Nitrogen supply effects on leaf dynamics and nutrient input into the soil of plant species in a sub-arctic tundra ecosystem

Global warming will lead to increased nitrogen supply in tundra ecosystems. How increased N supply affected leaf production, leaf turnover and dead leaf N input into the soil of Empetrum nigrum and Andromeda polifolia (evergreens), Eriophorum vaginatum (graminoid) and Betula nana (deciduous) in a sub-arctic tundra in northern Sweden between 2003 and 2007 was experimentally investigated. There was considerable interspecific variation in the response of leaf production to N addition, varying from negative, no response to a positive response. Nitrogen addition effects on leaf turnover also showed considerable variation among species, varying from no effect to increased leaf turnover (up to 27% in Eriophorum). Nitrogen addition resulted in a four to fivefold increase in N content in the dead leaves of both evergreens and a 65% increase in Eriophorum. Surprisingly, there was no increase in Betula. The response of dead leaf P contents to N addition was rather species specific. There was no response in Empetrum, whereas there were significant increases in Andromeda (+214%) and Eriophorum (+32%), and a decrease of 47% in Betula. As an overall result of the changes in leaf production, leaf turnover and dead leaf N and P contents, nitrogen addition increased in all species except Betula the amount of N and, for Andromeda and Eriophorum the amount of P transferred to the soil due to leaf litter inputs. However, the way in which this was achieved differed substantially among species due to interspecific differences in the response of the component processes (leaf production, leaf turnover, dead leaf nutrient content).     

Seasonal climate manipulations have only minor effects on litter decomposition rates and N dynamics but strong effects on litter P dynamics of sub-arctic bog species

Litter decomposition and nutrient mineralization in high-latitude peatlands are constrained by low temperatures. So far, little is known about the effects of seasonal components of climate change (higher spring and summer temperatures, more snow which leads to higher winter soil temperatures) on these processes. In a 4-year field experiment, we manipulated these seasonal components in a sub-arctic bog and studied the effects on the decomposition and N and P dynamics of leaf litter of Calamagrostis lapponica, Betula nana, and Rubus chamaemorus, incubated both in a common ambient environment and in the treatment plots. Mass loss in the controls increased in the order Calamagrostis < Betula < Rubus. After 4 years, overall mass loss in the climate-treatment plots was 10 % higher compared to the ambient incubation environment. Litter chemistry showed within each incubation environment only a few and species-specific responses. Compared to the interspecific differences, they resulted in only moderate climate treatment effects on mass loss and these differed among seasons and species. Neither N nor P mineralization in the litter were affected by the incubation environment. Remarkably, for all species, no net N mineralization had occurred in any of the treatments during 4 years. Species differed in P-release patterns, and summer warming strongly stimulated P release for all species. Thus, moderate changes in summer temperatures and/or winter snow addition have limited effects on litter decomposition rates and N dynamics, but summer warming does stimulate litter P release. As a result, N-limitation of plant growth in this sub-arctic bog may be sustained or even further promoted.     

Foliage litter quality and annual net N mineralization: comparison across North American forest sites

The feedback between plant litterfall and nutrient cycling processes plays a major role in the regulation of nutrient availability and net primary production in terrestrial ecosystems. While several studies have examined site-specific feedbacks between litter chemistry and nitrogen (N) availability, little is known about the interaction between climate, litter chemistry, and N availability across different ecosystems. We assembled data from several studies spanning a wide range of vegetation, soils, and climatic regimes to examine the relationship between aboveground litter chemistry and annual net N mineralization. Net N mineralization declined strongly and non-linearly as the litter lignin:N ratio increased in forest ecosystems (r ² = 0.74, P < 0.01). Net N mineralization decreased linearly as litter lignin concentration increased, but the relationship was significant (r ² = 0.63, P < 0.01) only for tree species. Litterfall quantity, N concentration, and N content correlated poorly with net N mineralization across this range of sites (r ² < 0.03, P = 0.17–0.26). The relationship between the litter lignin:N ratio and net N mineralization from forest floor and mineral soil was similar. The litter lignin:N ratio explained more of the variation in net N mineralization than climatic factors over a wide range of forest age classes, suggesting that litter quality (lignin:N ratio) may exert more than a proximal control over net N mineralization by influencing soil organic matter quality throughout the soil profile independent of climate. Received: 16 December 1996 / Accepted: 8 February 1997     

Influence of the phenolic compound bearing species Ledum palustre on soil N cycling in a boreal hardwood forest

The effects of the understory shrub Ledum palustre on soil N cycling were studied in a hardwood forest of Interior Alaska. This species releases high concentrations of phenolic compounds from green leaves and decomposing litter by rainfall. Organic and mineral soils sampled underneath L. palustre and at nearby non-Ledum sites were amended with L. palustre litter leachates and incubated at controlled conditions. We aimed to know (i) whether L. palustre presence and litter leachate addition changed net N cycling rates in organic and mineral soils, and (ii) what N cycling processes, including gross N mineralization, N immobilization and gross N nitrification, were affected in association with L. palustre. Our results indicate that N transformation rates in the surface organic horizon were not affected by L. palustre presence or leachate addition. However, mineral soils underneath L. palustre as well as soils amended with leachates had significantly higher C/N ratios and microbial respiration rates, and lower net N mineralization and N-to-C mineralization compared to no Ledum and no leachates soils. No nitrification was detected. Plant presence and leachate addition also tended to increase both gross N mineralization and immobilization. These results suggest that soluble C compounds present in L. palustre increased N immobilization in mineral soils when soil biota used them as a C source. Increases in gross N mineralization may have been caused by an enhanced microbial biomass due to C addition. Since both plant presence and leachate addition decreased soil C/N ratio and had similar effects on N transformation rates, our results suggest that litter leachates could be partially responsible for plant presence effects. The lower N availability under L. palustre canopy could exert negative interactions on the establishment and growth of other plant species.     

Mineralization and Decomposition Rates in Restored Pine Fens

Growing public interest in conserving peatlands has created a need for restoration and rapid indicators of progress in peat formation. Vegetation and hydrological indicators are commonly assessed, but changes in mineralization and decomposition rates might better indicate when peat formation is underway in restored peatlands. In Finland, we investigated differences in mineralization and decomposition in the upper peat layer of five undrained and eight drained Pinus‐dominated fens from 2006 to 2009. Forestry‐drained fens were restored in 2007 by harvesting either whole trees or only stems, and by damming and filling ditches. Before restoration, net N mineralization rate was slightly higher in the drained than in undrained fens, whereas soil pH and Betula leaf litter decomposition rate were lower. After restoration, net N mineralization rate was similar for the undrained and restored fens, except near ditches after stem harvest. Also, soil pH and decomposition rate of Betula leaf litter became similar for undrained and restored fens. We conclude that whole tree harvest is a more suitable method for peatland restoration than stem harvest and that mineralization and decomposition rates are suitable indicators for peat formation after restoration.     

Effects of understory removal, N fertilization, and litter layer removal on soil N cycling in a 13-year-old white spruce plantation infested with Canada bluejoint grass

Canada bluejoint grass [Calamagrostis canadensis (Michx.) Beauv., referred to as bluejoint below] is a competitive understory species widely distributed in the boreal region in North America and builds up a thick litter layer that alters the soil surface microclimate in heavily infested sites. This study examined the effects of understory removal, N fertilization, and litter layer removal on litter decomposition, soil microbial biomass N (MBN), and net N mineralization and nitrification rates in LFH (the sum of organic horizons of litter, partially decomposed litter and humus on the soil surface) and mineral soil (0–10 cm) in a 13-year-old white spruce [Picea glauca (Moench.) Voss] plantation infested with bluejoint in Alberta, Canada. Removal of the understory vegetation and the litter layer together significantly increased soil temperature at 10 cm below the mineral soil surface by 1.7 and 1.3°C in summer 2003 and 2004, respectively, resulting in increased net N mineralization (by 1.09 and 0.14 mg N kg⁻¹ day⁻¹ in LFH and mineral soil, respectively, in 2004) and net nitrification rates (by 0.10 and 0.20 mg N kg⁻¹ day⁻¹ in LFH and mineral soil, respectively, in 2004). When the understory vegetation was intact, nitrification might have been limited by NH₄ ⁺ availability due to competition for N from bluejoint and other understory species. Litter layer removal increased litter decomposition rate (percentage mass loss per month) from 2.6 to 3.0% after 15 months of incubation. Nitrogen fertilization did not show consistent effects on soil MBN, but increased net N mineralization and nitrification rates as well as available N concentrations in the soil. Clearly, understory removal combined with N fertilization was most effective in increasing rates of litter decomposition, net N mineralization and nitrification, and soil N availability. The management of understory vegetation dominated by bluejoint in the boreal region should consider the strong effects of understory competition and the accumulated litter layer on soil N cycling and the implications for forest management.     

The Effect of Phosphorus Availability on Decomposition Dynamics in a Seasonal Lowland Amazonian Forest

Once the weathering of parent material ceases to supply significant inputs of phosphorus (P), vegetation depends largely on the decomposition of litter and soil organic matter and the associated mineralization of organic P forms to provide an adequate supply of this essential nutrient. At the same time, the decomposition of litter is often characterized by the immobilization of nutrients, suggesting that nutrient availability is a limiting factor for this process. Immobilization temporally decouples nutrient mineralization from decomposition and may play an important role in nutrient retention in low-nutrient ecosystems. In this study, we used a common substrate to study the effects of native soil P availability as well as artificially elevated P availability on litter decomposition rates in a lowland Amazonian rain forest on highly weathered soils. Although both available and total soil P pools varied almost three fold across treatments, there was no significant difference in decomposition rates among treatments. Decomposition was rapid in all treatments, with approximately 50% of the mass lost over the 11-month study period. Carbon (C) and nitrogen (N) remaining and C:N ratios were the most effective predictors of amount of mass remaining at each time point in all treatments. Fertilized treatments showed significant amounts of P immobilization (P < 0.001). By the final collection point, the remaining litter contained a quantity equivalent to two-thirds of the initial P and N, even though only half of the original mass remained. In these soils, immobilization of nutrients in the microbial biomass, late in the decomposition process, effectively prevents the loss of essential nutrients through leaching or occlusion in the mineral soil.     

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

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

植被类型与坡位对喀斯特土壤氮转化速率的影响

土壤氮素转化对于植物氮素营养具有重要作用,尤其是对于受氮素限制的喀斯特退化生态系统。选取植被恢复过程中4种典型喀斯特植被类型(草丛、灌丛、次生林、原生林)和3个坡位(上、中、下坡位)表层土壤(0—15cm)为对象,利用室内培养的方法,研究不同植被类型和坡位下土壤氮素养分与氮转化速率(氮净矿化率、净硝化率和净氨化率)的特征及其影响因素。结果表明,植被类型对土壤硝态氮含量、无机氮含量、氮净矿化率、净硝化率和净氨化率均有显著影响(P0.01),即随着植被的正向演替(草丛—灌丛—次生林—原生林),土壤硝态氮含量、无机氮含量、土壤氮净矿化速率和净硝化速率整体上呈增加趋势,而坡位以及坡位与植被类型的交互作用对上述土壤氮素指标无显著影响(P0.05)。冗余分析结果表明凋落物氮含量、凋落物C∶N比和硝态氮含量对土壤氮转化速率有显著影响,其中凋落物氮含量是影响土壤氮转化速率的主要因子(F=35.634,P=0.002)。可见,尽管坡位影响喀斯特水土再分配过程,但植被类型决定的凋落物质量(如凋落物氮含量等)对喀斯特土壤氮素转化速率的作用更为重要。因此,在喀斯特退化生态系统植被恢复初期,应注重植被群落的优化配置(如引入豆科植物)和土壤质量的改善(如降低土壤C∶N),促进土壤氮素转化及氮素的有效供给。     

Factors affecting nitrogen dynamics in a semiarid woodland (Dry Chaco,Argentina)

In an effort to elucidate the factors affecting soil N dynamics in the Dry Chaco ecosystem, soil respiration and microbial biomass N were measured for one year underneath 5 vegetation types: a leguminous tree (Prosopis flexuosa DC), a non-leguminous tree (Aspidosperma quebracho-blanco Schlecht.), a non leguminous shrub (Larrea spp.), the open interspaces, and a pure grassland. Ammonifier and nitrifier densities and N content in litter were also measured in some cases. Results were compared with previously reported N mineralization rates and soil fertility.During the dry season microbial biomass N and net N mineralization were low, while accretion of easily mineralizable C occurred (estimated through soil respiration rates in lab under controlled temperature and moisture). With the onset of rain, microbial biomass N and N mineralization increased markedly, resulting in a decrease in easily mineralizable C. Throughout the wet season N mineralization varied with soil moisture while microbial biomass N remained consistently high. Mean values of immobilized N in this ecosystem were high (20–140 mg kg^–1), of about the same order of magnitude as accumulated net N mineralization (50–150 mg kg^–1 yr^–1). Microbial decay in the dry season, considered as a source of easily mineralizable N, accounted for only 40% of gross N mineralization increase at the beginning of the wet season. Ammonifier densities correlated significantly with soil moisture and N mineralization, but nitrifiers did not.The highest values of total N, N mineralization, inorganic N, microbial biomass N, nitrifier densities, N content in litter, total organic C and easily mineralizable C were found under Prosopis and the lowest values under shrubs and the interspaces. The main differences between tree species were in N mineralization at the beginning of the wet season, in total and inorganic N pools, and in nitrifier densities; all of which were significantly lower under Aspidosperma than under Prosopis.N mineralization in the pure grassland was very low despite high values of total N and C sources. Although N immobilized in microbial biomass was similarly high under Aspidosperma, Prosopis and the pure grassland, net N mineralization rates were quite different.     

Decomposition and nitrogen dynamics of 15N-labeled leaf, root, and twig litter in temperate coniferous forests

Litter nutrient dynamics contribute significantly to biogeochemical cycling in forest ecosystems. We examined how site environment and initial substrate quality influence decomposition and nitrogen (N) dynamics of multiple litter types. A 2.5-year decomposition study was installed in the Oregon Coast Range and West Cascades using ¹⁵N-labeled litter from Acer macrophyllum, Picea sitchensis, and Pseudotsuga menziesii. Mass loss for leaf litter was similar between the two sites, while root and twig litter exhibited greater mass loss in the Coast Range. Mass loss was greatest from leaves and roots, and species differences in mass loss were more prominent in the Coast Range. All litter types and species mineralized N early in the decomposition process; only A. macrophyllum leaves exhibited a net N immobilization phase. There were no site differences with respect to litter N dynamics despite differences in site N availability, and litter N mineralization patterns were species-specific. For multiple litter × species combinations, the difference between gross and net N mineralization was significant, and gross mineralization was 7–20 % greater than net mineralization. The mineralization results suggest that initial litter chemistry may be an important driver of litter N dynamics. Our study demonstrates that greater amounts of N are cycling through these systems than may be quantified by only measuring net mineralization and challenges current leaf-based biogeochemical theory regarding patterns of N immobilization and mineralization.     

Differential Controls of Water Input on Litter Decomposition and Nitrogen Dynamics in the Patagonian Steppe

Studies of the effects of precipitation on litter decomposition and nitrogen mineralization in arid and semiarid environments have demonstrated contradictory results. We conducted a manipulative experiment with rainout shelters in the semiarid Patagonian steppe, aimed at assessing the direct effects of water availability on litter decomposition and net nitrogen mineralization while isolating the indirect effects. We created four levels of precipitation input: control and three levels (30, 55 and 80%) of precipitation interception and we examined litter decomposition and nutrient release of a dominant grass species, Stipa speciosa, inorganic soil nitrogen, and in situ net nitrogen mineralization over two consecutive years. Litter decomposition rates (k, year⁻¹) varied significantly (P < 0.001) among precipitation interception treatments and were positively correlated with incoming annual precipitation (APPT, mm/year) (k = 0.0007 × APPT + 0.137). In contrast, net N mineralization was not correlated with incoming precipitation. Soil NO₃⁻ significantly decreased with increasing precipitation input, whereas soil NH₄⁺ concentration did not differ among precipitation interception treatments. Controls of water input on litter decomposition appear to be different from controls on N mineralization in the semiarid Patagonian steppe. We suggest that although water availability affects both the mineralization of C and N, it differentially affects the movement and fate of the inorganic products. A consequence of the accumulation of inorganic N during dry episodes is that periods of maximum water and soil nutrient availability may occur at different times. This asynchrony in the availability of N and water in the soil may explain the observed lags in the response of primary production to increases in water availability.     

Nitrogen transformation rates are affected by cover soil type but not coarse woody debris application in reclaimed oil sands soils

Forest floor mineral soil mix (FMM) and peat mineral soil mix (PMM) are cover soils commonly used for reclamation of open‐pit oil sands mining disturbed land in northern Alberta, Canada; coarse woody debris (CWD) is another source of organic matter for land reclamation. We investigated net nitrogen (N) transformation rates in FMM and PMM cover soils near and away from CWD 4–6 years after oil sands reclamation. Monthly net nitrification and N mineralization rates varied over time; however, mean rates across the incubation periods and microbial biomass were greater (p < 0.05) in FMM than in PMM. Net N mineralization rates were positively related to soil temperature (p < 0.001) and microbial biomass carbon (p = 0.045). Net N transformation rates and inorganic N concentrations were not affected by CWD; however, the greater ¹⁵N isotope ratio of ammonium near CWD than away from CWD indicates that CWD application increased both gross N mineralization/nitrification (causing N isotope fractionation) and gross N immobilization (no isotopic fractionation). Microbial biomass was greater near CWD than away from CWD, indicating the greater potential for N immobilization near CWD. We conclude that (1) CWD application affected soil microbial properties and would create spatial variability and diverse microsites and (2) cover soil type and CWD application had differential effects on net N transformation rates. Applying FMM with CWD for oil sands reclamation is recommended to increase N availability and microsites.     

Leaf litter production and decomposition in a poplar short-rotation coppice exposed to free air CO2 enrichment (POPFACE)

The capacity of forest ecosystems to sequester C in the soil relies on the net balance between litter production above, as well as, below ground, and decomposition processes. Nitrogen mineralization and its availability for plant growth and microbial activity often control the speed of both processes. Litter production, decomposition and N mineralization are strongly interdependent. Thus, their responses to global environmental changes (i.e. elevated CO₂, climate, N deposition, etc.) cannot be fully understood if they are studied in isolation. In the present experiment, we investigated litter fall, litter decomposition and N dynamics in decomposing litter of three Populus spp., in the second and third growing season of a short rotation coppice under FACE. Elevated CO₂ did not affect annual litter production but slightly retarded litter fall in the third growing season. In all species, elevated CO₂ lowered N concentration, resulting in a reduction of N input to the soil via litter fall, but did not affect lignin concentrations. Litter decomposition was studied in bags incubated in situ both in control and FACE plots. Litter lost between 15% and 18% of the original mass during the eight months of field incubation. On average, litter produced under elevated CO₂ attained higher residual mass than control litter. On the other end, when litter was incubated in FACE plots it exhibited higher decay rates. These responses were strongly species‐specific. All litter increased their N content during decomposition, indicating immobilization of N from external sources. Independent of the initial quality, litter incubated on FACE soils immobilized less N, possibly as a result of lower N availability in the soil. Indeed, our results refer to a short‐term decomposition experiment. However, according to a longer‐term model extrapolation of our results, we anticipate that in Mediterranean climate, under elevated atmospheric CO₂, soil organic C pool of forest ecosystems may initially display faster turnover, but soil N availability will eventually limit the process.     

Nitrogen‐dependent recovery of subarctic tundra vegetation after simulation of extreme winter warming damage to Empetrum hermaphroditum

Vast areas of (sub)arctic tundra are dominated by the ericoid dwarf shrub Empetrum hermaphroditum. Recent experimental and observational data have shown that Empetrum can be damaged heavily by recurrent extreme winter warming. In addition, summer warming leads to increased soil N availability in tundra ecosystems. In a 7‐year experiment, I investigated the recovery of subarctic Empetrum‐dominated tundra vegetation using a factorial combination of various degrees of aboveground Empetrum removal (simulating the damaging effects of extreme winter warming) and N addition (simulating one of the effects of summer warming). After 7 years no new species had established in the plots. The growth of planted Betula nana seedlings was stimulated by Empetrum removal and reduced by N addition. This Empetrum‐dominated tundra ecosystem was resilient against severe disturbances. Only when Empetrum was 100% removed did it fail to recover, and only in combination with high N supply the subordinate species (notably Eriophorum vaginatum and Rubus chamaemorus, a graminoid and a forb) could benefit. In the 50% removal treatment Empetrum recovered in 7 years when no N was supplied and the cover of the subordinate species did not change. However, when N was added Empetrum recovered faster (in 4 years) and the subordinates decreased. When Empetrum was not removed and N was added, Empetrum even increased in abundance at the expense of the subordinate species. Thus, profound changes in tundra ecosystems can only be expected when Empetrum is very heavily damaged as a result of recurrent extreme winter warming and when soil N availability is increased as a result of summer warming. These changes in species composition upon extreme disturbance events may lead to a wide variety of ecosystem feedbacks and cascade processes as this tundra system is relatively species‐poor, and can be hypothesized to have low functional redundancy.     

晋西北不同年限小叶锦鸡儿灌丛土壤氮矿化和硝化作用

利用PVC管顶盖埋管法研究了晋西北黄土高原区小叶锦鸡儿人工灌丛不同定植年限(5,10,20,30,40a)土壤氮矿化与硝化速率的动态和净矿化与硝化总量。结果表明,⑴小叶锦鸡儿灌丛土壤无机氮主要以NO_-~3-N形式存在,不同生长年限相同月份的土壤硝态氮(NO-3-N)含量分别是铵态氮(NH+4-N)含量的1.5—15.4倍;⑵土壤氮素硝化速率和矿化速率随生长年限延长而加快,30年生时达到高峰,数值达40.2,44.1 mg m~(-2)d~(-1)。从季节性变化看,7—8月份是硝化速率和矿化速率快速增长期,30年生小叶锦鸡儿灌丛土壤硝化速率和矿化速率分别达到86.9,93.1 mg m~(-2)d~(-1),显著高于其它生长年限(P0.05);(3)土壤氮素硝化与矿化总量同样随小叶锦鸡儿生长年限延长而增加,30年生时达到最高,与5年生相比,分别增加了3.7和3.1倍。(4)5—10月份小叶锦鸡儿生长期内,各年限土壤全氮量的2.3%被矿化成无机氮,其中87%最终被转化成NO-3-N形式存在于土体中。     

Original Articles: Differential Influence of Plant Species on Soil Nitrogen Transformations Within Moist Meadow Alpine Tundra

Plant species can influence nitrogen (N) cycling indirectly through the feedbacks of litter quality and quantity on soil N transformation rates. The goal of this research was to focus on small-scale (within-community) variation in soil N cycling associated with two community dominants of the moist meadow alpine tundra. Within this community, the small-scale patchiness of the two most abundant species (Acomastylis rossii and Deschampsia caespitosa) provides natural variation in species cover within a relatively similar microclimate, thus enabling estimation of the effects of plant species on soil N transformation rates. Monthly rates of soil N transformations were dependent on small-scale variation in both soil microclimate and species cover. The relative importance of species cover compared with soil microclimate increased for months 2 and 3 of the 3-month growing season. Growing-season net N mineralization rates were over ten times greater and nitrification rates were four times greater in Deschampsia patches than in Acomastylis patches. Variability in litter quality [carbon:nitrogen (C:N) and phenolic:N], litter quantity (aboveground and fine-root production), and soil quality (C:N) was associated with three principal components. Variability between the species in litter quality and fine-root production explained 31% of the variation in net N mineralization rates and 36% of net nitrification rates. Site variability across the landscape in aboveground production and soil C:N explained 33% of the variation in net N mineralization rates and 21% of net nitrification rates. Within the moist meadow community, the high spatial variability in soil N transformation rates was associated with differences in the dominant species' litter quality and fine-root production. Deschampsia-dominated patches consistently had greater soil N transformation rates than did Acomastylis-dominated patches across the landscape, despite site variability in soil moisture, soil C:N, and aboveground production. Plant species appear to be an important control of soil N transformation in the alpine tundra, and consequently may influence plant community structure and ecosystem function.     

Impacts of the exotic,nitrogen-fixing black locust (Robinia pseudoacacia) on nitrogen-cycling in a pine–oak ecosystem

We investigated the influence of the exotic nitrogen-fixing black locust (Robinia pseudoacacia) on nitrogen cycling in a pitch pine (Pinus rigida) −scrub oak (Quercus ilicifolia, Q. prinoides) ecosystem. Within paired pine-oak and adjacent black locust stands that were the result of a 20-35 year-old invasion, we evaluated soil nutrient contents, soil nitrogen transformation rates, and annual litterfall biomass and nitrogen concentrations. In the A horizon, black locust soils had 1.3-3.2 times greater nitrogen concentration relative to soils within pine-oak stands. Black locust soils also had elevated levels of P and Ca, net nitrification rates and total net N-mineralization rates. Net nitrification rates were 25-120 times greater in black locust than in pine-oak stands. Elevated net N-mineralization rates in black locust stands were associated with an abundance of high nitrogen, low lignin leaf litter, with 86 kg N ha^–1 yr^–1 in leaf litter returned compared with 19 kg N ha^–1 yr^–1 in pine-oak stands. This difference resulted from a two-fold greater litterfall mass combined with increased litter nitrogen concentration in black locust stands (1.1% and 2.6% N for scrub oak and black locust litter, respectively). Thus, black locust supplements soil nitrogen pools, increases nitrogen return in litterfall, and enhances soil nitrogen mineralization rates when it invades nutrient poor, pine-oak ecosystems. This revised version was published online in June 2006 with corrections to the Cover Date.     

The Response of Soil Processes to Climate Change: Results from Manipulation Studies of Shrublands Across an Environmental Gradient

Predicted changes in climate may affect key soil processes such as respiration and net nitrogen (N) mineralization and thus key ecosystem functions such as carbon (C) storage and nutrient availability. To identify the sensitivity of shrubland soils to predicted climate changes, we have carried out experimental manipulations involving ecosystem warming and prolonged summer drought in ericaceous shrublands across a European climate gradient. We used retractable covers to create artificial nighttime warming and prolonged summer drought to 20-m² experimental plots. Combining the data from across the environmental gradient with the results from the manipulation experiments provides evidence for strong climate controls on soil respiration, net N mineralization and nitrification, and litter decomposition. Trends of 0%–19% increases of soil respiration in response to warming and decreases of 3%–29% in response to drought were observed. Across the environmental gradient and below soil temperatures of 20°C at a depth of 5–10 cm, a mean Q₁₀ of 4.1 in respiration rates was observed although this varied from 2.4 to 7.0 between sites. Highest Q₁₀ values were observed in Spain and the UK and were therefore not correlated with soil temperature. A trend of increased accumulated surface litter mass loss was observed with experimental warming (2%– 22%) but there was no consistent response to experimental drought. In contrast to soil respiration and decomposition, variability in net N mineralization was best explained by soil moisture rather than temperature. When water was neither limiting or in excess, a Q₁₀ of 1.5 was observed for net N mineralization rates. These data suggest that key soil processes will be differentially affected by predicted changes in rainfall pattern and temperature and the net effect on ecosystem functioning will be difficult to predict without a greater understanding of the controls underlying the sensitivity of soils to climate variables.     

Nitrogen cycling in a northern hardwood forest: Do species matter?

To investigate the influence of individual tree species on nitrogen (N) cycling in forests, we measured key characteristics of the N cycle in small single-species plots of five dominant tree species in the Catskill Mountains of New York State. The species studied were sugar maple (Acer saccharum), American beech (Fagus grandifolia), yellow birch (Betula alleghaniensis), eastern hemlock (Tsuga canadensis), and red oak (Quercus rubra). The five species varied markedly in N cycling characteristics. For example, hemlock plots consistently showed characteristics associated with "slow" N cycling, including low foliar and litter N, high soil C:N, low extractable N pools, low rates of potential net N mineralization and nitrification and low NO ₃ ^– amounts trapped in ion-exchange resin bags buried in the mineral soil. Sugar maple plots had the lowest soil C:N, and the highest levels of soil characteristics associated with NO ₃ ^– production and loss (nitrification, extractable NO ₃ ^– , and resin bag NO ₃ ^– ). In contrast, red oak plots had near-average net mineralization rates and soil C:N ratios, but very low values of the variables associated with NO ₃ ^– production and loss. Correlations between soil N transformations and litter concentrations of N, lignin, lignin:N ratio, or phenolic constituents were generally weak. The inverse correlation between net nitrification rate and soil C:N that has been reported in the literature was present in this data set only if red oak plots were excluded from the analysis. This study indicates that tree species can exert a strong control on N cycling in forest ecosystems that appears to be mediated through the quality of soil organic matter, but that standard measures of litter quality cannot explain the mechanism of control.