and highest in the 74‐ and 154‐year‐old stands. Total net primary production (TNPP) was reduced in the 1‐ and 6‐year‐old stands, highest in the 23‐ through 74‐year‐old stands and somewhat reduced in the 154‐year‐old stand. The NPP decline at the 154‐year‐old stand was related to increased autotrophic respiration rather than decreased gross primary production (GPP). Net ecosystem production (NEP), calculated by integrated eddy covariance, indicated the 1‐ and 6‐year‐old stands were losing carbon, the 15‐year‐old stand was gaining a small amount of carbon, the 23‐ and 74‐year‐old stands were gaining considerable carbon, and the 40‐ and 154‐year‐old stands were gaining modest amounts of carbon. The recovery from fire was rapid; a linear fit through the NEP observations at the 6‐ and 15‐year‐old stands indicated the transition from carbon source to sink occurred within 11–12 years. The NEP decline at the 154‐year‐old stand appears related to increased losses from C_live by tree mortality and possibly from C_{forest floor} by decomposition. Our findings support the idea that NPP, carbon production efficiency (NPP/GPP), NEP, and carbon storage efficiency (NEP/TNPP) all decrease in old boreal stands.     

高寒沙地乌柳防护林碳库随林龄的变化

植被恢复是改善脆弱生态系统的有效方式。长期的植被恢复能够提高沙地生态系统的服务功能。以青海共和高寒沙地不同林龄乌柳(Salix cheilophila)防护林生态系统为研究对象,研究植被恢复过程中植被碳库与土壤碳库的动态变化,探讨乌柳防护林生态系统的碳汇功能。结果表明:随林龄增加,乌柳各组分碳浓度变化规律并不显著(P0.05),而碳贮量显著增加(P0.05),且不同林龄乌柳各组分碳库的分配比例不同,树干碳贮量占林分碳贮量的百分比最高。各林龄(6、11、16、21a)乌柳林碳贮量分别为4.95、9.93、14.67 t/hm2和21.99 t/hm2。土壤碳库随植被恢复时间的增加而增加,各林龄土壤碳库(0—200cm)分别为9.54、13.03、17.18和19.05 t/hm2。较之6、11a土壤碳库增加26.78%,16a较之11a提高24.16%,21a较16a提高9.82%。地被物层(植被残体)固碳量分别为0.27、0.29、0.33、0.43 t/hm2。不同林龄乌柳林生态系统碳库分别为14.76、23.25、32.18 t/hm2和41.48 t/hm2。各林龄乌柳植被层碳库分别占该林龄总碳库的33.54%、42.71%、45.59%和53.01%,土壤碳库分别占该林龄总碳库的64.63%、56.04%、53.39%和45.93%,而地被物层分别占该林龄总碳库的1.83%、1.25%、1.03%和1.03%。较之恢复前的,各林龄碳库依次增加57.05%、36.52%、27.75%和22.42%。植被恢复各阶段年净碳累积速率分别为1.41、1.70、1.79、1.86 t C hm-2a-1。乌柳防护林生态系统具有"碳汇"功能。     

Above-ground biomass and structure of pristine Siberian Scots pine forests as controlled by competition and fire

The study presents a data set of above-ground biomass (AGB), structure, spacing and fire regime, for 24 stands of pristine Siberian Scots pine (Pinus sylvestris) forests with lichens (n = 20) or Vaccinium/mosses (n = 4) as ground cover, along four chronosequences. The stands of the “lichen” site type (LT) were stratified into three chronosequences according to stand density and fire history. Allometric equations were established from 90 sample trees for stem, coarse branch, fine branch, twig and needle biomass. The LT stands exhibited a low but sustained biomass accumulation until a stand age of 383 years. AGB reached only 6–10 kg_dw m⁻² after 200 years depending on stand density and fire history compared to 20 kg_dw m⁻² in the “Vaccinium” type (VT) stands. Leaf area index (LAI) in the LT stands remained at 0.5–1.5 and crown cover was 30–60%, whereas LAI reached 2.5 and crown cover was >100% in the VT stands. Although nearest-neighbour analyses suggested the existence of density-dependent mortality, fire impact turned out to have a much stronger effect on density dynamics. Fire scar dating and calculation of mean and initial fire return intervals revealed that within the LT stands differences in structure and biomass were related to the severity of fire regimes, which in turn was related to the degree of landscape fragmentation by wetlands. Self-thinning analysis was used to define the local carrying capacity for biomass. A series of undisturbed LT stands was used to characterise the upper self-thinning boundary. Stands that had experienced a moderate fire regime were positioned well below the self-thinning boundary in a distinct fire-thinning band of reduced major axis regression slope −0.26. We discuss how this downward shift resulted from alternating phases of density reduction by fire and subsequent regrowth. We conclude that biomass in Siberian Scots pine forests is strongly influenced by fire and that climate change will affect ecosystem functions predominantly via changes in fire regimes. Received: 2 July 1998 / Accepted: 10 June 1999     

Ecosystem Carbon Remains Low for Three Decades Following Fire and Constrains Soil CO2 Responses to Precipitation in Southwestern Ponderosa Pine Forests

The fire regime of ponderosa pine forests in the southwestern United States has shifted over the past century from historically frequent, low-intensity surface fires to infrequent, stand-replacing crown fires. We quantified plant and soil carbon (C) responses to this new fire regime and assessed interactions between changes in fire regime and changes in precipitation regime predicted by some climate models (specifically, an earlier monsoon rain season). We hypothesized that soil C pools and carbon dioxide (CO₂) efflux rates would decrease initially following stand-replacing fires (due to low plant C inputs and the loss of the soil surficial organic (O) horizon), but then increase with time-after-fire (as plant C inputs increase). Water availability often limits soil biological activity in these forests, but we predicted that low soil C availability following fire would constrain soil CO₂ efflux responses to precipitation. In a series of sites with histories of stand-replacing fires that burned between 2 and 34?years prior to sampling, burned patches had lower soil C pools and fluxes than adjacent unburned patches, but there was no evidence of a trend with time-after-fire. Burned forests had 7,500?g C m^?2 less live plant biomass C (P?<?0.001), 1,600?g C m^?2 less soil total C (P?<?0.001) and 90?g C m^?2 less soil labile C (P?<?0.001) than unburned forests. Lower soil labile C in burned patches was due to both a loss of O horizon mass with fire and lower labile C concentrations (g labile C kg^?1 soil total C) in the mineral soil. During the annual drought that precedes summer monsoon rains, both burned and unburned patches had soil CO₂ efflux rates ranging from 0.9 to 1.1?g CO₂-C m^?2 day^?1. During the monsoon season, soil CO₂ efflux in unburned patches increased to approximately 4.8?g CO₂-C m^?2 day^?1 and rates in paired burned patches (3.4?g CO₂-C m^?2 day^?1) were lower (P?<?0.001). We also used field irrigation to experimentally create an earlier and longer monsoon season, and soil CO₂ efflux rates at both burned and unburned plots increased initially in response to watering, but decreased to below control (plots without irrigation) rates within weeks. Watering did not significantly change cumulative growing season soil CO₂ efflux, supporting our prediction that C availability constrains soil CO₂ efflux responses to precipitation. This research advances our understanding of interactions among climate, fire, and C in southwestern forests, suggesting that climate-induced shifts toward more stand-replacing fires will decrease soil C for decades, such that a single fire can constrain future soil biological responses to precipitation regime changes.     

Methane efflux in relation to plant biomass and sediment characteristics in stands of three common emergent macrophytes in boreal mesoeutrophic lakes

Methane efflux was studied in stands of three emergent macrophyte species (Equisetum fluviatile, Schoenoplectus lacustris and Phragmites australis) commonly found in the littoral zone of boreal lakes. In vegetation stands with relatively low methane (CH₄) emissions (<0.3 mol m^?2 (ice‐free period)^?1), the seasonal variation of CH₄ efflux was better correlated with the dynamics of plant growth than variation in sediment temperature. In dense and productive vegetation stands that released high amounts of CH₄ (2.3–7.7 mol m^?2 (ice‐free period)^?1), the seasonal variation in CH₄ efflux was correlated with sediment temperature, indicating that methanogens were more limited by temperature than substrate supply. The bottom type at the growth site of the emergent plants significantly influenced the ratio of CH₄ efflux to aboveground biomass of plants (Eff : B). The lowest Eff : B ratio was found in E. fluviatile stands growing on sand bottom under experimental conditions and the highest in P. australis‐dominated littoral areas accumulating detritus from external sources. The future changes expected in the hydrology of boreal lakes and rivers because of climatic warming may impact the growth conditions of aquatic macrophytes as well as decomposition and accumulation of detritus and, thus, CH₄ effluxes from boreal lakes.     

Long-term impact of a stand-replacing fire on ecosystem CO2 exchange of a ponderosa pine forest

Ponderosa pine (Pinus ponderosa) forests of the southwestern United States are a mosaic of stands where undisturbed forests are carbon sinks, and stands recovering from wildfires may be sources of carbon to the atmosphere for decades after the fire. However, the relative magnitude of these sinks and sources has never been directly measured in this region, limiting our understanding of the role of fire in regional and US carbon budgets. We used the eddy covariance technique to measure the CO₂ exchange of two forest sites, one burned by fire in 1996, and an unburned forest. The fire was a high‐intensity stand‐replacing burn that killed all trees. Ten years after the fire, the burned site was still a source of CO₂ to the atmosphere [109±6 (SEM) g C m^?2 yr^?1], whereas the unburned site was a sink (?164±23 g C m^?2 yr^?1). The fire reduced total carbon storage and shifted ecosystem carbon allocation from the forest floor and living biomass to necromass. Annual ecosystem respiration was lower at the burned site (480±5 g C m^?2 yr^?1) than at the unburned site (710±54 g C m^?2 yr^?1), but the difference in gross primary production was even larger (372±13 g C m^?2 yr^?1 at the burned site and 858±37 g C m^?2 yr^?1at the unburned site). Water availability controlled carbon flux in the warm season at both sites, and the burned site was a source of carbon in all months, even during the summer, when wet and warm conditions favored respiration more than photosynthesis. Our study shows that carbon losses following stand‐replacing fires in ponderosa pine forests can persist for decades due to slow recovery of the gross primary production. Because fire exclusion is becoming increasingly difficult in dry western forests, a large US forest carbon sink could shift to a decadal‐scale carbon source.