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.     

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.     

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.     

Carbon Accumulation Patterns During Post-Fire Succession in Cajander Larch (Larix cajanderi) Forests of Siberia

Increased fire activity within boreal forests could affect global terrestrial carbon (C) stocks by decreasing stand age or altering tree recruitment, leading to patterns of forest regrowth that differ from those of pre-fire stands. To improve our understanding of post-fire C accumulation patterns within boreal forests, we evaluated above- and belowground C pools within 17 Cajander larch (Larix cajanderi) stands of northeastern Siberia that varied in both years since fire and stand density. Early-successional stands (<20-year old) exhibited low larch recruitment, and consequently, low density, aboveground larch biomass, and aboveground net primary productivity (ANPP_tree). Mid-successional stands (21- to 70-year old) were even-aged with considerable variability in stand density. High-density mid-successional stands had 21 times faster rates of ANPP_tree than low-density stands (252 vs. 12?g?C?m^?2?y^?1) and 26 times more C in aboveground larch biomass (2,186 vs. 85?g?C?m^?2). Density had little effect on total soil C pools. During late-succession (>70-year old), aboveground larch biomass, ANPP_tree, and soil organic layer C pools increased with stand age. These stands were low density and multi-aged, containing both mature trees and new recruits. The rapid accumulation of aboveground larch biomass in high-density, mid-successional stands allowed them to obtain C stocks similar to those in much older low-density stands (~8,000?g?C?m^?2). If fire frequency increases without altering stand density, landscape-level C storage could decline, but if larch density also increases, large aboveground C pools within high-density stands could compensate for a shorter successional cycle.     

Modeling above‐ground litterfall in eastern Mediterranean conifer forests using fractional tree cover,and remotely sensed and ground data

Question: How can we model above‐ground litterfall in Mediterranean conifer forests using remotely sensed and ground data, and geographic information systems (GIS)? Location: Eastern Mediterranean conifer forest of Turkey. Methods: Above‐ground litterfall from Mediterranean forest stands of Pinus nigra, Cedrus libani, Pinus brutia and Juniperus excelsa and mixed Abies cilicica, C. libani and P. nigra was modeled as a function of fractional tree cover using a regression tree algorithm, based on IKONOS and Landsat TM/ETM+data. Landsat TM/ETM+images for the study area were used to map actual stand patterns, based on a land‐cover map of species stands using a supervised classification. Results: Total amount of annual above‐ground litterfall for the entire study area (12 260 km²) was estimated at 417.2 Mg ha^?1 for P. brutia, 291.1 Mg ha^?1 for the mixed stand, 115.5 Mg ha^?1 for P. nigra, 54.6 Mg ha^?1 for J. excelsa and 45.9 Mg ha^?1 for C. libani. The maps generated indicate the distribution of the seasonal amount of total above‐ground litterfall for different species and the distribution of species stands in the study area. There was an increase in the amount of above‐ground litterfall for P. brutia stand in summer, for J. excelsa in autumn and for C. libani, P. nigra and the mixed stand of A. cilicica, P. nigra and C. libani in winter. Conclusion: Application of this model helps to improve the accuracy of estimated litterfall input to soil organic carbon pools in the Mediterranean conifer forests.     

Fine Root Biomass,Production, Turnover Rates,and Nutrient Contents in Boreal Forest Ecosystems in Relation to Species,Climate, Fertility,and Stand Age: Literature Review and Meta-Analyses

Fine roots <2 mm in diameter play a key role in regulating the biogeochemical cycles of ecosystems and are important to our understanding of ecosystem responses to global climate changes. Given the sensitivity of fine roots, especially in boreal region, to climate changes, it is important to assess whether and to what extent fine roots in this region change with climates. Here, in this synthesis, a data set of 218 root studies were complied to examine fine root patterns in the boreal forest in relation to site and climatic factors. The mean fine root biomass in the boreal forest was 5.28 Mg ha^?1, and the production of fine roots was 2.82 Mg ha^?1 yr^?1, accounting for 32% of annual net primary production of the boreal forest. Fine roots in the boreal forest on average turned over 1.07 times per year. Fine roots contained 50.9 kg ha^?1 of nitrogen (N) and 3.63 kg ha^?1 of phosphorous (P). In total, fine roots in the boreal forest ecosystems contain 6.1 × 10⁷ Mg N and 4.4×10⁶Mg P pools, respectively, about 10% of the global nutrients of fine roots. Fine root biomass, production, and turnover rate generally increased with increasing mean annual temperature and precipitation. Fine root biomass in the boreal forest decreased significantly with soil N and P availability. With increasing stand age, fine root biomass increased until about 100 years old for forest stands and then leveled off or decreased thereafter. These results of meta analysis suggest that environmental factors strongly influence fine root biomass, production, and turnover in boreal forest, and future studies should place a particular emphasis on the root-environment relationships.     

Pyrogenic organic matter production from wildfires: a missing sink in the global carbon cycle

Wildfires release substantial quantities of carbon (C) into the atmosphere but they also convert part of the burnt biomass into pyrogenic organic matter (PyOM). This is richer in C and, overall, more resistant to environmental degradation than the original biomass, and, therefore, PyOM production is an efficient mechanism for C sequestration. The magnitude of this C sink, however, remains poorly quantified, and current production estimates, which suggest that ~1‐5% of the C affected by fire is converted to PyOM, are based on incomplete inventories. Here, we quantify, for the first time, the complete range of PyOM components found in‐situ immediately after a typical boreal forest fire. We utilized an experimental high‐intensity crown fire in a jack pine forest (Pinus banksiana) and carried out a detailed pre‐ and postfire inventory and quantification of all fuel components, and the PyOM (i.e., all visually charred, blackened materials) produced in each of them. Our results show that, overall, 27.6% of the C affected by fire was retained in PyOM (4.8 ± 0.8 t C ha^?1), rather than emitted to the atmosphere (12.6 ± 4.5 t C ha^?1). The conversion rates varied substantially between fuel components. For down wood and bark, over half of the C affected was converted to PyOM, whereas for forest floor it was only one quarter, and less than a tenth for needles. If the overall conversion rate found here were applicable to boreal wildfire in general, it would translate into a PyOM production of ~100 Tg C yr^?1 by wildfire in the global boreal regions, more than five times the amount estimated previously. Our findings suggest that PyOM production from boreal wildfires, and potentially also from other fire‐prone ecosystems, may have been underestimated and that its quantitative importance as a C sink warrants its inclusion in the global C budget estimates.     

The influence of fire on carbon distribution and net primary production of boreal Larix gmelinii forests in north-eastern China

The boreal larch forest of Eurasia is a widespread forest ecosystem and plays an important role in the carbon budget of boreal forests. However, few carbon budgets exist for these forests, and the effects of wildfire, the dominant natural disturbance in this region, on carbon budgets are poorly understood. The objective of this study was to quantify the effects of wildfire on carbon distribution and net primary production (NPP) for three major Dahurian larch (Larix gmelinii Rupr.) forest ecosystems in Tahe, Daxing'anling, north‐eastern China: Larix gmelinii–Ledum palustre, Larix gmelinii–grass and Larix gmelinii–Rhododendron dahurica forests. The experimental design included mature forests (unburned), and lightly and heavily burned forests from the 1.3‐million‐ha 1987 wildfire. We measured carbon distribution and above‐ground NPP, and estimated fine root production from literature values. Total ecosystem carbon content for the mature forests was greatest for Larix–Ledum forests (251.4 t C ha^?1) and smallest for Larix–grass forests (123.8 t C ha^?1). Larix–Ledum forests contained the smallest vegetation carbon (13.5%), while Larix–Rhododendron contained the largest vegetation carbon (63.1%). Fires tended to transfer carbon from vegetation to detritus and soil. Total NPP did not differ significantly between the lightly burned and unburned stands, and averaged 1.58, 1.29 and 1.01 t C ha^?1 year^?1 for Larix–grass, Larix–Rhododendron and Larix–Ledum lightly burned stands, respectively. Above‐ground net primary production (ANPP) of heavily burned stands was 92–95% less than unburned and lightly burned stands. The estimated carbon loss during the 1987 fire showed substantial variability among forest types and fire severity levels. Depending upon the assumptions made about the fraction of the landscape occupied by the three larch forest types, the 1987 conflagration in north‐east China released 2.5 × 10⁷?4.9 × 10⁷ t C to the atmosphere. This study illustrates the need to distinguish between the different larch forests for developing general carbon budgets.     

The legacy of harvest and fire on ecosystem carbon storage in a north temperate forest

Forest harvesting and wildfire were widespread in the upper Great Lakes region of North America during the early 20th century. We examined how long this legacy of disturbance constrains forest carbon (C) storage rates by quantifying C pools and fluxes after harvest and fire in a mixed deciduous forest chronosequence in northern lower Michigan, USA. Study plots ranged in age from 6 to 68 years and were created following experimental clear‐cut harvesting and fire disturbance. Annual C storage was estimated biometrically from measurements of wood, leaf, fine root, and woody debris mass, mass losses to herbivory, soil C content, and soil respiration. Maximum annual C storage in stands that were disturbed by harvest and fire twice was 26% less than a reference stand receiving the same disturbance only once. The mechanism for this reduction in annual C storage was a long‐lasting decrease in site quality that endured over the 62‐year timeframe examined. However, during regrowth the harvested and burned forest rapidly became a net C sink, storing 0.53 Mg C ha⁻¹ yr⁻¹ after 6 years. Maximum net ecosystem production (1.35 Mg C ha⁻¹ yr⁻¹) and annual C increment (0.95 Mg C ha⁻¹ yr⁻¹) were recorded in the 24‐ and 50‐year‐old stands, respectively. Net primary production averaged 5.19 Mg C ha⁻¹ yr⁻¹ in experimental stands, increasing by < 10% from 6 to 50 years. Soil heterotrophic respiration was more variable across stand ages, ranging from 3.85 Mg C ha⁻¹ yr⁻¹ in the 6‐year‐old stand to 4.56 Mg C ha⁻¹ yr⁻¹ in the 68‐year‐old stand. These results suggest that harvesting and fire disturbances broadly distributed across the region decades ago caused changes in site quality and successional status that continue to limit forest C storage rates.     

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.     

Quantifying above‐ and belowground biomass carbon loss with forest conversion in tropical lowlands of Sumatra (Indonesia)

Natural forests in South‐East Asia have been extensively converted into other land‐use systems in the past decades and still show high deforestation rates. Historically, lowland forests have been converted into rubber forests, but more recently, the dominant conversion is into oil palm plantations. While it is expected that the large‐scale conversion has strong effects on the carbon cycle, detailed studies quantifying carbon pools and total net primary production (NPP_total) in above‐ and belowground tree biomass in land‐use systems replacing rainforest (incl. oil palm plantations) are rare so far. We measured above‐ and belowground carbon pools in tree biomass together with NPP_total in natural old‐growth forests, ‘jungle rubber’ agroforests under natural tree cover, and rubber and oil palm monocultures in Sumatra. In total, 32 stands (eight plot replicates per land‐use system) were studied in two different regions. Total tree biomass in the natural forest (mean: 384 Mg ha^?1) was more than two times higher than in jungle rubber stands (147 Mg ha^?1) and >four times higher than in monoculture rubber and oil palm plantations (78 and 50 Mg ha^?1). NPP_total was higher in the natural forest (24 Mg ha^?1 yr^?1) than in the rubber systems (20 and 15 Mg ha^?1 yr^?1), but was highest in the oil palm system (33 Mg ha^?1 yr^?1) due to very high fruit production (15–20 Mg ha^?1 yr^?1). NPP_total was dominated in all systems by aboveground production, but belowground productivity was significantly higher in the natural forest and jungle rubber than in plantations. We conclude that conversion of natural lowland forest into different agricultural systems leads to a strong reduction not only in the biomass carbon pool (up to 166 Mg C ha^?1) but also in carbon sequestration as carbon residence time (i.e. biomass‐C:NPP‐C) was 3–10 times higher in the natural forest than in rubber and oil palm plantations.     

Biomass and nutrient distribution in a 15.5 year old forest growing on a rehabilitated bauxite mine

Abstract Above-ground biomass and its nutrient content were measured in a 15.5 year old rehabilitated bauxite mine in the jarrah forest of southwest Australia. The litterfall and its nutrient content was also measured on the same site, 4–6 years and 8–10 years after rehabilitation. The total mass of the vegetation and litter was 103t ha^?1, 25% of that in a nearby jarrah forest. The mass of the trees was 58t ha^?1, approximately 22% of that in the jarrah forest. The amount of nitrogen in the soil, in the litter and in the understorey, and in the total system, exceeded that in the nearby jarrah forest. Annual litterfall in the rehabilitated area was comparable to that measured in the jarrah forest and other dry sclerophyll forests. The nitrogen content of the litterfall in the rehabilitated area was around three to four times that in other dry sclerophyll forests.     

Litterfall of epiphytic macrolichens in Nothofagus forests of northern Patagonia,Argentina: Relation to stand age and precipitation

Abstract: The objective of this study was to analyse how stand age and precipitation influence abundance and diversity of epiphytic macrolichens in southern beech Nothofagus forests, estimated by lichen litter sampling. Five sites of Nothofagus dombeyi (Mirbel) Oersted were selected in Nahuel Huapi National Park, Argentina. At each site, lichen fragments from the forest floor were collected at 12.5 m² plots in pairs of young and mature N. dombeyi forest. Additionally, two sites with multi‐aged subalpine Nothofagus pumilio (Poepp. et Endl.) Krasser forest were investigated in a similar manner. Average litterfall biomass per stand varied from less than 1 kg ha^?1 in a young low‐precipitation stand to a maximum of 20 kg ha^?1 in a mature high‐precipitation stand. In places with higher precipitation, litterfall biomass in N. dombeyi forest was considerably higher in old stands as compared with young ones. In places with less than 2000 mm of precipitation, differences in biomass were less pronounced. Old humid stands contained about twice as many taxa in the litter as old low‐precipitation stands and young stands in general. Mature stands in low‐precipitation sites only contained 17% of the litter biomass as compared with mature stands in high‐precipitation sites. Epiphytic lichen composition changed from predominating fruticose lichens (Usnea spp. and Protousnea spp.) in low‐precipitation stands to Pseudocyphellaria spp., Nephroma spp. and other foliose lichens, in the high‐precipitation stands. There were no clear differences in the proportion of fruticose and foliose lichens between young and old stands. Fruticose lichens dominated litter biomass in both N. pumilio sites.     

The Contribution of Litterfall to Net Primary Production During Secondary Succession in the Boreal Forest

Litterfall is a fundamental process in the nutrient cycle of forest ecosystems and a major component of annual net primary production (NPP). Despite its importance for understanding ecosystem energetics and carbon accounting, the dynamics of litterfall production following disturbance and throughout succession remain poorly understood in boreal forest ecosystems. Using a replicated chronosequence spanning 209 years following fire and 33 years following logging in Ontario, Canada, we examined the dynamics of litterfall production associated with stand development, overstory composition type (broadleaf, mixedwood, and conifer), and disturbance origin. We found that total annual litterfall production increased with stand age following fire and logging, plateauing in post-fire stands approximately 98 years after fire. Neither total annual litterfall production nor any of its constituents differed between young fire- or logging-originated stands. Litterfall production was generally higher in broadleaf stands compared with mixedwood and conifer stands, but varied seasonally, with foliar litterfall highest in broadleaf stands in autumn, and epiphytic lichen litterfall highest in conifer stands in spring. Contrary to previous assumptions, we found that the contribution of litterfall production to net primary production increased with stand age, highlighting the need for modeling studies of net primary productivity to account for the effects of stand age on litterfall dynamics.     

Allocation of carbon in a mature eucalypt forest and some effects of soil phosphorus availability

Pools and annual fluxes of carbon (C) were estimated for a mature Eucalyptus pauciflora (snowgum) forest with and without phosphorus (P) fertilizer addition to determine the effect of soil P availability on allocation of C in the stand. Aboveground biomass was estimated from allometric equations relating stem and branch diameters of individual trees to their biomass. Biomass production was calculated from annual increments in tree diameters and measurements of litterfall. Maintenance and construction respiration were calculated for each component using equations given by Ryan (1991a). Total belowground C flux was estimated from measurements of annual soil CO₂ efflux less the C content of annual litterfall (assuming forest floor and soil C were at approximate steady state for the year that soil CO₂ efflux was measured). The total C content of the standing biomass of the unfertilized stand was 138 t ha^-1, with approximately 80% aboveground and 20% belowground. Forest floor C was 8.5 t ha^-1. Soil C content (0–1 m) was 369 t ha^-1 representing 70% of the total C pool in the ecosystem. Total gross annual C flux aboveground (biomass increment plus litterfall plus respiration) was 11.9 t ha^-1 and gross flux belowground (coarse root increment plus fine root production plus root respiration) was 5.1 t ha^-1. Total annual soil efflux was 7.1 t ha^-1, of which 2.5 t ha^-1 (35%) was contributed by litter decomposition.The short-term effect of changing the availability of P compared with C on allocation to aboveground versus belowground processes was estimated by comparing fertilized and unfertilized stands during the year after treatment. In the P-fertilized stand annual wood biomass increment increased by 30%, there was no evidence of change in canopy biomass, and belowground C allocation decreased by 19% relative to the unfertilized stand. Total annual C flux was 16.97 and 16.75 t ha^-1 yr^-1 and the ratio of below- to aboveground C allocation was 0.43 and 0.35 in the unfertilized and P-fertilized stands, respectively. Therefore, the major response of the forest stand to increased soil P availability appeared to be a shift in C allocation; with little change in total productivity. These results emphasise that both growth rate and allocation need to be estimated to predict changes in fluxes and storage of C in forests that may occur in response to disturbance or climate change.     

Secondary forest growth deviation from chronosequence predictions in central Amazonia

Nearly all published rates of secondary forest (SF) regrowth for Amazonia are inferred from chronosequences. We examined SF regrowth on abandoned pastures over a 4‐year period to determine if measured rates of forest recovery differ from chronosequence predictions. We studied the emergence, development and death of over 1300 stems in 10 SFs representing three age classes (<1–5, 6–10 and 11–14 years old). Mean tree biomass accumulation in both the <1–5 and 6–10 years old (4.4 and 5.7 Mg ha⁻¹ yr⁻¹, respectively) abandoned pastures was lower than predicted and deviated significantly (57% and 41%) from rates estimated from the chronosequence. The older SFs, with a mean growth rate of 9.9 Mg ha⁻¹ yr⁻¹ followed the rate predicted by the chronosequence. Understocking was the primary cause of low biomass recovery rates in the youngest forests; although the youngest stands had a diameter at breast height increment three times the oldest stands, the youngest stands lacked sufficient density to cumulatively produce high biomass accumulation rates. Four years of measurement indicated that the youngest stands had developed 59% of the stems measured in the older stands during the same time period. The 6–10‐year‐old stands were rapidly self‐thinning and approached stem density values measured in the same aged stands at the onset of the study. Mortality was high for all stands, with 54% of the original stems remaining after 4 years in intermediate‐aged stands. The forests were dominated by the tree Vismia, which represented 55–66% of the biomass in all stands. The Vismia share of the biomass was decreasing over time, with other genera replacing the pioneer. Our measured rates of regrowth indicate that generalized estimates of forest regrowth through chronosequence studies will overestimate forest regrowth for the youngest forests that were under land use for longer time‐periods before abandonment. Certified Emission Reductions under the Clean Development Mechanism of the Kyoto protocol should consider these results when predicting and compensating for carbon sequestered under natural forest management.     

Partitioning direct and indirect human-induced effects on carbon sequestration of managed coniferous forests using model simulations and forest inventories

Temperate forest ecosystems have recently been identified as an important net sink in the global carbon budget. The factors responsible for the strength of the sinks and their permanence, however, are less evident. In this paper, we quantify the present carbon sequestration in Thuringian managed coniferous forests. We quantify the effects of indirect human‐induced environmental changes (increasing temperature, increasing atmospheric CO₂ concentration and nitrogen fertilization), during the last century using BIOME‐BGC, as well as the legacy effect of the current age‐class distribution (forest inventories and BIOME‐BGC). We focused on coniferous forests because these forests represent a large area of central European forests and detailed forest inventories were available. The model indicates that environmental changes induced an increase in biomass C accumulation for all age classes during the last 20 years (1982–2001). Young and old stands had the highest changes in the biomass C accumulation during this period. During the last century mature stands (older than 80 years) turned from being almost carbon neutral to carbon sinks. In high elevations nitrogen deposition explained most of the increase of net ecosystem production (NEP) of forests. CO₂ fertilization was the main factor increasing NEP of forests in the middle and low elevations. According to the model, at present, total biomass C accumulation in coniferous forests of Thuringia was estimated at 1.51 t C ha^?1 yr^?1 with an averaged annual NEP of 1.42 t C ha^?1 yr^?1 and total net biome production of 1.03 t C ha^?1 yr^?1 (accounting for harvest). The annual averaged biomass carbon balance (BCB: biomass accumulation rate‐harvest) was 1.12 t C ha^?1 yr^?1 (not including soil respiration), and was close to BCB from forest inventories (1.15 t C ha^?1 yr^?1). Indirect human impact resulted in 33% increase in modeled biomass carbon accumulation in coniferous forests in Thuringia during the last century. From the forest inventory data we estimated the legacy effect of the age‐class distribution to account for 17% of the inventory‐based sink. Isolating the environmental change effects showed that these effects can be large in a long‐term, managed conifer forest.     

Litterfall and forest floor dynamics in Eucalyptus pilularis forests

Abstract Calculations relating the input of litterfall to litter or forest floor mass in forests generally assume that the forest floor reaches an equilibrium state. Based on this assumption, a decomposition factor (k) can be calculated. In the present paper, this basic assumption is questioned and the implications considered. Data on litterfall and forest floor from blackbutt (Eucalyptus pilularis) regrowth forests and plantations were collated from publications and the authors' studies to evaluate both assumptions and relationships. Blackbutt grows over a wide environmental range but its main distribution is in mild temperate to subtropical conditions. Data were from single‐plot studies, sequential studies and chronosequences in both plantations and native regrowth forests. Stands ranged in age from 3 years to maturity in the case of pure, or almost pure blackbutt stands. The forest floor biomass increased up to 12.3 tha^?1 at 33 years of age with no evidence of steady state. Litterfall increased up to 7.8 t ha^?1 year^?1 and was correlated with crown biomass. Regrowth stands were relatively undisturbed and more than 20 years of age, and litterfall ranged from 4.1 to 11.6 tha^?1 year^?1 and was correlated with stand basal area. Forest floor mass in regrowth forests was variable between the different aged stands but did not exceed 18 tha^?1, and there was no evidence that steady state was achieved. The forest floor mass was related to, and approximately 1.7 times the input of litterfall. Although the assumption of steady state was not valid, a k' factor was estimated that related input to forest floor mass and this was relatively constant across all stands and correlated with generalized environmental data. Although assumptions of forest floor equilibrium cannot be supported for E. pilularis, it still should be possible to predict forest floor mass and decomposition from stand conditions and general environmental data.     

Biomass,Carbon, and Nitrogen Pools in Mexican Tropical Dry Forest Landscapes

Tropical dry forest is the most widely distributed land-cover type in the tropics. As the rate of land-use/land-cover change from forest to pasture or agriculture accelerates worldwide, it is becoming increasingly important to quantify the ecosystem biomass and carbon (C) and nitrogen (N) pools of both intact forests and converted sites. In the central coastal region of México, we sampled total aboveground biomass (TAGB), and the N and C pools of two floodplain forests, three upland dry forests, and four pastures converted from dry forest. We also sampled belowground biomass and soil C and N pools in two sites of each land-cover type. The TAGB of floodplain forests was as high as 416 Mg ha^–1, whereas the TAGB of the dry forest ranged from 94 to 126 Mg ha^–1. The TAGB of pastures derived from dry forest ranged from 20 to 34 Mg ha^–1. Dead wood (standing and downed combined) comprised 27%–29% of the TABG of dry forest but only about 10% in floodplain forest. Root biomass averaged 32.0 Mg ha^–1 in floodplain forest, 17.1 Mg ha^–1 in dry forest, and 5.8 Mg ha^–1 in pasture. Although total root biomass was similar between sites within land-cover types, root distribution varied by depth and by size class. The highest proportion of root biomass occurred in the top 20 cm of soil in all sites. Total aboveground and root C pools, respectively, were 12 and 2.2 Mg ha^–1 in pasture and reached 180 and 12.9 Mg ha^–1 in floodplain forest. Total aboveground and root pools, respectively, were 149 and 47 kg ha^–1 in pasture and reached 2623 and 264 kg ha^–1 in floodplain forest. Soil organic C pools were greater in pastures than in dry forest, but soil N pools were similar when calculated for the same soil depths. Total ecosystem C pools were 306. The Mg ha^–1 in floodplain forest, 141 Mg ha^–1 in dry forest, and 124 Mg ha^–1 in pasture. Soil C comprised 37%–90% of the total ecosystem C, whereas soil N comprised 85%–98% of the total. The N pools lack of a consistent decrease in soil pools caused by land-use change suggests that C and N losses result from the burning of aboveground biomass. We estimate that in México, dry forest landscapes store approximately 2.3 Pg C, which is about equal to the C stored by the evergreen forests of that country (approximately 2.4 Pg C). Potential C emissions to the atmosphere from the burning of biomass in the dry tropical landscapes of México may amount to 708 Tg C, as compared with 569 Tg C from evergreen forests.     

Bryophytes attenuate anthropogenic nitrogen inputs in boreal forests

Productivity in boreal ecosystems is primarily limited by available soil nitrogen (N), and there is substantial interest in understanding whether deposition of anthropogenically derived reactive nitrogen (N_r) results in greater N availability to woody vegetation, which could result in greater carbon (C) sequestration. One factor that may limit the acquisition of N_r by woody plants is the presence of bryophytes, which are a significant C and N pool, and a location where associative cyanobacterial N‐fixation occurs. Using a replicated stand‐scale N‐addition experiment (five levels: 0, 3, 6, 12, and 50 kg N ha^?1 yr^?1; n=6) in the boreal zone of northern Sweden, we tested the hypothesis that sequestration of N_r into bryophyte tissues, and downregulation of N‐fixation would attenuate N_r inputs, and thereby limit anthropogenic N_r acquisition by woody plants. Our data showed that N‐fixation per unit moss mass and per unit area sharply decreased with increasing N addition. Additionally, the tissue N concentrations of Pleuorzium schreberi increased and its biomass decreased with increasing N addition. This response to increasing N addition caused the P. schreberi N pool to be stable at all but the highest N addition rate, where it significantly decreased. The combined effects of changed N‐fixation and P. schreberi biomass N accounted for 56.7% of cumulative N_r additions at the lowest N_r addition rate, but only a minor fraction for all other treatments. This ‘bryophyte effect’ can in part explain why soil inorganic N availability and acquisition by woody plants (indicated by their δ¹⁵N signatures) remained unchanged up to N addition rates of 12 kg ha^?1 yr^?1 or greater. Finally, we demonstrate that approximately 71.8% of the boreal forest experiences N_r deposition rates at or below 3 kg ha^?1 yr^?1, suggesting that bryophytes likely limit woody plant acquisition of ambient anthropogenic N_r inputs throughout a majority of the boreal forest.