Jack pine foliar δ15N indicates shifts in plant nitrogen acquisition after severe wildfire and through forest stand development

Background and aims

Natural abundance of the stable nitrogen (N) isotope ¹⁵N can elucidate shifts in plant N acquisition and ecosystem N cycling following disturbance events. This study examined the potential relationship between foliar δ¹⁵N and depth of plant N acquisition (surface organic vs. mineral soil) and nitrification as conifer stands develop following stand-replacing wildfire.

Methods

We measured foliar δ¹⁵N along an 18-site chronosequence of jack pine (Pinus banksiana) stands, 1 to 72 years in age post-wildfire. Foliar δ¹⁵N was compared to total δ¹⁵N of the organic (Oe + Oa) and mineral (0–15 cm) soil horizons, and organic horizon N mineralization and nitrification as functions of total mineralization.

Results

Foliar δ¹⁵N declined with stand age, yet wildfire effects were heterogeneous. Jack pine seedlings on burned, mineral soil patches in the youngest stand were significantly more enriched than those on unburned, organic patches (P?=?0.007). High foliar values in the youngest stands relative to mineral-horizon δ¹⁵N indicate that nitrification also likely contributed to seedling enrichment.

Conclusions

Our results suggest jack pine seedlings on burned patches obtain N from the mineral soil with potentially high nitrification rates, whereas seedlings on unburned patches and increasingly N-limited, mature jack pine acquire relatively more N from organic horizons.     

Carbon sequestration in boreal jack pine stands following harvesting

A large area of boreal jack pine (Pinus banksiana Lamb.) forest in Canada is recovering from clear‐cut harvesting, and the carbon (C) balance of these regenerating forests remains uncertain. Net ecosystem CO₂ exchange was measured using the eddy‐covariance technique at four jack pine sites representing different stages of stand development: three postharvest sites (HJP02, HJP94, and HJP75) and one preharvest site (OJP). The four sites, located in the southern Canadian boreal forest, Saskatchewan, Canada, are typical of low productivity jack pine stands and were 2, 10, 29, and 90 years old in 2004, respectively. Mean annual net ecosystem production (NEP) for 2004 and 2005 was ?137±11, 19±16, 73±28, and 22±30 g C m^?2 yr^?1 at HJP02, HJP94, HJP75 and OJP, respectively, showing the postharvest jack pine stands to be moderate C sources immediately after harvesting, weak sinks at 10 years, moderate C sinks at 30 years, then weak C sinks at 90 years. Mean annual gross ecosystem photosynthesis (GEP) for the 2 years was 96±10, 347±20, 576±34, and 583±35 g C m^?2 yr^?1 at HJP02, HJP94, HJP75, and OJP, respectively. The ratio of annual ecosystem respiration (R) to annual GEP was 2.51±0.15, 0.95±0.04, 0.87±0.03, and 0.96±0.03. Seasonally, NEP peaked in May or June at all four sites but GEP and R were highest in July. R at a reference soil temperature of 10 °C, ecosystem quantum yield and photosynthetic capacity were lowest for the 2‐year‐old stand. R was most sensitive to soil temperature for the 90‐year‐old stand. The primary source of variability in NEP over the course of succession of the jack pine ecosystem following harvesting was stand age due to the changes in leaf area index. Intersite variability in GEP and R was an order of magnitude greater than interannual variability at OJP. For both young and old stands, GEP had greater interannual variability than R and played a more important role than R in interannual variation in NEP. Based on year‐round flux measurements from 2000 to 2005, the 10‐year stand had larger interannual variability in GEP and R than the 90‐year stand. Interannual variability in NEP was driven primarily by early‐growing‐season temperature and growing‐season length. Photosynthesis played a dominant role in the rapid rise in NEP early in stand development. Late in stand development, however, the subtle decrease in NEP resulted primarily from increasing respiration.     

Soil CO2 efflux in contrasting boreal deciduous and coniferous stands and its contribution to the ecosystem carbon balance

Similar nonsteady‐state automated chamber systems were used to measure and partition soil CO₂ efflux in contrasting deciduous (trembling aspen) and coniferous (black spruce and jack pine) stands located within 100 km of each other near the southern edge of the Boreal forest in Canada. The stands were exposed to similar climate forcing in 2003, including marked seasonal variations in soil water availability, which provided a unique opportunity to investigate the influence of climate and stand characteristics on soil CO₂ efflux and to quantify its contribution to the net ecosystem CO₂ exchange (NEE) as measured with the eddy‐covariance technique. Partitioning of soil CO₂ efflux between soil respiration (including forest‐floor vegetation) and forest‐floor photosynthesis showed that short‐ and long‐term temporal variations of soil CO₂ efflux were related to the influence of (1) soil temperature and water content on soil respiration and (2) below‐canopy light availability, plant water status and forest‐floor plant species composition on forest‐floor photosynthesis. Overall, the three stands were weak to moderate sinks for CO₂ in 2003 (NEE of ?103, ?80 and ?28 g C m^?2 yr^?1 for aspen, black spruce and jack pine, respectively). Forest‐floor respiration accounted for 86%, 73% and 75% of annual ecosystem respiration, in the three respective stands, while forest‐floor photosynthesis contributed to 11% and 14% of annual gross ecosystem photosynthesis in the black spruce and jack pine stands, respectively. The results emphasize the need to perform concomitant measurements of NEE and soil CO₂ efflux at longer time scales in different ecosystems in order to better understand the impacts of future interannual climate variability and vegetation dynamics associated with climate change on each component of the carbon balance.     

Effectiveness of soil N availability indices in predicting site productivity in the oil sands region of Alberta

Background and aims

Quantitative relationships between soil N availability indices and tree growth are lacking in the oil sands region of Alberta and this can hinder the development of guidelines for the reclamation of the disturbed landscape after oil sands extraction. The aim of this paper was to establish quantitative relationships between soil N availability indices and tree growth in the oil sands region of Alberta.

Methods

In situ N mineralization rates, in situ N availability measured in the field using Plant Root Simulators (PRS? probes), laboratory aerobic and anaerobic soil N mineralization rates, and soil C/N and N content were determined for both the forest floor and the 0–20?cm mineral soil in eight jack pine (Pinus banksiana Lamb.) stands in the oil sands region in northern Alberta. Tree growth rates were determined based on changes in tree ring width in the last 6?years and as mean annual aboveground biomass increment.

Results

Soil N availability indices across those forest stands varied and for each stand it was several times higher in the forest floor than in the mineral soil. The in situ and laboratory aerobic and anaerobic soil N mineralization rates, soil mineralized N, in situ N availability measured using PRS probes, soil C/N ratio and N content in both the forest floor and mineral soil, as well as stand age were linearly correlated with tree ring width of jack pine trees across the selected forest stands, consistent with patterns seen in other published studies and suggesting that N availability could be a limiting factor in the range of jack pine stands studied.

Conclusions

In situ and laboratory aerobic and anaerobic N mineralization rates and soil C/N ratio and N content can be used for predicting tree growth in jack pine forests in the oil sand region. Laboratory based measurements such as aerobic and anaerobic N mineralization rates and soil C/N ratio and N content would be preferable as they are more cost effective and equally effective for predicting jack pine growth.     

Long-Term Nitrogen Additions and Nitrogen Saturation in Two Temperate Forests

This article reports responses of two different forest ecosystems to 9 years (1988–96) of chronic nitrogen (N) additions at the Harvard Forest, Petersham, Massachusetts. Ammonium nitrate (NH₄NO₃) was applied to a pine plantation and a native deciduous broad-leaved (hardwood) forest in six equal monthly doses (May–September) at four rates: control (no fertilizer addition), low N (5 g N m^-2 y^-1), high N (15 g N m^-2 y^-1), and low N + sulfur (5 g N m^-2 y^-1 plus 7.4 g S m^-2 y^-1). Measurements were made of net N mineralization, net nitrification, N retention, wood production, foliar N content and litter production, soil C and N content, and concentrations of dissolved organic carbon (DOC) and nitrogen (DON) in soil water. In the pine stand, nitrate losses were measured after the first year of additions (1989) in the high N plot and increased again in 1995 and 1996. The hardwood stand showed no significant increases in nitrate leaching until 1995 (high N only), with further increases in 1996. Overall N retention efficiency (percentage of added N retained) over the 9-year period was 97–100% in the control and low N plots of both stands, 96% in the hardwood high N plot, and 85% in the pine high N plot. Storage in aboveground biomass, fine roots, and soil extractable pools accounted for only 16–32% of the added N retained in the amended plots, suggesting that the one major unmeasured pool, soil organic matter, contains the remaining 68–84%. Short-term redistribution of ¹⁵N tracer at natural abundance levels showed similar division between plant and soil pools. Direct measurements of changes in total soil C and N pools were inconclusive due to high variation in both stands. Woody biomass production increased in the hardwood high N plot but was significantly reduced in the pine high N plot, relative to controls. A drought-induced increase in foliar litterfall in the pine stand in 1995 is one possible factor leading to a measured increase in N mineralization, nitrification, and nitrate loss in the pine high N plot in 1996. Received 2 April 1999; Accepted 29 October 1999.     

Net ecosystem productivity of boreal jack pine stands regenerating from clearcutting under current and future climates

Life cycle analysis of climate and disturbance effects on forest net ecosystem productivity (NEP) is necessary to assess changes in forest carbon (C) stocks under current or future climates. Ecosystem models used in such assessments need to undergo well-constrained tests of their hypotheses for climate and disturbance effects on the processes that determine CO₂ exchange between forests and the atmosphere. We tested the ability of the model ecosys to simulate diurnal changes in CO₂ fluxes under changing air temperatures (T_a) and soil water contents during forest regeneration with eddy covariance measurements over boreal jack pine (Pinus banksiana) stands along a postclearcut chronosequence. Model hypotheses for hydraulic and nutrient constraints on CO₂ fixation allowed ecosys to simulate the recovery of C cycling during the transition of boreal jack pine stands from C sources following clearcutting (NEP from −150 to −200 g C m⁻² yr⁻¹) to C sinks at maturity (NEP from 20 to 80 g C m⁻² yr⁻¹) with large interannual variability. Over a 126-year logging cycle, annualized NEP, C harvest, and net biome productivity (NBP=NEP–harvest removals) of boreal jack pine averaged 47, 33 and 14 g C m⁻² yr⁻¹. Under an IPCC SRES climate change scenario, rising T_a exacerbated hydraulic constraints that adversely affected NEP of boreal jack pine after 75 years. These adverse effects were avoided in the model by replacing the boreal jack pine ecotype with one adapted to warmer T_a. This replacement raised annualized NEP, C harvest, and NBP to 81, 56 and 25 g C m⁻² yr⁻¹ during a 126-year logging cycle under the same climate change scenario.     

Carbon and Nitrogen Dynamics of Boreal Jack Pine Stands With and Without a Green Alder Understory

We compared the species composition, structure and selected components of the carbon (C) and nitrogen (N) budgets of similar-aged, mature boreal jack pine (Pinus banksiana Lamb.) forests with and without green alder [Alnus crispa (Ait.) Pursh.] in two different boreal environments. The C and N content of the overstory biomass components (for example, stem, branch, and foliage), total vegetation, forest floor, and mineral soil were greater (P= 0.05 to P= 0.10) for jack pine with alder (JPA) stands than for jack pine without alder (JP) stands at both study areas. Jack pine foliage N isotopic discrimination (δ¹⁵N) and annual litterfall N content were significantly greater (P < 0.05) in the JPA than the JP stands at both study areas, suggesting that alder was fixing N and that N availability was greater in the JPA than the JP stands. The greater leaf area index (LAI) and overstory C accumulation in the JPA than the JP stands (P < 0.05) is likely because of the greater N availability in the JPA stands, but the effect of soil texture discontinuity on water availability in the JPA stands can not be dismissed. Percent ground cover by feathermoss varied among the jack pine communities and was positively correlated with overstory LAI (r ²= 0.83, P< 0.05). One index of N-use efficiency (NUE), defined as aboveground net primary productivity (ANPP) per litterfall N, was significantly greater (P < 0.05) for the JP than the JPA stands, but a second index of NUE, ANPP/N uptake, did not differ between the two jack pine communities. Jack pine trees growing without alder produced more organic matter per unit of N, but percent N retranslocation from senescing foliage and N mean residence time in the overstory did not differ between the JPA and the JP stands. A conceptual model is presented that illustrates the potential influence of alder on the species composition, structure, and function of boreal jack pine forests. Received 6 January 1998; accepted 15 April 1998.     

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.     

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.     

Litterfall and fine root biomass contribution to nutrient dynamics in second- and old-growth Douglas-fir ecosystems

Litterfall and fine root production were measured for three years as part of a carbon balance study of three forest stands in the Pacific Northwest of the United States. A young second-growth Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] stand, a second-growth Douglas-fir with red alder (Alnus rubra Bong.) stand, and an old-growth (∼550 years) Douglas-fir stand were monitored for inputs of carbon and nitrogen into the soil from litterfall and fine root production, as well as changes in soil C and N. Fine root production and soil nutrient changes were measured through the use of soil ingrowth bags containing homogenized soil from the respective stands. Litterfall biomass was greatest in the Douglas-fir-alder stand (527 g m⁻² yr⁻¹) that annually returned nearly three times the amount of N as the other stands. Mean residence time for forest floor material was also shortest at this site averaging 4.6 years and 5.5 years for C an N, respectively. Fine root production in the upper 20 cm ranged from 584 g m⁻² in the N rich Douglas-fir-alder stand to 836 g m⁻² in the old-growth stand. Fine root production (down to one meter) was always greater than litterfall with a below:above ratio ranging from 3.73 for the young Douglas-fir stand to 1.62 for the Douglas-fir-alder stand. The below:above N ratios for all three stands closely approximate those for biomass. Soil changes in both C and N differed by site, but the soil C changes in the old-growth stand mirrored those obtained in an ongoing CO₂ flux study. Results from the soil ingrowth bags strongly suggest that this method provides a simple, but sufficient device for measuring potential fine root biomass production as well as soil chemical changes.     

Variation in fine root biomass of three European tree species: Beech (Fagus sylvatica L.), Norway spruce (Picea abies L. Karst.), and Scots pine (Pinus sylvestris L.)

Abstract

Fine roots (<2 mm) are very dynamic and play a key role in forest ecosystem carbon and nutrient cycling and accumulation. We reviewed root biomass data of three main European tree species European beech, (Fagus sylvatica L.), Norway spruce (Picea abies L. Karst.) and Scots pine (Pinus sylvestris L.), in order to identify the differences between species, and within and between vegetation zones, and to show the relationships between root biomass and the climatic, site and stand factors. The collected literature consisted of data from 36 beech, 71 spruce and 43 pine stands. The mean fine root biomass of beech was 389 g m^?2, and that of spruce and pine 297 g m^?2 and 277 g m^?2, respectively. Data from pine stands supported the hypothesis that root biomass is higher in the temperate than in the boreal zone. The results indicated that the root biomass of deciduous trees is higher than that of conifers. The correlations between root biomass and site fertility characteristics seemed to be species specific. There was no correlation between soil acidity and root biomass. Beech fine root biomass decreased with stand age whereas pine root biomass increased with stand age. Fine root biomass at tree level correlated better than stand level root biomass with stand characteristics. The results showed that there exists a strong relationship between the fine root biomass and the above-ground biomass.     

Stand age-related effects on soil respiration in a first rotation Sitka spruce chronosequence in central Ireland

The effect of stand age on soil respiration and its components was studied in a first rotation Sitka spruce chronosequence composed of 10‐, 15‐, 31‐, and 47‐year‐old stands established on wet mineral gley in central Ireland. For each stand age, three forest stands with similar characteristics of soil type and site preparation were used. There were no significant differences in total soil respiration among sites of the same age, except for the case of a 15‐year‐old stand that had lower soil respiration rates due to its higher productivity. Soil respiration initially decreased with stand age, but levelled out in the older stands. The youngest stands had significantly higher respiration rates than more mature sites. Annual soil respiration rates were modelled by means of temperature‐derived functions. The average Q ₁₀ value obtained treating all the stands together was 3.8. Annual soil respiration rates were 991, 686, 556, and 564 g C m^?2 for the 10‐, 15‐, 31‐, and 47‐year‐old stands, respectively. We used the trenching approach to separate soil respiration components. Heterotrophic respiration paralleled soil organic carbon dynamics over the chronosequence, decreasing with stand age to slightly increase in the oldest stand as a result of accumulated aboveground litter and root inputs. Root respiration showed a decreasing trend with stand age, which was explained by a decrease in fine root biomass over the chronosequence, but not by nitrogen concentration of fine roots. The decrease in the relative contribution of autotrophic respiration to total soil CO₂ efflux from 59.3% in the youngest stand to 49.7% in the oldest stand was explained by the higher activity of the root system in younger stands. Our results show that stand age should be considered if simple temperature‐based models to predict annual soil respiration in afforestation sites are to be used.     

重庆缙云山两种林分土壤呼吸对模拟氮沉降的季节响应差异性

氮沉降对土壤呼吸的影响仍然存在着争论,需要进一步研究。选择重庆缙云山的马尾松林和柑橘林开展了氮添加实验,分别设置3个氮添加水平(低氮T_5:20 g N m~(-2)a~(-1),中氮T_(10):40 g N m~(-2)a~(-1)和高氮T_(15):60 g N m~(-2)a~(-1))和对照(T_0:0 g N m~(-2)a~(-1))共4个水平的处理,各林分每个处理各9次重复,每个处理量分4次,在每个季度开始各施1次。采用ACE(Automated Soil CO_2 Exchange Station,UK)自动土壤呼吸监测系统测定两林分土壤表层(0—10 cm)的呼吸、温度和湿度,分别在当年的7月、9月、11月、第2年的1月、2月、3月、5月、6月各连续测定4d,每天(8:00—18:00)4次,以揭示两种林分土壤呼吸对模拟氮沉降的季节动态响应及其差异性。结果表明:(1)柑橘林与马尾松林林下土壤表层呼吸表现出一致的季节变化动态趋势:夏季春季秋季冬季,但柑橘林土壤呼吸显著高于马尾松林(P0.05)。(2)总体上氮沉降抑制了2种林分土壤表层呼吸,而N沉降量大抑制程度越高。只在冬季土壤湿度低的马尾松林下氮沉降促进了土壤呼吸。(3)土壤温度与土壤呼吸有极显著的正相关指数关系(P0.01),而土壤水分与土壤呼吸有显著的二次模型拟合关系,但均受到氮沉降量处理的影响。综合分析表明,在亚热带山区2类森林下的典型案例结果支持氮沉降抑制土壤呼吸的认识。     

Nitrogen dynamics of a boreal black spruce wildfire chronosequence

This study examined the nitrogen (N) dynamics of a black spruce (Picea mariana (Mill.) BSP)-dominated chronosequence in Manitoba, Canada. The seven sites studied each contained separate well- and poorly drained stands, originated from stand-killing wildfires, and were between 3 and 151 years old. Our goals were to (i) measure total N concentration ([N]) of all biomass components and major soil horizons; (ii) compare N content and select vegetation N cycle processes among the stands; and (iii) examine relationships between ecosystem C and N cycling for these stands. Vegetation [N] varied significantly by tissue type, species, soil drainage, and stand age; woody debris [N] increased with decay state and decreased with debris size. Soil [N] declined with horizon depth but did not vary with stand age. Total (live + dead) biomass N content ranged from 18.4 to 99.7 g N m⁻² in the well-drained stands and 37.8–154.6 g N m⁻² in the poorly drained stands. Mean soil N content (380.6 g N m⁻²) was unaffected by stand age. Annual vegetation N requirement (5.9 and 8.4 g N m⁻² yr⁻¹ in the middle-aged well- and poorly drained stands, respectively) was dominated by trees and fine roots in the well-drained stands, and bryophytes in the poorly drained stands. Fraction N retranslocated was significantly higher in deciduous than evergreen tree species, and in older than younger stands. Nitrogen use efficiency (NUE) was significantly lower in bryophytes than in trees, and in deciduous than in evergreen trees. Tree NUE increased with stand age, but overall stand NUE was roughly constant (∼ ∼150 g g⁻¹ N) across the entire chronosequence.     

Soil N mineralization and nitrification in relation to nitrogen solution chemistry in a small forested watershed

Spatial variations in soil processes regulating mineral N losses to streams were studied in a small watershed near Toronto, Ontario. Annual net N mineralization in the 0–8 cm soil was measured in adjacent upland and riparian forest stands using in situ soil incubations from April 1985 to 1987. Mean annual rates of soil N mineralization and nitrification were higher in a maple soil (93.8 and 87.0 kg.ha^–1) than in a pine soil (23.3 and 8.2 kg.ha^–1 ). Very low mean rates of mineralization (3.3 kg.ha^–1) and nitrification (3.4 kg.ha^–1) were found in a riparian hemlock stand. Average NO₃-N concentrations in soil solutions were 0.3–1.0 mg.L^–1 in the maple stand and >0.06mg.L^–1 in the pine stand. Concentrations of NO₃–N in shallow ground water and stream water were 3–4× greater in a maple subwatershed than in a pine subwatershed. Rapid N uptake by vegetation was an important mechanism reducing solution losses of NO₃–N in the maple stand. Low rates of nitrification were mainly responsible for negligible NO₃–N solution losses in the pine stand.     

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.     

Carbon pools in a boreal mixedwood logging chronosequence

Mixedwood forests are an ecologically and economically important forest type in central Canada, but the ecology of these forests is not as well studied as that of single-species dominated stands in the boreal forest. Northern boreal mixedwood forests have only recently been harvested and the effects of harvesting on carbon content in these stands are unknown. We quantified the carbon content and aboveground net primary production (NPP) for four different-aged mixedwood boreal forest stands in northern Manitoba, Canada. The stands included 11-, 18-, and 30-year-old stands that originated from harvesting and a 65-year-old fire-originated stand that typifies the origin of all northern boreal mixed-wood forests that are coming under management. Trees included black spruce (Picea mariana (Mill.) B.S.P.), jack pine (Pinus banksiana Lamb.), balsam poplar (Populus balsamifera L.), and quaking aspen (Populus tremuloides Michx.). Overstory biomass was estimated using species-specific allometric models that generally explained greater than 95% of the observed variation in biomass. Carbon content of the overstory vegetation was greatest in the 65-year-old stand and was 74% larger than the 11-year-old stand and showed a positive relationship with stand age (F_{1, 2}=122.62, P=0.0081 R²=0.99). The slope of mineral soil carbon did not differ significantly among stands (F_{1, 2}=0.39, P=0.5956, R²=0.16). Coarse woody debris carbon content followed a U-shaped pattern among stands. Aboveground NPP differed by 24% between the youngest and oldest stand. Mean annual carbon accumulation and aboveground NPP rates of the mixedwood forests were on average two times greater than nearby relatively pure stands studied during the BOREAS (BOReal Ecosystem Atmospheric Study) project. The trends in the results, along with other field studies, suggest that harvesting does not significantly affect the total soil carbon content. The results of this study suggest that scientists should be cautious about extrapolating results from BOREAS stands to a broader region until more data on other forest types and regions are available.     

Soil respiration in a Mediterranean oak forest at different developmental stages after coppicing

To assess the variation of soil respiration at different forest stages we measured it in a coppiced oak (Quercus cerris L.) chronosequence in central Italy during two campaigns, spanning 2 successive years, in four stands at different stages of the rotation: 1 year (S1), 5 years (S5), 10 years (S10) and 17 years (S17) after coppicing. The contribution of the different components of soil respiration flux (aboveground litter, belowground decomposition soil organic matter and root respiration) was estimated by a paired comparison of manipulative experiments between the recently coppiced stand (S1) and mature stand (S17). Ninety percent of soil respiration values were between 1.7 and 7.8 μmol m^?2 s^?1, with an overall mean (±SD) of 4.0±2.7 μmol m^?2 s^?1. Spatial variation of soil respiration was high (CV=44.9%), with a mean range (i.e. patch size) of 4.8±2.7 m, as estimated from a semivariance analysis. In the absence of limitation by soil moisture, soil respiration was related to soil temperature with the exponential Q₁₀ model (average Q₁₀=2.25). During summer, soil moisture constrained soil respiration and masked its dependence on soil temperature. Soil respiration declined over the years after coppicing. Assuming a linear decline with stand age, we estimated a reduction of 24% over a 20‐year‐rotation cycle. The response of soil respiration to temperature also changed with age of the stands: the Q₁₀ was estimated to decrease from 2.90 in S1 to 2.42 in S17, suggesting that different components or processes may be involved at different developmental stages. The contribution of heterotrophic respiration to total soil respiration flux was relatively larger in the young S1 stand than in the mature S17 stand.     

Soil Organic Carbon and Water Retention after Conversion of Grasslands to Pine Plantations in the Ecuadorian Andes

Tree plantations in the high elevations of the tropics constitute a growing land use, but their effect on ecosystem processes and services is not well known. We examined changes in soil organic carbon (C) and water retention in a chronosequence of Pinus radiata stands planted in páramo grasslands in Cotopaxi province, Ecuador. Water retention at 10, 33, and 1,500 kPa declined with stand age, with soils in the oldest pine stands retaining 39%, 55%, and 63% less water than grassland soils at the three pressures tested. Soil organic C in the 0–10-cm depth also declined with stand age, from 5.0 kg m^–2 in grasslands to 3.5 kg m^–2 in 20–25-year-old pine stands (P < 0.001); at greater depth in the A horizon, C contents decreased from 2.8 to 1.2 kg m^–2 (P = 0.047). There were no significant differences among age classes in the AC and C horizons (P = 0.15 and P = 0.34, respectively), where little or no weathering of the primary material has occurred. Inputs of C may be affected by the significantly higher carbon–nitrogen (C:N) ratio of the litter under older pine stands (P = 0.005), whereas outputs are influenced by substrate quality as well as soil environmental factors. Soil ratios at the 0–10 cm depth were significantly higher in grasslands and young pine stands (P < 0.001), whereas carbon–phosphorous (C:P) ratios at 0–10-cm depth followed a similar but not significant trend. However, there was no significant difference in short-term decomposition rates (P = 0.60) when the soils were incubated under uniform temperature and moisture conditions. In páramo ecosystems, where high soil moisture plays an important role in retarding decomposition and driving high C storage, the loss of water retention after afforestation may be the dominant factor in C loss. These results suggest that soil C buildup and water retention respond rapidly to changes in biota and need to be assessed with regard to implications for C sequestration and watershed management.     

Detritus Production and Soil N Transformations in Old-Growth Eastern Hemlock and Sugar Maple Stands

To examine the linkage between forest cover type, litter inputs, and patterns of net N mineralization versus the turnover of N among soil microbes, we measured both the net and gross rates of N mineralization in replicated, adjacent old-growth eastern hemlock [Tsuga canadensis(L.) Carr.] or sugar maple (Acer saccharum Marsh.) stands in upper Michigan. Mean aboveground net primary production and annual litterfall mass were significantly higher (P < 0.01) in the maple forests (870 g·m^-2·y^-1 and 439 g·m^-2·y^-1, respectively) than in the hemlock forests (480 g·m^-2·y^-1 and 344 g·m^-2·y^-1, respectively). Forest floor and coarse woody debris mass, however, were significantly lower (P < 0.05) in the maple forests (2.2 and 0.1 kg·m^-2, respectively) than in the hemlock forests (2.9 and 0.2 kg·m^-2, respectively). Litterfall N concentration was not significantly different (P > 0.10) between the two forest types. In situ gross rates of N mineralization were higher (P < 0.06) in the maple forests than in the hemlock forests (7.5 and 6.1 mg N·kg soil^-1·d^-1 respectively), but in situ net N mineralization varied independently of forest type and stand-level litterfall N concentration. Cover type–dependent differences in detritus production and detritus C quality appear to result in different N turnover rates, but the balance between gross mineralization and immobilization of N is very sensitive to within stand variability and varies at a scale smaller than cover type alone can predict. Received 3 Feburary 1999; accepted 27 August 1999.