Postfire nitrogen balance of Mediterranean shrublands: Direct combustion losses versus gaseous and leaching losses from the postfire soil mineral nitrogen flush

Fire is a major factor controlling global carbon (C) and nitrogen (N) cycling. While direct C and N losses caused by combustion have been comparably well established, important knowledge gaps remain on postfire N losses. Here, we quantified both direct C and N combustion losses as well as postfire gaseous losses (N₂O, NO and N₂) and N leaching after a high‐intensity experimental fire in an old shrubland in central Spain. Combustion losses of C and N were 9.4 Mg C/ha and 129 kg N/ha, respectively, representing 66% and 58% of initial aboveground vegetation and litter stocks. Moreover, fire strongly increased soil mineral N concentrations by several magnitudes to a maximum of 44 kg N/ha 2 months after the fire, with N largely originating from dead soil microbes. Postfire soil emissions increased from 5.4 to 10.1 kg N ha^?1 year^?1 for N₂, from 1.1 to 1.9 kg N ha^?1 year^?1 for NO and from 0.05 to 0.2 kg N ha^?1 year^?1 for N₂O. Maximal leaching losses occurred 2 months after peak soil mineral N concentrations, but remained with 0.1 kg N ha^?1 year^?1 of minor importance for the postfire N mass balance. ¹⁵N stable isotope labelling revealed that 33% of the mineral N produced by fire was incorporated in stable soil N pools, while the remainder was lost. Overall, our work reveals significant postfire N losses dominated by emissions of N₂ that need to be considered when assessing fire effects on ecosystem N cycling and mass balance. We propose indirect N gas emissions factors for the first postfire year, equalling to 7.7% (N₂‐N), 2.7% (NO‐N) and 5.0% (N₂O‐N) of the direct fire combustion losses of the respective N gas species.     

Effects of nitrogen deposition and management practices on leaf litterfall and N and P return in a Moso bamboo forest

The effects of nitrogen (N) deposition and management practices on leaf litterfall and N and P return in Moso bamboo forest are not yet known. In this study, we investigated the effects of four levels of simulated N deposition, including low-N (N30, 30 kg ha^?1 year^?1), medium-N (N60, 60 kg ha^?1 year^?1), and high-N (N90, 90 kg ha^?1 year^?1), and a control with no N (N-free addition). The experiment was performed in a Moso bamboo forest under conventional management (CM) and intensive management (IM). The results showed that leaf litterfall and N and P return occurred mainly from March to June and accounted for 78.2–82.2, 78.5–82.1, and 85.6–94.6% of annual leaf litterfall, N return, and P return, respectively. Unlike CM, IM significantly increased leaf litterfall and N and P return. The positive effects were further amplified by low- and medium-N deposition, but not high-N deposition. The combination of low- and medium-N deposition and IM significantly increased N and P return, but not litterfall. Our results indicated that the interaction of anthropogenic management practices and N deposition need to be considered when estimating the effects of N deposition on the biogeochemical cycle of a forest ecosystem.     

Fluxes of nitrogen and phosphorus in a gallery forest in the Cerrado of central Brazil

The Gallery forests of the Cerrado biome play a critical role in controlling stream chemistry but little information about biogeochemical processes in these ecosystems is available. This work describes the fluxes of N and P in solutions along a topographic gradient in a gallery forest. Three distinct floristic communities were identified along the gradient: a wet community nearest the stream, an upland dry community adjacent to the woodland savanna and an intermediate community between the two. Transects were marked in the three communities for sampling. Fluxes of N from bulk precipitation to these forests resulted in deposition of 12.6 kg ha^?1 y^?1 of total N of which 8.8 kg ha^?1 was as inorganic N. The throughfall flux of total N was generally <8.4 kg ha^?1 year^?1. Throughfall NO₃?CN fluxes were higher (7?C32%) while NH₄?CN and organic N fluxes were lower (54?C69% and 5?C46%) than those in bulk precipitation. The throughfall flux was slightly lower for the wet forest community compared to other communities. Litter leachate fluxes differed among floristic communities with higher NH₄?CN in the wet community. The total N flux was greater in the wet forest than in the dry forest (13.5 vs. 9.4 kg ha^?1 year^?1, respectively). The stream water had total N flux of 0.3 kg ha^?1 year^?1. The flux of total P through bulk precipitation was 0.7 kg ha^?1 year^?1 while the mean fluxes of total P in throughfall (0.6 kg ha^?1 year^?1) and litter leachate (0.5 kg ha^?1 year^?1) declined but did not differ between communities. The low concentrations presented in soil solution and low fluxes in stream water (0.3 and 0.1 kg ha^?1 year^?1 for N and P, respectively) relative to other flowpaths emphasize the conservative nutrient cycling of these forests and the importance of internal recycling processes for the maintenance and conservation of riparian and stream ecosystems in the Cerrado.     

Nitrogen biogeochemistry of a mature Scots pine forest subjected to high nitrogen loads

Nitrogen (N) biogeochemistry of a mature Scots pine (Pinus sylvestris L.) stand subjected to an average total atmospheric N deposition of 48 kg ha^?1 year^?1 was studied during the period 1992–2007. The annual amount of dissolved inorganic nitrogen (DIN) in throughfall (TF) averaged 34 kg ha^?1 year^?1 over the 16-year monitoring period. The throughfall fluxes contained also considerable amounts of dissolved organic nitrogen (DON) (5–8.5 kg N ha^?1 year^?1), which should be incorporated in the estimate of N flux using throughfall collectors. Throughfall DIN fluxes declined at a rate of ?0.9 kg N ha^?1 year^?1, mainly due to the decreasing TF fluxes of ammonium (NH₄), which accounted for 70% to TF DIN. The decrease in TF DIN was accompanied by a decrease in DIN leaching in the seepage water (?1.6 kg N ha^?1 year^?1), which occurred exclusively as nitrate (NO₃ ^?). Nitrate losses in the leachate of the forest floor (LFH) equalled the TF NO₃ ^? delivered to the LFH-layer. On the contrary, about half of the TF NH₄ ⁺ was retained within the LFH-layer. Approximately 60% of the TF DIN fluxes were leached indicating that N inputs were far in excess of the N requirements of the forest. For DON, losses were only substantial from the LFH-layer, but no DON was leached in the seepage water. Despite the high N losses through nitrate leaching and NO _x emission, the forest was still accumulating N, especially in the aggrading LFH-layer. The forest stand, on the contrary, was found to be a poor N sink.     

Effects of experimental N addition on plant diversity in an old‐growth temperate forest

Temperate forest ecosystems have experienced mounting negative effects due to increasing levels of nitrogen (N) deposition. We examined the effects of experimental N addition on plant diversity in an old‐growth temperate forest to test the following hypothesis: Long‐term excessive N addition decreases plant diversity by affecting the growth of plants, which results from changes in the soil nutrient content and a decrease in the soil pH in temperate forests. Experimental N additions were administered at the following levels since 2008: control (0 kg N ha^?1 year^?1), low N (30 kg N ha^?1 year^?1), medium N (60 kg N ha^?1 year^?1), and high N (120 kg N ha^?1 year^?1). Additionally, plant diversity was studied from 2014 to 2016. The results showed that the experimental N additions had significant effects on plant diversity and soil properties in an old‐growth temperate forest. The high‐N treatment decreased the density, cover, and diversity of understory plants, and some herbs even appeared to undergo premature aging, whereas the species diversity of herbs and ferns in the low‐N treatment plots showed a slight increasing tendency. This may have been because the old‐growth temperate forest is an N‐limited ecosystem, so the moderate N input did not show a large influence on plant diversity. However, the long‐term high‐N treatment ultimately reduced plant diversity by changing the soil nutrient contents, decreasing the pH values, and damaging plant growth. Our results suggested that the long‐term excessive N addition negatively affected the forest ecosystem in an N‐limited temperature forest.     

Simulated nitrogen deposition significantly suppresses the decomposition of forest litter in a natural evergreen broad-leaved forest in the Rainy Area of Western China

Background and aims

Litter, an essential component of forest ecosystems, plays an important role in maintaining soil fertility, sequestering carbon (C) and improving soil biodiversity. However, litter decomposition is affected by increased nitrogen (N) deposition. Numerous reports have presented N deposition experiments in different forest ecosystems to investigate the effects of N deposition on litter decomposition, but the effects remain unclear, especially in ecosystems receiving increasingly higher levels of ambient N deposition. To address this gap, we performed a litterbag experiment to understand the effects of increasing N deposition on the litter decomposition process in natural evergreen broad-leaved forest in the Rainy Area of Western China.

Methods

A 2-year field litter decomposition experiment was conducted using the litterbag method. Four levels of N deposition were established: control (CK; 0 kg·N·ha^?1·year^?1), low N deposition (LN; 50 kg·N·ha^?1·year^?1), medium N deposition (MN; 150 kg·N·ha^?1·year^?1), and high N deposition (HN; 300 kg·N·ha^?1·year^?1). The simulated N depositions ranged from 50% to 320% of the ambient rate of wet N deposition.

Results

Simulated N deposition significantly increased the remaining mass, C, N, lignin and cellulose of the litter. The LN treatment decreased the remaining phosphorus (P); conversely, the HN treatment increased it. In the late stage of the study period, the mass remaining was positively closely correlated to the lignin and cellulose remaining during the decomposition process.

Conclusions

Simulated N deposition significantly suppressed the litter decomposition in the natural evergreen broad-leaved forest, despite the high rate of ambient N deposition, and the inhibitory effects increased with the N deposition levels. The suppressive effect of N deposition on litter decomposition may be primarily explained by the inhibition of lignin and cellulose degradation by the exogenous inorganic N. With ongoing N deposition in future, N deposition may have a potentially significant impact on C and N cycles in such forest ecosystems.

     

Symptoms of nitrogen saturation in an aggrading forested watershed in western Maryland

The objective of this study was to evaluate the nitrogen (N) biogeochemistry of an 18–22 year old forested watershed in western Maryland. We hypothesized that this watershed should not exhibit symptoms of N saturation. This watershed was a strong source of nitrate (NO₃ ⁻) to the stream in all years, with a mean annual export of 9.5 kg N ha⁻¹ year⁻¹ and a range of 4.4–18.4 kg N ha⁻¹ year⁻¹. During the 2001 and 2002 water years, wet deposition of inorganic N was 9.0 kg N ha⁻¹ year⁻¹ and 6.3 kg N ha⁻¹ year⁻¹, respectively. Watershed N export rates in 2001 and 2002 water years were 4.2 kg N ha⁻¹ year⁻¹ and 5.3 kg N ha⁻¹ year⁻¹, respectively. During the wetter water years of 2003 and 2004, the watershed exported 15.0 kg N ha⁻¹ year⁻¹ and 18.4 kg N ha⁻¹ year⁻¹, rates that exceeded annual wet deposition of N by a factor of two (7.5 kg N ha⁻¹ year⁻¹ in 2003) and three (5.5 kg N ha⁻¹ year⁻¹ in 2004). Consistent with the high rates of N export, were high concentrations (2.1–3.3%) of N in foliage, wood (0.3%) and fine roots, low C:N ratios in the forest floor (17–24) and mineral soil (14), high percentages (83–96%) of the amount of mineralized N that was nitrified and elevated N concentrations (up to 3 mg N l⁻¹) in soil solution. Although this watershed contained a young aggrading forest, it exhibited several symptoms of N saturation commonly observed in more mature forests.     

Exotic grass invasion alters potential rates of N fixation in Hawaiian woodlands

Exotic grass invasion promotes fire which drives the conversion of native woodlands to exotic grasslands in the seasonally dry submontane forests of the island of Hawai'i. We compared potential rates of N fixation in an unburned forest site and a converted grassland site using the acetylene reduction assay. In addition to measuring rates of N fixation on separate and mixed substrates in each site, we tested the effect of abiotic factors on rates of N fixation of specific substrates. We hypothesized that rates of N fixation would be higher in the converted grassland site. N fixation estimates were 4.9 kg N ha⁻¹ year⁻¹ for the unburned forest, and 0.10 kg N ha⁻¹ year⁻¹ for the grassland site, so our hypothesis was rejected. The N fixation in the unburned forest occurs mostly on the leaf litter of native woody species. These substrates are absent from the grassland site, except for wood debris which was not consumed during the fires. No nitrogenase activity was detected in the rhizosphere and litter of grasses, the rhizospheres of shrubs or in soil. Although wood debris is not a significant contributor to the N fixed in the unburned forest, it contributes the majority of N fixed in the grassland. The response of nitrogenase activity to varying conditions of moisture and temperature suggests that microclimatic differences between sites do not control differences in N fixation activity; rather, these differences are due to the abundance of N-fixing substrates. The substantial decrease in N fixation activity after the conversion from woodland to grassland implies that ecosystem-level rates of N accretion are decreased by fire in these sites so much that the N lost during volatilization due to fire is not replenished over the long term by N fixation. Received: 10 January 1997 / Accepted: 7 August 1997     

Differences in Englemann Spruce Forest Biogeochemistry East and West of the Continental Divide in Colorado, USA

We compared Englemann spruce biogeochemical processes in forest stands east and west of the Continental Divide in the Colorado Front Range. The divide forms a natural barrier for air pollutants such that nitrogen (N) emissions from the agricultural and urban areas of the South Platte River Basin are transported via upslope winds to high elevations on the east side but rarely cross over to the west side. Because there are far fewer emissions sources to the west, atmospheric N deposition is 1–2 kg N ha⁻¹ y⁻¹ on the west side, as compared with 3–5 kg N ha⁻¹ y⁻¹ on the east side. Species composition, elevation, aspect, parent material, site history, and climate were matched as closely as possible across six east and six west side old-growth forest stands. Higher N deposition sites had significantly lower organic horizon C:N and lignin:N ratios, lower foliar C:N ratios, as well as greater %N, higher N:Ca, N:Mg, and N:P ratios, and higher potential net mineralization rates. When C:N ratios dropped below 29, as they did in east-side organic horizon soils, mineralization rates increased linearly. Our results are comparable to those from studies of the northeastern United States and Europe that have found changes in forest biogeochemistry in response to N deposition inputs between 3 and 60 kg ha⁻¹ y⁻¹. Though they are low by comparison with more densely populated and agricultural regions, current levels of N deposition, have caused measurable changes in Englemann spruce forest biogeochemistry east of the Continental Divide in Colorado. Received 22 January 2001; accepted 11 June 2001.     

Do Forests Receive Occult Inputs of Nitrogen?

The nitrogen (N) cycle of forest ecosystems is understood relatively well, and few scientists expect that major revisions will be necessary; most current work on N cycling focuses on improving the precision estimates of pools and fluxes, or measuring the magnitudes of well-known pools in response to management or disturbances. However, in the past few decades more than a dozen articles in refereed journals have claimed very high rates of N input, far beyond the rates expected for known sources of N. In this review, we summarize the literature on N accretion rates in forests that lack substantial contributions from symbiotic N-fixing plants. We critique each study for the strength of the experimental design behind the estimate of N accretion and consider whether unexpectedly large inputs of N really occur in forests. Only 6 of 24 estimates of N accretion had strong experimental designs, and only 2 of these 6 yielded estimates of >5 kgN ha^-1 y^-1. The high accretion estimates with a strong experimental design come from repeated sampling at the Walker Branch watersheds in Tennessee, where N accretion rates ranged from 50 to 80 kg N ha^-1 y^-1 over 15 years after harvesting. At the same location, an unharvested stand showed no significant change. We conclude that there is no widespread evidence of high rates of occult N input in forests. Too few studies have carefully tested for balanced N budgets in forests (inputs minus outputs plus change in storage), and we recommend that at least a few of these studies be undertaken on soils that permit high precision sampling. Received 14 December 1999; accepted 22 March 2000.     

Urban influences on the nitrogen cycle in Puerto Rico

Anthropogenic actions are altering fluxes of nitrogen (N) in the biosphere at unprecedented rates. Efforts to study these impacts have concentrated in the Northern hemisphere, where experimental data are available. In tropical developing countries, however, experimental studies are lacking. This paper summarizes available data and assesses the impacts of human activities on N fluxes in Puerto Rico, a densely populated Caribbean island that has experienced drastic landscape transformations over the last century associated with rapid socioeconomic changes. N yield calculations conducted in several watersheds of different anthropogenic influences revealed that disturbed watersheds export more N per unit area than undisturbed forested watersheds. Export of N from urban watersheds ranged from 4.8 kg ha^?1 year^?1 in the Río Bayamón watershed to 32.9 kg ha^?1 year^?1 in the highly urbanized Río Piedras watershed and 33.3 kg ha^?1 year^?1 in the rural-agricultural Río Grande de Añasco watershed. Along with land use, mean annual runoff explained most of the variance in fluvial N yield. Wastewater generated in the San Juan Metropolitan Area receives primary treatment before it is discharged into the Atlantic Ocean. These discharges are N-rich and export large amounts of N to the ocean at a rate of about 140 kg ha^?1 year^?1. Data on wet deposition of inorganic N ( $\hbox{NH}_{4}^{+}+\hbox{NO}_{3}^{-}Anthropogenic actions are altering fluxes of nitrogen (N) in the biosphere at unprecedented rates. Efforts to study these impacts have concentrated in the Northern hemisphere, where experimental data are available. In tropical developing countries, however, experimental studies are lacking. This paper summarizes available data and assesses the impacts of human activities on N fluxes in Puerto Rico, a densely populated Caribbean island that has experienced drastic landscape transformations over the last century associated with rapid socioeconomic changes. N yield calculations conducted in several watersheds of different anthropogenic influences revealed that disturbed watersheds export more N per unit area than undisturbed forested watersheds. Export of N from urban watersheds ranged from 4.8 kg ha⁻¹ year⁻¹ in the Río Bayamón watershed to 32.9 kg ha⁻¹ year⁻¹ in the highly urbanized Río Piedras watershed and 33.3 kg ha⁻¹ year⁻¹ in the rural-agricultural Río Grande de A?asco watershed. Along with land use, mean annual runoff explained most of the variance in fluvial N yield. Wastewater generated in the San Juan Metropolitan Area receives primary treatment before it is discharged into the Atlantic Ocean. These discharges are N-rich and export large amounts of N to the ocean at a rate of about 140 kg ha⁻¹ year⁻¹. Data on wet deposition of inorganic N () suggest that rates of atmospheric N deposition are increasing in the pristine forests of Puerto Rico. Stationary and mobile sources of NO _x (NO+NO₂) and N₂O generated in the large urban centers may be responsible for this trend. Comprehensive measurements are required in Puerto Rico to quantitatively characterize the local N cycle. More research is required to assess rates of atmospheric N deposition, N fixation in natural and human-dominated landscapes, N-balance associated with food and feed trade, and denitrification.     

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.     

Effect of nitrogen and phosphorus fertilization on the composition of rhizobacterial communities of two Chilean Andisol pastures

The effect of nitrogen (N) and phosphorus (P) fertilization on composition of rhizobacterial communities of volcanic soils (Andisols) from southern Chile at molecular level is poorly understood. This paper investigates the composition of rhizobacterial communities of two Andisols under pasture after 1- and 6-year applications of N (urea) and P (triple superphosphate). Soil samples were collected from two previously established sites and the composition of rhizobacterial communities was determined by denaturing gradient gel electrophoresis (PCR–DGGE). The difference in the composition and diversity between rhizobacterial communities was assessed by nonmetric multidimensional scaling (MDS) analysis and the Shannon–Wiener index. In Site 1 (fertilized for 1 year), PCR–DGGE targeting 16S rRNA genes and MDS analysis showed that moderate N application (270 kg N ha^?1 year^?1) without P significantly changed the composition of rhizobacterial communities. However, no significant community changes were observed with P (240 kg P ha^?1 year^?1) and N–P application (270 kg N ha^?1 year^?1 plus 240 kg P ha^?1 year^?1). In Site 2 (fertilized for 6 years with P; 400 kg P ha^?1 year^?1), PCR–DGGE targeting rpoB, nifH, amoA and alkaline phosphatase genes and MDS analysis showed changes in rhizobacterial communities only at the highest rate of N application (600 kg N ha^?1 year^?1). Quantitative PCR targeting 16S rRNA genes also showed higher abundance of bacteria at higher N application. In samples from both sites, the Shannon–Wiener index did not show significant difference in the diversity of rhizobacterial communities. The changes observed in rhizobacterial communities coincide in N fertilized pastures with lower soil pH and higher pasture yields. This study indicates that N–P application affects the soil bacterial populations at molecular level and needs to be considered when developing fertilizer practices for Chilean pastoral Andisols.     

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.     

Fire effects on ecosystem nitrogen cycling in a Californian bishop pine forest

Fire can cause severe nitrogen (N) losses from grassland, chaparral, and temperate and boreal forest ecosystems. Paradoxically, soil ammonium levels are markedly increased by fire, resulting in high rates of primary production in re-establishing plant communities. In a manipulative experiment, we examined the influence of wild-fire ash residues on soil, microbial and plant N pools in a recently burned Californian bishop pine (Pinus muricata D. Don) forest. Ash stimulated post-fire primary production and ecosystem N retention through direct N inputs from ash to soils, as well as indirect ash effects on soil N availability to plants. These results suggest that redistribution of surface ash after fire by wind or water may cause substantial heterogeneity in soil N availability to plants, and could be an important mechanism contributing to vegetation patchiness in fire-prone ecosystems. In addition, we investigated the impact of fire on ecosystem N cycling by comparing ¹⁵N natural abundance values from recently burned and nearby unburned P. muricata forest communities. At the burned site, ¹⁵N natural abundance in recolonising species was similar to that in bulk soil organic matter. By contrast, there was a marked ¹⁵N depletion in the same species relative to the total soil N pool at the unburned site. These results suggest that plant uptake of nitrate (which tends to be strongly depleted in ¹⁵N because of fractionation during nitrification) is low in recently burned forest communities but could be an important component of eco- system N cycling in mature conifer stands. Received: 29 June 1999 / Accepted: 24 October 1999     

Nitrogen Addition Shapes Soil Phosphorus Availability in Two Reforested Tropical Forests in Southern China

Scant information is available on how soil phosphorus (P) availability responds to atmospheric nitrogen (N) deposition, especially in the tropical zones. This study examined the effect of N addition on soil P availability, and compared this effect between forest sites of contrasting land‐use history. Effects of N addition on soil properties, litterfall production, P release from decomposing litter, and soil P availability were studied in a disturbed (reforested pine forest with previous understory vegetation and litter harvesting) and a rehabilitated (reforested mixed pine/broadleaf forest with no understory vegetation and litter harvesting) tropical forest in southern China. Experimental N‐treatments (above ambient) were the following: Control (no N addition), N50 (50 kg N ha^?1 yr^?1), and N100 (100 kg N ha^?1 yr^?1). Results indicated that N addition significantly decreased soil P availability in the disturbed forest. In the rehabilitated forest, however, soil P availability was significantly increased by N addition. Decreases in soil P availability may be correlated with decreases in rates of P release from decomposing litter in the N‐treated plots, whereas the increase in soil P availability was correlated with an increase in litterfall production. Our results suggest that response of soil P availability to N deposition in the reforested tropical forests in southern China may vary greatly with temporal changes in tree species composition and soil nutrient status, caused by different land‐use practices.     

Wildfire effects on carbon and nitrogen in inland coniferous forests

A ponderosa pine/Douglas-fir forest (Pinus ponderosa Dougl., Pseudotsuga menziesii (Mirb.) Franco; PP/DF) and a lodgepole pine/Engelmann spruce forest (Pinus contorta Loud., Picea engelmannii Parry ex Engelm.; LP/ES) located on the eastern slopes of the Cascade Mountains in Washington state, USA, were examined following severe wildfire to compare total soil carbon and nitrogen capitals with unburned (control) forests. One year after fire, the average C content (60 cm depth) of PP/DF and LP/ES soil was 30% (25 Mg ha^-1) and 10% (7 Mg ha^-1) lower than control soil. Average N content on the burned PP/DF and LP/ES plots was 46% (3.0 Mg ha^-1) and 13% (0.4 Mg ha^-1) lower than control soil. The reduction in C and N in the PP/DF soil was largely the result of lower nutrient capitals in the burned Bw horizons (12–60 cm depth) relative to control plots. It is unlikely that the 1994 fire substantially affected nutrient capitals in the Bw horizons; however, natural variability or past fire history could be responsible for the varied nutrient capitals observed in the subsurface soils. Surface erosion (sheet plus rill) removed between 15 and 18 Mg ha^-1 of soil from the burned plots. Nutrient losses through surface erosion were 280 kg C ha^-1 and 14 kg N ha^-1 in the PP/DF, whereas LP/ES losses were 640 and 22 kg ha^-1 for C and N, respectively. In both forests, surface erosion of C and N was _∼1% to 2% of the A-horizon capital of these elements in unburned soil. A bioassay (with lettuce as an indicator plant) was used to compare soils from low-, moderate- and high-severity burn areas relative to control soil. In both forests, low-severity fire increased lettuce yield by 70–100% of controls. With more severe fire, yield decreased in the LP/ES relative to the low-intensity burn soil; however, only in the high-severity treatment was yield reduced (14%) from the control. Moderate- and high-severity burn areas in the PP/DF were fertilized with _∼56 kg ha^-1 of N four months prior to soil sampling. In these soils, yield was 70–80% greater than the control. These results suggest that short-term site productivity can be stimulated by low-severity fire, but unaffected or reduced by more severe fire in the types of forests studied. Post-fire fertilization with N could increase soil productivity where other environmental factors do not limit growth. This revised version was published online in June 2006 with corrections to the Cover Date.     

Impact of prescribed burning on the nutrient balance of heathlands with particular reference to nitrogen and phosphorus

Question: Can prescribed winter burning compensate atmospheric nutrient loads for dry heathlands? What effects does prescribed burning have on nutrient balances, particularly as regards the limiting nutrients N and P? Location: Lueneburg Heath, NW Germany. Methods: In two burning experiments (in 10/15 year old Calluna‐stands) nutrient balances (for N, Ca, K, Mg, P) were calculated by analysing nutrient inputs (atmospheric deposition, ash deposition), nutrient stores (above‐ground biomass, organic horizon) and nutrient outputs (biomass combustion, leaching). Results: Atmospheric nutrient deposition amounted to 22.8 kg.ha^‐1.a^‐1 for N and < 0.5 kg.ha^‐1.a^‐1 for P. Nutrient stores in the above‐ground biomass were 95/197 kg.ha^‐1 for N and 5/13 kg.ha^‐1 for P (first/second experiment, respectively). From these stores 90/53% (for N) and 25/14% (for P) were removed by burning. Effects of leaching on nutrient balances were low. In the first two years after burning, leaching rates of N increased by about 4/6 kg.ha^‐1, whereas leaching rates of P did not change significantly. Input/output‐ratios showed that prescribed burning leads to positive nutrient balances for N, Ca and Mg in the long term. For example, the amounts of N removed by prescribed burning are equivalent to ca. five years of atmospheric inputs. Applied in ten‐year cycles, this measure alone cannot prevent N accumulation in the long term. Conclusion: Regarding 10/15 year old Calluna‐heaths, we assume that prescribed burning cannot compensate for atmospheric N inputs, thus making long‐term changes in the nutritional state inevitable. Therefore, prescribed burning should be applied in combination with high‐intensity management measures.     

The relationship between fire history and an exotic fungal disease in a deciduous forest

Exotic diseases have fundamentally altered the structure and function of forest ecosystems. Controlling exotic diseases across large expanses of forest has proven difficult, but fire may reduce the levels of diseases that are sensitive to environmental conditions. We examined Cornus florida populations in burned and unburned Quercus–Carya stands to determine if burning prior to anthracnose infection has reduced the impacts of an exotic fungal disease, dogwood anthracnose, caused by Discula destructiva. We hypothesized that fire has altered stand structure and created open conditions less conducive to dogwood anthracnose. We compared C. florida density, C. florida health, and species composition and density among four sampling categories: unburned stands, and stands that had burned once, twice, and 3 times over a 20-year period (late 1960s to late 1980s). Double burn stands contained the greatest density of C. florida stems (770 stems ha⁻¹) followed by triple burn stands (233 stems ha⁻¹), single burn stands (225 stems ha⁻¹) and unburned stands (70 stems ha⁻¹; P < 0.01). We observed less crown dieback in small C. florida trees (<5 cm diameter at breast height) in burned stands than in unburned stands (P < 0.05). Indicator species analysis showed that burning favored species historically associated with Quercus–Carya forests and excluded species associated with secondary succession following nearly a century of fire suppression. Our results suggest that fire may mitigate the decline of C. florida populations under attack by an exotic pathogen by altering forest structure and composition. Further, our results suggest that the burns we sampled have had an overall restorative effect on forest communities and were within the fire return interval of the historic fire regime. Consequently, prescribed fire may offer a management tool to reduce the impacts of fungal disease in forest ecosystems that developed under historic burning regimes.     

Carbon cycling and net ecosystem production at an early stage of secondary succession in an abandoned coppice forest

Secondary mixed forests are one of the dominant forest cover types in human-dominated temperate regions. However, our understanding of how secondary succession affects carbon cycling and carbon sequestration in these ecosystems is limited. We studied carbon cycling and net ecosystem production (NEP) over 4 years (2004–2008) in a cool-temperate deciduous forest at an early stage of secondary succession (18 years after clear-cutting). Net primary production of the 18-year-old forest in this study was 5.2 tC ha^?1 year^?1, including below-ground coarse roots; this was partitioned into 2.5 tC ha^?1 year^?1 biomass increment, 1.6 tC ha^?1 year^?1 foliage litter, and 1.0 tC ha^?1 year^?1 other woody detritus. The total amount of annual soil surface CO₂ efflux was 6.8 tC ha^?1 year^?1, which included root respiration (1.9 tC ha^?1 year^?1) and heterotrophic respiration (RH) from soils (4.9 tC ha^?1 year^?1). The 18-year forest at this study site exhibited a great increase in biomass pool as a result of considerable total tree growth and low mortality of tree stems. In contrast, the soil organic matter (SOM) pool decreased markedly (?1.6 tC ha^?1 year^?1), although further study of below-ground detritus production and RH of SOM decomposition is needed. This young 18-year forest was a weak carbon sink (0.9 tC ha^?1 year^?1) at this stage of secondary succession. The NEP of this 18-year forest is likely to increase gradually because biomass increases with tree growth and with the improvement of the SOM pool through increasing litter and dead wood production with stand development.