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.     

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

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

Soil nitrogen dynamics three years after a severe Araucaria–Nothofagus forest fire

Wildfires have shaped the biogeography of south Chilean Araucaria–Nothofagus rainforest vegetation patterns, but their impact on soil properties and associated nutrient cycling remains unclear. Nitrogen (N) availability shows a site‐specific response to wildfire events indicating the need for an increased understanding of underlying mechanisms that drive changes in soil N cycling. In this study, we selected unburned and burned sites in a large area of the National Park Tolhuaca that was affected by a stand‐replacing wildfire in February 2002. We conducted net N cycling flux measurements (net ammonification, net nitrification and net N mineralization assays) on soils sampled 3 years after fire. In addition, samples were physically fractionated and natural abundance of C and N, and ¹³C‐NMR analyses were performed. Results indicated that standing inorganic N pools were greater in the burned soil, but that no main differences in net N cycling fluxes were observed between unburned and burned sites. In both sites, net ammonification and net nitrification fluxes were low or negative, indicating N immobilization. Multiple linear regression analyses indicated that soil N cycling could largely be explained by two parameters: light fraction (LF) soil organic matter N content and aromatic Chemical Oxidation Resistant Carbon (COREC_arom), a relative measure for char. The LF fraction, a strong NH₄⁺ sink, decreased as a result of fire, while COREC_arom increased in the burned soil profile and stimulated NO₃^‐ production. The absence of increased total net nitrification might relate to a decrease in heterotrophic nitrification after wildfire. We conclude that (i) wildfire induced a shift in N transformation pathways, but not in total net N mineralization, and (ii) stable isotope measurements are a useful tool to assess post‐fire soil organic matter dynamics.     

Changes in leaf nutrient traits in a wildfire chronosequence

The effect of wildfire on ecosystem function is gaining interest since climate change is expected to increase fire frequency and intensity in many forest systems. Fire alters the nutritional status of forest ecosystems, affecting ecosystem function and productivity, but further studies evaluating changes in leaf nutrient traits induced by forest wildfires are still needed. We used a 17-year-old Pinus canariensis wildfire chronosequence to elucidate the nature of nutrient limitations in natural and unmanaged pine forest in the Canary Islands. Pine needles were sampled in winter and spring and analysed for N and P concentrations. As expected, we found the lowest leaf N and leaf P in recently burned plots. However, the leaf N:P ratio was higher in burned versus unburned plots, suggesting that the decrease in P availability due to the fire is larger than that of N. For all leaf traits and sampling dates, leaf trait values in burned plots matched those observed in unburned plots 17 years after a fire. The N:P ratio found in P. canariensis needles was one of the lowest values reported in the literature for woody species, and suggests that all pine trees in the chronosequence are unambiguously limited by low N availability. Our results show that these N-limited pine forests retained N more efficiently than P 4 years after a wildfire; however, leaf N recovery is slower than P recovery, suggesting that the mechanisms responsible for pine N limitation operate continuously in these forests.     

Recovery of forest-floor vegetation after a wildfire in a Picea mariana forest

We aimed to detect the trajectories of forest-floor vegetation recovery in a Picea mariana forest after a wildfire. Since fire severity in boreal forests is expected to increase because of climate changes, we investigated the effects of ground-surface burn severity, a surrogate for overall fire severity, on the revegetation. We annually monitored vegetation <1.3 m high in 80 1 m × 1 m quadrats at Poker Flat Research Range (65°12′N, 147°46′W, 650 m a.s.l.) near Fairbanks, interior Alaska, where a large wildfire occurred in the summer of 2004, from 2005 to 2009. Sphagnum mosses were predominant on the unburned ground surface. In total, 66 % of the ground surface was burned completely by the wildfire. Total plant cover increased from 48 % in 2005 to 83 % in 2009. The increase was derived mostly by the vegetative reproduction of shrubs on the unburned surface and by the immigration of non-Sphagnum mosses and deciduous trees on the burned surface. Deciduous trees, which had not been established before the wildfire, colonized only on the burned surface and grew faster than P. mariana. Although species richness decreased with increasing slope gradient, these deciduous trees became established even on steep slopes. The wildfire that completely burned the ground surface distorted the revegetation, particularly on steep slopes. The restoration of the Sphagnum surface was a prerequisite after the severe wildfire occurred, although the Sphagnum cover had difficulty returning to predominance in the short term.     

Factors Regulating Nitrogen Retention During the Early Stages of Recovery from Fire in Coastal Chaparral Ecosystems

Fire is a fundamental reorganizing force in chaparral and other Mediterranean-type ecosystems. Postfire nutrient redistribution and cycling are frequently invoked as drivers of ecosystem recovery. The extent to which N is transported from slopes to streams following fire is a function of the balance between the rate at which soil microbes retain and metabolize N into forms that readily dissolve or leach, and how rapidly recovering plants sequester this mobilized N. To better understand how fire impacts this balance, we sampled soil and plant N dynamics in 17 plots distributed across two burned, chaparral-dominated watersheds in Santa Barbara County, California. We measured a variety of ecosystem properties in both burned and unburned plots on a periodic basis for 2 years, including soil water content, pH, soil and plant carbon and nitrogen, extractable inorganic nitrogen, dissolved organic nitrogen, and microbial biomass. In burned plots, nitrification was significantly enhanced relative to rates measured in unburned plots. Ephemeral herbs established quickly following the first postfire rain events. Aboveground plant biomass assimilated N commensurate with soil net mineralization, implying tight N cycling during the early stages of recovery. Microbial biomass N, on the other hand, remained low throughout the study. These findings highlight the importance of herbaceous species in conserving ecosystem nutrients as shrubs gradually recover.     

Wildfire Effects on Soil Gross Nitrogen Transformation Rates in Coniferous Forests of Central Idaho, USA

Forest fires often result in a series of biogeochemical processes that increase soil nitrate (NO₃ ^?) concentrations for several years; however, the dynamic nature of inorganic nitrogen (N) cycling in the plant–microbe–soil complex makes it challenging to determine the direct causes of increased soil NO₃ ^?. We measured gross inorganic N transformation rates in mineral soils 2 years after wildfires in three central Idaho coniferous forests to determine the causes of the elevated soil NO₃ ^?. We also measured key factors that could affect the soil N processes, including temperature during soil incubation in situ, soil water content, pH and carbon (C) availability. We found no significant differences (P = 0.461) in gross nitrification rates between burned and control soils. However, microbial NO₃ ^? uptake rates were significantly lower (P = 0.078) in burned than control soils. The reduced consumption of NO₃ ^? caused slightly elevated NO₃ ^? concentrations in the burned soils. C availability was positively correlated with microbial NO₃ ^? uptake rates. Despite reduced microbial NO₃ ^? uptake capacity in the burned soils, soil microbes were a strong enough N sink to maintain low soil NO₃ ^? concentrations 2 years post fire. Soil NH₄ ⁺ concentrations between the treatments were not significantly different (P = 0.673). However, gross NH₄ ⁺ production and microbial uptake rates in burned soils were significantly lower (P = 0.028 and 0.035, respectively) than in the controls, and these rates were positively correlated with C availability. Our results imply that C availability is an important factor regulating soil N cycling of coniferous forests in the region.     

Variation in Aboveground Cover Influences Soil Nitrogen Availability at Fine Spatial Scales Following Severe Fire in Subalpine Conifer Forests

Following fire, fine-scale variation in early successional vegetation and soil nutrients may influence development of ecosystem structure and function. We studied conifer forests burned by stand-replacing wildfire in Greater Yellowstone (Wyoming, USA) to address two questions: (1) How do the variability and spatial structure of aboveground cover and soil nitrogen availability change during the first 4 years following stand-replacing fire? (2) At fine scales (2–20 m), are postfire soil inorganic N pools and fluxes related to aboveground cover? Aboveground cover, soil N pools, and annual net N transformations were measured from 2001 to 2004 using a spatially explicit sampling design in four 0.25-ha plots that burned during summer 2000. Within-stand variability (coefficient of variation) in postfire live vegetative cover declined with time since fire, whereas variability in bare mineral soil, charred litter and fresh litter was greatest 2-3 years postfire. The soil nitrate pool was more variable than the soil ammonium pool, but annual net nitrification was less variable than annual net N mineralization. Spatial structure (quantified by semivariograms) was observed in some aboveground cover variables (for example, graminoids and fresh litter), but there was little spatial structure in soil N variables and no obvious congruence in spatial scales of autocorrelation for soil N and aboveground cover. Significant Spearman correlations (at the sample point) indicated that aboveground cover and soil N were coupled following severe fire, and the dominant influence was from aboveground cover to soil N, rather than from soil N to vegetation. Initial patterns of fire severity and re-vegetation contributed to fine-scale heterogeneity in soil N availability for at least 4 years after severe fire.     

Changes to the N cycle following bark beetle outbreaks in two contrasting conifer forest types

Outbreaks of Dendroctonus beetles are causing extensive mortality in conifer forests throughout North America. However, nitrogen (N) cycling impacts among forest types are not well known. We quantified beetle-induced changes in forest structure, soil temperature, and N cycling in Douglas-fir (Pseudotsuga menziesii) forests of Greater Yellowstone (WY, USA), and compared them to published lodgepole pine (Pinus contorta var. latifolia) data. Five undisturbed stands were compared to five beetle-killed stands (4–5 years post-outbreak). We hypothesized greater N cycling responses in Douglas-fir due to higher overall N stocks. Undisturbed Douglas-fir stands had greater litter N pools, soil N, and net N mineralization than lodgepole pine. Several responses to disturbance were similar between forest types, including a pulse of N-enriched litter, doubling of soil N availability, 30–50 % increase in understory cover, and 20 % increase in foliar N concentration of unattacked trees. However, the response of some ecosystem properties notably varied by host forest type. Soil temperature was unaffected in Douglas-fir, but lowered in lodgepole pine. Fresh foliar %N was uncorrelated with net N mineralization in Douglas-fir, but positively correlated in lodgepole pine. Though soil ammonium and nitrate, net N mineralization, and net nitrification all doubled, they remained low in both forest types (<6 μg N g soil^?1 NH₄ ⁺or NO₃ ^?; <25 μg N g soil^?1 year^?1 net N mineralization; <8 μg N g soil^?1 year^?1 net nitrification). Results suggest that beetle disturbance affected litter and soil N cycling similarly in each forest type, despite substantial differences in pre-disturbance biogeochemistry. In contrast, soil temperature and soil N–foliar N linkages differed between host forest types. This result suggests that disturbance type may be a better predictor of litter and soil N responses than forest type due to similar disturbance mechanisms and disturbance legacies across both host–beetle systems.     

Fire legacies in eastern ponderosa pine forests

Disturbance legacies structure communities and ecological memory, but due to increasing changes in disturbance regimes, it is becoming more difficult to characterize disturbance legacies or determine how long they persist. We sought to quantify the characteristics and persistence of material legacies (e.g., biotic residuals of disturbance) that arise from variation in fire severity in an eastern ponderosa pine forest in North America. We compared forest stand structure and understory woody plant and bird community composition and species richness across unburned, low‐, moderate‐, and high‐severity burn patches in a 27‐year‐old mixed‐severity wildfire that had received minimal post‐fire management. We identified distinct tree densities (high: 14.3 ± 7.4 trees per ha, moderate: 22.3 ± 12.6, low: 135.3 ± 57.1, unburned: 907.9 ± 246.2) and coarse woody debris cover (high: 8.5 ± 1.6% cover per 30 m transect, moderate: 4.3 ± 0.7, low: 2.3 ± 0.6, unburned: 1.0 ± 0.4) among burn severities. Understory woody plant communities differed between high‐severity patches, moderate‐ and low‐severity patches, and unburned patches (all p < 0.05). Bird communities differed between high‐ and moderate‐severity patches, low‐severity patches, and unburned patches (all p < 0.05). Bird species richness varied across burn severities: low‐severity patches had the highest (5.29 ± 1.44) and high‐severity patches had the lowest (2.87 ± 0.72). Understory woody plant richness was highest in unburned (5.93 ± 1.10) and high‐severity (5.07 ± 1.17) patches, and it was lower in moderate‐ (3.43 ± 1.17) and low‐severity (3.43 ± 1.06) patches. We show material fire legacies persisted decades after the mixed‐severity wildfire in eastern ponderosa forest, fostering distinct structures, communities, and species in burned versus unburned patches and across fire severities. At a patch scale, eastern and western ponderosa system responses to mixed‐severity fires were consistent.     

火烧对草地土壤氮总矿化、硝化及无机氮消耗速率的影响

采用同位素^15N库稀释技术结合扰动较小的管型取样法,测定了羊草草地火烧区与未烧区不同季节土壤氮的总矿化速率、总硝化速率、无机氮消耗速率．结果表明,火烧地的氮总矿化与硝化速率在牧草返青后的4、5月份均高于未烧地,7月份差异不显著,到生长季末的9月份又低于未烧地;火烧地NH^4-N的消耗速率7月份以前均高于未烧地,9月份低于未烧地,N03^--N的消耗4、5份火烧地要高于未烧地,7、9月份又低于未烧地;火烧地土壤NH4^ -N含量在4、5和7月份均高于未烧地,9月份基本没有差别,而N03^--N在4、5月份无大差别,7、9月份高于未烧地．     

Soil nitrogen dynamics as an indicator for longleaf pine restoration

Assessing the status of soil nutrients with their corresponding microbial communities provides important information about degraded soils during the restoration of coastal wet pine forests. Net nitrogen mineralization, nitrogen‐oxidizing bacteria (NOB), and soil microbial biomass were compared with patch‐derived volume along a 110‐year longleaf pine (Pinus palustris Mill.) chronosequence for identifying a trajectory and ecological benchmark during forest restoration. Net nitrogen mineralization rates decreased significantly in the maturing‐aged, pine patches, driven by a larger drop in net nitrification. Net nitrification and abundance of NOB were higher in young pine patches compared to soils from the maturing (86–110 years) pine patches. Gross nitrate fluxes followed the nonfungal portion of the soil microbial biomass along the chronosequence, declining in 64‐year‐old pine patches. Microbial biomass peaked in patches 17–34 years of age, but significantly declined in the older patches. Fungal biomass leveled off without decline. Ammonium was the major source of nitrogen within the maturing pine patches as well as the wetland patches, indicating that ammonium maintains longleaf pine during growth‐limiting conditions. Nitrate dominated during rapid tree growth, optimally in mesic conditions. The relative amounts of available ammonium to nitrate can be used to model nitrogen cycling in facultative‐wetland pine forests of the coastal United States as soils alternate between wet and mesic conditions. A key restoration benchmark occurred after 86 years of pine development when pine patch growth rates slowed, with lower numbers of NOB, when the nonfungal biomass leveled off, and net nitrification rates are at a minimum, during pine maturation.     

Post-fire Aleppo pine growth,C and N isotope composition depend on site dryness

Key message

The post-fire growth responses and changes in wood C and N isotope composition depend on site water availability and fire severity in Mediterranean Aleppo pine forests.

Abstract

Mediterranean forests are subjected to recurrent wildfires and summer droughts. Under warmer and drier conditions, it is required to determine how Mediterranean pines recover after wildfires, and how this translates into changes in tree radial growth and function (e.g. intrinsic water-use efficiency—iWUE). We analysed four Aleppo pine areas located in SE Spain affected by 1994 wildfires and subjected to different water availability, ranging from mesic to semi-arid conditions. We combined dendrochronological analyses with δ¹³C and δ¹⁵N wood isotopes to quantify the changes in radial growth (expressed as Basal Area Increment—BAI) and functional responses (iWUE and N cycling) to three fire severities (unburned sites, low and medium severities). We expected that the post-fire release in nutrients and a reduction in competition for water would enhance radial growth. We found that fire did not significantly alter growth patterns at the driest sites, but increased BAI at the wettest sites. δ¹³C was significantly (P ≤ 0.01) more negative only in burned stands located at the wettest site indicating a decreased iWUE and thus improved water availability. However, the δ¹⁵N was higher in severely burned than in unburned plots from all sites but the wettest site, indicating a potential fertilization effect of fire in sites subjected to mild drought severity. Site water availability determined how fire affected subsequent modifications in growth and tree functioning of Aleppo pine forests, that is, changes in iWUE and N cycling. Therefore, site dryness should be explicitly considered to forecast the growth and functioning responses of Mediterranean pine forests to the predicted increasing recurrence of fire events due to global warming.

     

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

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

Correlation between leaf litter and fine root decomposition among subtropical tree species

Sampling disturbance has been shown to rapidly increase net nitrification rates in some forest soils. To gain insight on mechanisms, we investigated both gross and net rates of ammonification and nitrification in intact cores and mixed composite samples. Using the isotope pool dilution method, we studied samples from two northeastern USA watersheds, Brush Brook and Sleepers River in Vermont, where previous work had found high net nitrification rates. Gross ammonification was usually not significantly different between intact cores and mixed samples. However, gross and net nitrification rates in mixed samples were similar (mean ～24?µmol N?kg^?1?hr^?1 or ～8 mg N kg^?1 d^?1) and significantly higher than in intact cores (7.7 and 3.4?µmol N kg^?1?h^?1 for means of gross and net respectively). Nitrate consumption was decreased somewhat by disturbance but did not account for the large differences in net rates. Because there were similar gross ammonification rates in both treatments, increased nitrification in these disturbed soils must be a result of an increase in the utilization of ammonium by the ammonia oxidizers at the expense of other ammonium consumption pathways. Different mechanisms may operate in different soils; increased nitrification appears to be the primary pathway in these soils with high N cycling rates.     

In situ gross nitrogen transformations differ between temperate deciduous and coniferous forest soils

Despite long-term enhanced nitrogen (N) inputs, forests can retain considerable amounts of N. While rates of N inputs via throughfall and N leaching are increased in coniferous stands relative to deciduous stands at comparable sites, N leaching below coniferous stands is disproportionally enhanced relative to the N input. A better understanding of factors affecting N retention is needed to assess the impact of changing N deposition on N cycling and N loss of forests. Therefore, gross N transformation pathways were quantified in undisturbed well-drained sandy soils of adjacent equal-aged deciduous (pedunculate oak (Quercus robur L.)) and coniferous (Scots pine (Pinus sylvestris L.)) planted forest stands located in a region with high N deposition (north Belgium). In situ inorganic ¹⁵N labelling of the mineral topsoil (0–10?cm) combined with numerical data analysis demonstrated that (i) all gross N transformations differed significantly (p?<?0.05) between the two forest soils, (ii) gross N mineralization in the pine soil was less than half the rate in the oak soil, (iii) meaningful N immobilization was only observed for ammonium, (iv) nitrate production via oxidation of organic N occurred three times faster in the pine soil while ammonium oxidation was similar in both soils, and (v) dissimilatory nitrate reduction to ammonium was detected in both soils but was higher in the oak soil. We conclude that the higher gross nitrification (including oxidation of organic N) in the pine soil compared to the oak soil, combined with negligible nitrate immobilization, is in line with the observed higher nitrate leaching under the pine forest.     

Stand-replacing wildfires alter the community structure of wood-inhabiting fungi in southwestern ponderosa pine forests of the USA

Increases in stand-replacing wildfires in the western USA have widespread implications for ecosystem carbon (C) cycling, in part because the decomposition of trees killed by fire can be a long-term source of CO₂ to the atmosphere. Knowledge of the composition and function of decay fungi communities may be important to understanding how wildfire alters C cycles. We assessed the effects of stand-replacing wildfires on the community structure of wood-inhabiting fungi along a 32-yr wildfire chronosequence. Fire was associated with low species richness for up to 4 yr and altered species composition relative to unburned forest for the length of the chronosequence. A laboratory incubation demonstrated that species varied in their capacity to decompose wood; Hypocrea lixii, an indicator of the most recent burn, caused the lowest decomposition rate. Our results show that stand-replacing wildfires have long-term effects on fungal communities, which may have consequences for wood decomposition and C cycling.     

Nitrogen-15 signals of leaf-litter-soil continuum as a possible indicator of ecosystem nitrogen saturation by forest succession and N loads

Understanding forest carbon cycling responses to atmospheric N deposition is critical to evaluating ecosystem N dynamics. The natural abundance of ¹⁵N (??¹⁵N) has been suggested as an efficient and non-invasive tool to monitor N pools and fluxes. In this study, three successional forests in southern China were treated with four levels of N addition. In each treatment, we measured rates of soil N mineralization, nitrification, N₂O emission and inorganic N leaching as well as N concentration and ?? ¹⁵N of leaves, litters and soils. We found that foliar N concentration and ??¹⁵N were higher in the mature broadleaf forest than in the successional pine or mixed forests. Three-year continuous N addition did not change foliar N concentration, but significantly increased foliar ?? ¹⁵N (p < 0.05). Also, N addition decreased the ?? ¹⁵N of top soil in the N-poor pine and mixed forests and significantly increased that of organic and mineral soils in N-rich broadleaf forests (p < 0.05). In addition, the soil N₂O emission flux and inorganic N leaching rate increased with increasing N addition and were positively correlated with the ¹⁵N enrichment factor (?? _p/s) of forest ecosystems. Our study indicates that ?? ¹⁵N of leaf, litter and soil integrates various information on plant species, forest stand age, exogenous N input and soil N transformation and loss, which can be used to monitor N availability and N dynamics in forest ecosystems caused by increasing N deposition in the future.     

Responses of nitrous oxide fluxes and soil nitrogen cycling to nutrient additions in montane forests along an elevation gradient in southern Ecuador

Tropical montane forests are commonly limited by N or co-limited by N and P. Projected increases in N deposition in tropical montane regions are thought to be insufficient for vegetation demand and are not therefore expected to affect soil N availability and N₂O emissions. We established a factorial N- and P-addition experiment (i.e., N, P, N + P, and control) across an elevation gradient of montane forests in Ecuador to test these hypotheses: (1) moderate rates of N and P additions are able to stimulate soil-N cycling rates and N₂O fluxes, and (2) the magnitude and timing of soil N₂O-flux responses depend on the initial nutrient status of the forest soils. Moderate rates of nutrients were added: 50 kg N ha^?1 year^?1 (in the form of urea) and 10 kg P ha^?1 year^?1 (in the form of NaH₂PO ₄ ^. 2H₂O) split in two equal applications. We tested the hypotheses by measuring changes in net rates of soil–N cycling and N₂O fluxes during the first 2 years (2008–2009) of nutrient manipulation in an old-growth premontane forest at 1,000 m, growing on a Cambisol soil with no organic layer, in an old-growth lower montane forest at 2,000 m, growing on a Cambisol soil with an organic layer, and an old-growth upper montane rainforest at 3,000 m, growing on a Histosol soil with a thick organic layer. Among the control plots, net nitrification rates were largest at the 1,000-m site whereas net nitrification was not detectable at the 2,000- and 3,000-m sites. The already large net nitrification at the 1,000-m site was not affected by nutrient additions, but net nitrification became detectable at the 2,000- and 3000-m sites after the second year of N and N + P additions. N₂O emissions increased rapidly following N and N + P additions at the 1,000-m site whereas only smaller increases occurred at the 2,000- and 3,000-m sites during the second year of N and N + P additions. Addition of P alone had no effect on net rates of soil N cycling and N₂O fluxes at any elevation. Our results showed that the initial soil N status, which may also be influenced by presence or absence of organic layer, soil moisture and temperature as encompassed by the elevation gradient, is a good indicator of how soil N cycling and N₂O fluxes may respond to future increases in nutrient additions.     

The establishment patterns of tree seedlings are determined immediately after wildfire in a black spruce (Picea mariana) forest

Fire severity is predicted to increase in boreal regions due to global warming. We hypothesized that these extreme events will alter regeneration patterns of black spruce (Picea mariana). To test this hypothesis, we monitored seed dispersal and seedling emergence, survival and growth for 6 years from 2005 to 2010 after the 2004 wildfire on Poker Flat, interior Alaska, using 96 1 × 1 m plots. A total of 1,300 seedlings of black spruce and three broad-leaved deciduous trees (Populus tremuloides, Betula papyrifera, and Salix spp.) were recorded. Black spruce seedlings colonized burned and unburned ground surfaces for the first 2 years after the wildfire and established on any topographical surface, while the broad-leaved trees emerged less in areas of lower elevation, slope gradient and canopy openness and only on burned surfaces. Vascular plant cover on the ground floor increased the seedling establishment of black spruce and broad-leaved trees, most likely because of seed-trap effects. Black spruce grew faster on burned surface than on unburned surfaces. However, broad-leaved trees grew faster than black spruce on burned surfaces. Black spruce regenerates even after severe wildfire when the microtopography restricts the colonization of broad-leaved trees. The regeneration trajectories are determined soon after wildfire by a combination of seed limitation for black spruce and habitat preference for broad-leaved trees.