Ungulate stimulation of nitrogen cycling and retention in Yellowstone Park grasslands

We studied how ungulates and a large variation in site conditions influenced grassland nitrogen (N) dynamics in Yellowstone National Park. In contrast to most grassland N studies that have examined one or two soil N processes, we investigated four rates, net N mineralization, nitrification, denitrification, and inorganic N leaching, at seven paired sites inside and outside long-term (33+ year) exclosures. Our focus was how N fluxes were related to one another among highly variable grasslands and how grazers influenced those relationships. In addition, we examined variation in soil δ¹⁵N among grasslands and the relationships between soil ¹⁵N abundance and N processes. Previously, ungulates were reported to facilitate net N mineralization across variable Yellowstone grasslands and denitrification at mesic sites. In this study, we found that herbivores also promoted nitrification among diverse grasslands. Furthermore, net N mineralization, nitrification, and denitrification (kg N ha^–1 year^–1, each variable) were postively and linearly related to one another among all grasslands (grazed and fenced), and grazers reduced the nitrification/net N mineralization and denitrification/net N mineralization ratios, indicating that ungulates inhibited the proportion of available NH₄ ⁺ that was nitrified and denitrified. There was no relationship between net N mineralization or nitrification with leaching (indexed by inorganic N adsorbed to resin buried at the bottom of rooting zones) and leaching was unaffected by grazers. Soil δ¹⁵N was positively and linearly related to in situ net N mineralization and nitrification in ungrazed grasslands; however, there was no relationship between isotopic composition of N and those rates among grazed grasslands. The results suggested that grazers simultaneously increased N availability (stimulated net N mineralization and nitrification per unit area) and N conservation (reduced N loss from the soil per unit net N mineralization) in Yellowstone grasslands. Grazers promoted N retention by stimulating microbial productivity, probably caused by herbivores promoting labile soil C. Process-level evidence for N retention by grazers was supported by soil δ¹⁵N data. Grazed grassland with high rates of N cycling had substantially lower soil δ¹⁵N relative to values expected for ungrazed grassland with comparable net N mineralization and nitrification rates. These soil ¹⁵N results suggest that ungulates inhibited N loss at those sites. Such documented evidence for consumer control of N availability to plants, microbial productivity, and N retention in Yellowstone Park is further testimony for the widespread regulation of grassland processes by large herbivores. Received: 5 May 1999 / Accepted: 1 November 1999     

Herbivore influence on soil microbial biomass and nitrogen mineralization in a northern grassland ecosystem: Yellowstone National Park

Microorganisms are largely responsible for soil nutrient cycling and energy flow in terrestrial ecosystems. Although soil microorganisms are affected by topography and grazing, little is known about how these two variables may interact to influence microbial processes. Even less is known about how these variables influence microorganisms in systems that contain large populations of free-roaming ungulates. In this study, we compared microbial biomass size and activity, as measured by in situ net N mineralization, inside and outside 35- to 40-year exclosures across a topographic gradient in northern Yellowstone National Park. The objective was to determine the relative effect of topography and large grazers on microbial biomass and nitrogen mineralization. Microbial C and N varied by almost an order of magnitude across sites. Topographic depressions that contained high plant biomass and fine-textured soils supported the greatest microbial biomass. We found that plant biomass accurately predicted microbial biomass across our sites suggesting that carbon inputs from plants constrained microbial biomass. Chronic grazing neither depleted soil C nor reduced microbial biomass. We hypothesize that microbial populations in grazed grasslands are sustained mainly by inputs of labile C from dung deposition and increased root turnover or root exudation beneath grazed plants. Mineral N fluxes were affected more by grazing than topography. Net N mineralization rates were highest in grazed grassland and increased from dry, unproductive to mesic, highly productive communities. Overall, our results indicate that topography mainly influences microbial biomass size, while mineral N fluxes (microbial activity) are affected more by grazing in this grassland ecosystem. Received: 4 June 1997 / Accepted: 16 December 1997     

Evidence for top predator control of a grazing ecosystem

The importance of top predators in controlling ecological processes in large, intact ecosystems is unclear. In grasslands that support abundant ungulates, top–down control by predators may be particularly important, because of the tight biogeochemical linkages of ungulate prey with plants and soil microbes. Here, I examined the effects of the recent reintroduction of the gray wolf Canis lupus on ecosystem processes in Yellowstone National Park, where herds of grazing ungulates previously have been shown to stimulate several processes, including soil net nitrogen (N) mineralization. Rates of ungulate grazing intensity and soil net N mineralization were compared before and after wolf reintroduction in grasslands ranging five‐fold in aboveground production. Grazing intensity and grassland net N mineralization declined after wolf reintroduction, a likely partial function of fewer ungulates; wolf predation has been one of several factors implicated in causing the decline in Yellowstone ungulates. In addition, the spatial pattern of grazing and net N mineralization changed after reintroduction. A shift in the spatial patterns of grazer‐associated processes is consistent with a growing body of work indicating that wolves have changed habitat use patterns of ungulates in Yellowstone National Park. These findings suggest widespread wolf effects on ungulate prey, plants, and microbial activity that have spatially reorganized grassland energy and nutrient dynamics in Yellowstone Park.     

Grazers and soil moisture determine the fate of added 15NH4 + in Yellowstone grasslands

The influence of ungulate grazers on nutrient cycling and ecosystem productivity in grasslands has been shown to differ with moisture, nutrient availability, and feedbacks between above- and belowground activities. We examined the movement of nitrogen (N), applied as (¹⁵NH₄)₂SO₄, through both dry and mesic sites in the northern range of Yellowstone National Park to test the hypothesis that plants were more able to acquire added N in grazed relative to ungrazed sites. Previous studies showed enhanced N mineralization in grazed areas, and detritus removal by grazers was predicted to enhance early-season plant growth. Thirteen months after tracer addition, there were no differences in plant ¹⁵N as a function of grazing, but historically ungrazed sites retained more ¹⁵N in accumulated litter than at grazed sites. This result demonstrated the importance of detritus in regulating redistribution of incoming N and the role of grazers in this process. Site moisture status influenced ¹⁵N recovery in all pools—soils, microbial biomass, and plants—and greater plant ¹⁵N acquisition occurred in roots at dry relative to mesic sites. Understanding how grazers influence nutrient cycling at the landscape scale requires further investigation of interactions among soil moisture, plant production, litter accumulation, grazing intensity, and belowground processes.     

Interactive Effects of Ungulate Herbivores, Soil Fertility, and Variable Rainfall on Ecosystem Processes in a Semi-arid Savanna

Large herbivores can both positively and negatively affect primary productivity and rates of nutrient cycling in different ecosystems. Positive effects of grazers in grasslands have been attributed to migratory behavior of the dominant ungulate species and soil fertility. We studied the effects of grazers on aboveground net primary productivity (ANPP) and N cycling on central Kenyan rangeland characterized by intense, chronic grazing by a mixed community of cattle and resident native ungulates. Exclosure studies conducted at high and low levels of soil fertility showed that both soil fertility and annual rainfall patterns mediate the effects of grazers on ANPP and N cycling. In a low-rainfall year with short (1 month) growing seasons, grazers reduced aboveground productivity regardless of soil nutrient availability. However, in a high-rainfall year with a 5-month growing season, grazers increased ANPP on nutrient-rich glades and suppressed ANPP on nutrient-poor bushland sites. Concomitant studies of grazer effects on N cycling revealed complex interactions with the seasonal pattern of N-mineralization and inorganic N availability. Grazers increased the size of the inorganic N pool available to plants at the onset of the growing season, particularly in nutrient-rich glades. However, grazers also decreased N mineralization rates at all sites early in the growing season. Measures of N availability via ion-exchange resin bags suggested that the combined effects of grazers on inorganic N pool fluctuations and N-mineralization rates resulted in a net increase in N availability at glade sites and a net decrease in N availability at bushland sites. The net effect of grazers on soil N availability mirrored grazer effects on ANPP in the high-rainfall year. Overall, our results suggest that grazer effects on N dynamics are closely linked to effects on productivity and resilience to drought. Furthermore, even under optimal conditions of high soil fertility and above-average rainfall, grazer promotion of ANPP in this chronically grazed system dominated by resident ungulates was small compared to systems dominated by migratory ungulates.     

Ungulate and topographic control of nitrogen: phosphorus stoichiometry in a temperate grassland; soils, plants and mineralization rates

Although the link between the nitrogen (N): phosphorus (P) stoichiometry of biota and availability has received considerable attention in aquatic systems, there has been relatively little effort to compare the elemental composition of biota and supply in terrestrial habitats. In this study, I explored the effects of a prominent topo-edaphic gradient, from dry hilltop to wet slope-base, and native ungulates on N and P of soils, plants, and rates of in situ net mineralization in grasslands of Yellowstone National Park. Nitrogen and P measurements were made May–September, 2000, in paired, grazed and 38–42 year fenced, ungrazed grassland at five topographically variable sites. Similar to findings from other grassland ecosystems, several site factors associated with organic activity, including soil moisture, C, and plant biomass, covaried with soil N concentration and/or net N mineralization. Soil P concentration and net P mineralization, however, were unrelated to those factors. Instead, net P mineralization was negatively related to soil pH, which is known to control the form of inorganic P and its availability, and soil P was uncorrelated with any soil or plant variable measured in the study. Because of being influenced by different soil properties, N and P net mineralization were unrelated among grasslands. Furthermore, supply and plant N:P ratios were uncorrelated in this grassland system. Based on critical N:P ratios reflecting nutritional limitation of plants, Yellowstone grassland vegetation ranged from being N limited to N-P co-limited. Grazers increased N-P co-limitation by enhancing plant N concentrations and the soil pH gradient across grassland sites regulated plant nutritional limitation by affecting plant-available P. These findings showed how ungulates and a landscape factor, i.e. soil pH, determined plant nutrient status among YNP grasslands differently by influencing plant N concentration versus plant P concentration, respectively.     

Soil nitrogen dynamics in a holm oak forest

Soil nitrogen (N) dynamics were studied in a dense, holm oak (Quercus ilex ssp. ilex) stand in the Montseny mountains to determine annual and seasonal patterns of N availability and uptake in an undisturbed Mediterranean forest on acidic soil. Soil mineral N content, net N mineralization (NNM), and net nitrification (NN) were determined by monthly sampling at two soil depths followed by in situ incubation in polyethylene bags. NNM per unit of soil mass was much higher at 0–5 cm than at 5–20 cm (annual means 24 and 2.5 mg N/kg, respectively) but on an area basis NNM was similar at both depths. A total of 80 kg N/ha/yr were mineralized from the first 20 cm of soil. NN amounted to only 9% of the annual NNM (7.5 kg N/ha/yr) and it occurred only in the upper 5 cm. NNM was maximum in June and July, while the NN peaked in May. Despite favourable soil temperature and moisture, NNM was negative in autumn because of microbial immobilization. Seasonal and depth variations of NNM appeared to be controlled more by substrate quality than by organic matter quantity, temperature or moisture. NN was not limited by ammonium availability. Calculated N uptake amounted to 91 kg/ha yr, peaking in June and July. The investigated stand showed a moderately high N availability, but ammonium was the major form of mineral N supply for holm oak.     

Nutrient composition and inventory in a tropical grassland

Summary The mineral structure and its dynamics in forest ecosystems have been undertaken by a number of workers. Such studies are meagre for tropical grassland. The purpose of this study was to ascertain the nutrient structure for the producer components, and to estimate the annual uptake, retention and release for N, P, K and Ca. Data for these elements are given for plant and soil. The soil contains many more times nutrient than held by plant biomass. Inventory of the minerals indicates that, of the four elements i.e. N, P, K and Ca studied, annually about 58, 16, 18 and 38 kg/ha, respectively are taken up by the grassland, of which 29, 7, 6 and 23 kg/ha are released and about 29, 9, 12 and 14 kg/ha are retained in the vegetation compartments. re]19750516     

Biogeochemical approaches to assessment of East Asian ecosystem sensitivity to acid depositon

We used the critical load (CL) concept to calculate ecosystem response to acid deposition in East Asia. The calculation of critical loads to assess the sensitivity of ecosystems to acidic deposition was made using a biogeochemical approach, which took into consideration both rates of biogeochemical cycling and temperature responses. On the basis of these data the soil-biogeochemical mapping has been carried out for the area of East Asia and the CL values for acid-forming compounds have been calculated using modified steady-state mass balance (SSMB) equations. In the north-eastern ecosystems of the Asian part of Russia these values of critical loads for N [CL(N)] and S [CL(S)] compounds are shown to be less than in Europe due to peculiarities of climate, soil and vegetation. The minimum values of both CL(N) and CL(S) are <50 eq/ha/yr (which occur in 8.3% and 40.5% of this area for N and S, correspondingly) and the maximum values are >300 eq/ha/yr. These values are occasionally lower than for corresponding European ecosystems. For the south-eastern ecosystems of the northern part of Thailand the minimum values are <200 eq/ha/yr and maximum values are >700 eq/ha/yr. The minimum CL values (<200 eq/ha/yr) occur in more than 75% of the studied Thai area.     

草地生态系统中土壤氮素矿化影响因素的研究进展

氮素是各种植物生长和发育所需的大量营养元素之一,也是牧草从土壤吸收最多的矿质元素．土壤中的氮大部分以有机态形式存在,而植物可以直接吸收利用的是无机态氮．这些有机态氮在土壤动物和微生物的作用下。由难以被植物直接吸收利用的有机态转化为可被植物直接吸收利用的无机态的过程就是土壤氮的矿化．氮素矿化受多种因子的影响,这些因子可以归结为生物因子和非生物因子．生物因子包括：土壤动物、土壤微生物和植物种类．土壤动物可以促进土壤有机质的矿化;土壤微生物种类、结构及功能与氮的分解、矿化有密切的关系;不同的植物种类对土壤氮素的矿化作用是不相同的,一般来说。有豆科植物生长的土壤比其它种类土氮素矿化的作用大．非生物因素一般可以分为环境因子和人类活动干扰．环境因子中土壤温度和含水量对土壤氮素矿化的影响是国内外众多科学家研究的方向．尽管如此,在此方面的研究还没有取得一致意见,仍然需要进行这方面的研究,而在其他诸如：不同的土壤质地与土壤类型方面,研究报道的结论也很不一致,草地生态系统中人类活动对土壤氮素矿化的影响主要包括,不同强度的放牧,割草以及施肥、火烧强度等．非生物因子对氮素矿化的影响非常直接和明显,尤其是人类活动．本文综述了近年来影响草地生态系统土壤氮素矿化有关因素的一些进展．     

Soil C and N responses to woody plant expansion in a mesic grassland

Changes in land management and reductions in fire frequency have contributed to increased cover of woody species in grasslands worldwide. These shifts in plant community composition have the potential to alter ecosystem function, particularly through changes in soil processes and properties. In semi-arid grasslands, the invasion of shrubs and trees is often accompanied by increases in soil resources and more rapid N and C cycling. We assessed the effects of shrub encroachment in a mesic grassland in Kansas (USA) on soil CO₂ flux, extractable inorganic N, and N mineralization beneath shrub communities (Cornus drummondii) and surrounding undisturbed grassland sites. In this study, a shift in plant community composition from grassland to shrubland resulted in a 16% decrease in annual soil CO₂ flux(4.78 kg CO₂ m^–2 year^–1 for shrub dominated sites versus 5.84 kg CO₂ m^–2 year^–1 for grassland sites) with no differences in total soil C or N or inorganic N. There was considerable variability in N mineralization rates within sites, which resulted in no overall difference in cumulative N mineralized during this study (4.09 g N m^–2 for grassland sites and 3.03 g N m^–2 for shrub islands). These results indicate that shrub encroachment into mesic grasslands does not significantly alter N availability (at least initially), but does alter C cycling by decreasing soil CO₂ flux.     

Original Articles: Nitrogen Mineralization in Two Unpolluted Old-Growth Forests of Contrasting Biodiversity and Dynamics

Studies in unpolluted, old-growth forests in the coastal range of southern Chile (42°30′S) can provide a baseline for understanding how forest ecosystems are changing due to the acceleration of nitrogen (N) inputs that has taken place over the last century. Chilean temperate forests, in contrast to their northern hemisphere counterparts, exhibit extremely low losses of inorganic N to stream waters. The objectives of this study were (a) to determine whether low inorganic N outputs in these forests were due to low rates of N mineralization or nitrification, and (b) to examine how biodiversity (defined as number of dominant tree species) and forest structure influence N mineralization and overall patterns of N cycling. Studies were conducted in a species-poor, conifer-dominated (Fitzroya cupressoides) forest with an even-aged canopy, and in a mixed-angiosperm (Nothofagus nitida) forest with a floristically more diverse and unstable canopy. Nitrogen mineralization rates measured in laboratory assays varied seasonally, reaching 6.0 μg N/g DW/day in both forests during late summer. Higher values were related to higher microbial activity, larger pools of labile inorganic N, and increased fine litter inputs. Field assays, conducted monthly, indicated positive net flux from N mineralization mainly from December to January in both forests. Annual net flux of N from mineralization varied from 20 to 23 kg/ha/year for the Fitzroya forest and from 31 to 37 kg/ha/year for the Nothofagus forest. Despite low losses of inorganic N to streams, N mineralization and nitrification are not inhibited in these forests, implying the existence of strong sinks for NO₃ ⁻ in the ecosystem. Field N mineralization rates were two times higher in the Nothofagus forest than in the Fitzroya forest, and correlated with greater N input via litterfall, slightly higher soil pH, and narrower carbon (C)–nitrogen ratios of soils and litter in the former. Differences in N mineralization between the two forest types are attributed to differences in biotic structure, stand dynamics, and site factors. Median values of net N mineralization rates in these southern hemisphere forests were lower than median rates for forests in industrialized regions of North America, such as the eastern and central USA. We suggest that these high N mineralization rates may be a consequence of enhanced atmospheric N deposition.     

Ammonia volatilization from a seasonally and spatially variable grazed grassland: Yellowstone National Park

We measured ammonia volatilization at three topographic positions(hilltop, mid-slope, slope-bottom) on three grassland landscapes at threetimes during 1995 (April, May, July) on the northern winter range ofYellowstone National Park that supports large herds of native ungulates.Percent ammonia-N lost from all sites during the study ranged 1–24%of urea-N applied. Volatilization among sites was negatively related tosoil cation-exchange capacity (r = –0.85) and rates were highest inJuly. We used the relationship between soil CEC and percent Nvolatilized from urea-amended plots to estimate annual ammonia-Nvolatilization from 5 sites for which annual ungulate urine inputs werepreviously determined (Frank et al. 1994). Estimated mean annualammonia-N volatilized from those sites was 1.4 kg/ha/yr, which wasless than a previously reported regional atmospheric deposition rate (2kg/ha/yr; Swank 1984). Results indicate the need to understand theinteraction between (1) spatially heterogeneous patterns of soilprocesses, and 2) nonuniform patterns of ungulate use of landscapes todetermine rates of ecosystem-level N-gaseous loss. Findings alsosuggest that ammonia-N volatilized from urine patches should not leadto a decline in soil N in this ecosystem.     

Nutrient losses in runoff from grassland and shrubland habitats in Southern New Mexico: I. rainfall simulation experiments

Rainfall simulation experiments were performed in areas of semiarid grassland (Bouteloua eriopoda) and arid shrubland (Larrea tridentata) in the Chihuahuan desert of New Mexico. The objective was to compare the runoff of nitrogen (N) and phosphorus (P) from these habitats to assess whether losses of soil nutrients are associated with the invasion of grasslands by shrubs. Runoff losses from grass- and shrub-dominated plots were similar, and much less than from bare plots located in the shrubland. Weighted average concentrations of total dissolved N compounds in runoff were greatest in the grassland (1.72 mg/1) and lowest in bare plots in the shrubland (0.55 mg/1). More than half of the N transported in runoff was carried in dissolved organic compounds. In grassland and shrub plots, the total N loss was highly correlated to the total volume of discharge. We estimate that the total annual loss of N in runoff is 0.25 kg/ha/yr in grasslands and 0.43 kg/ha/yr in shrublands — consistent with the depletion of soil N during desertification of these habitats. Losses of P from both habitats were very small.     

Faunal isotope records reveal trophic and nutrient dynamics in twentieth century Yellowstone grasslands

Population sizes and movement patterns of ungulate grazers and their predators have fluctuated dramatically over the past few centuries, largely owing to overharvesting, land-use change and historic management. We used δ(13)C and δ(15)N values measured from bone collagen of historic and recent gray wolves and their potential primary prey from Yellowstone National Park to gain insight into the trophic dynamics and nutrient conditions of historic and modern grasslands. The diet of reintroduced wolves closely parallels that of the historic population. We suggest that a significant shift in faunal δ(15)N values over the past century reflects impacts of anthropogenic environmental changes on grassland ecosystems, including grazer-mediated shifts in grassland nitrogen cycle processes.     

干扰对典型草原生态系统土壤净呼吸特征的影响

由于土地利用格局的改变和人类干扰活动的加剧,草地生态系统CO2排放与固定的平衡、碳循环特征以及碳储量越来越受到人们的重视。尤其是定量区分土壤净呼吸与土壤总呼吸量之间的比例关系,以及定量描述草地生态系统碳循环过程等方面的研究尚不够完善。以河北沽源的典型草原为研究对象,测定了火烧、灌溉、施肥、刈割干扰下的天然草地土壤净呼吸变化动态及其与主要控制因素之间的关系。结果表明:不同处理土壤净呼吸均表现出明显的季节性变化规律,变化趋势基本一致。火烧、灌溉和刈割处理分别比对照的土壤净呼吸通量降低了28.93%、16.25%和36.82%。土壤温度、土壤湿度与土壤净呼吸通量呈指数相关(P0.01)。对地上生物量、地下生物量、土壤有机碳含量和土壤全氮含量与土壤净呼吸之间进行逐步回归分析表明,土壤有机碳含量(SC)和土壤全氮含量(SN)是土壤净呼吸通量的主要影响因素。     

Nitrogen response efficiency increased monotonically with decreasing soil resource availability: a case study from a semiarid grassland in northern China

The concept of nutrient use efficiency is central to understanding ecosystem functioning because it is the step in which plants can influence the return of nutrients to the soil pool and the quality of the litter. Theory suggests that nutrient efficiency increases unimodally with declining soil resources, but this has not been tested empirically for N and water in grassland ecosystems, where plant growth in these ecosystems is generally thought to be limited by soil N and moisture. In this paper, we tested the N uptake and the N use efficiency (NUE) of two Stipa species (S. grandis and S. krylovii) from 20 sites in the Inner Mongolia grassland by measuring the N content of net primary productivity (NPP). NUE is defined as the total net primary production per unit N absorbed. We further distinguished NUE from N response efficiency (NRE; production per unit N available). We found that NPP increased with soil N and water availability. Efficiency of whole-plant N use, uptake, and response increased monotonically with decreasing soil N and water, being higher on infertile (dry) habitats than on fertile (wet) habitats. We further considered NUE as the product of the N productivity (NP the rate of biomass increase per unit N in the plant) and the mean residence time (MRT; the ratio between the average N pool and the annual N uptake or loss). The NP and NUE of S. grandis growing usually in dry and N-poor habitats exceeded those of S. krylovii abundant in wet and N-rich habitats. NUE differed among sites, and was often affected by the evolutionary trade-off between NP and MRT, where plants and communities had adapted in a way to maximize either NP or MRT, but not both concurrently. Soil N availability and moisture influenced the community-level N uptake efficiency and ultimately the NRE, though the response to N was dependent on the plant community examined. These results show that soil N and water had exerted a great impact on the N efficiency in Stipa species. The intraspecific differences in N efficiency within both Stipa species along soil resource availability gradient may explain the differences in plant productivity on various soils, which will be conducive to our general understanding of the N cycling and vegetation dynamics in northern Chinese grasslands.     

Asymmetric effects of grazing intensity on macroelements and microelements in grassland soil and plants in Inner Mongolia Grazing alters nutrient dynamics of grasslands

Grazing is a traditional grassland management technique and greatly alters ecosystem nutrient cycling. The effects of grazing intensity on the nutrient dynamics of soil and plants in grassland ecosystems remain uncertain, especially among microelements. A 2‐year field grazing experiment was conducted in a typical grassland with four grazing intensities (ungrazed control, light, moderate, and heavy grazing) in Inner Mongolia, China. Nutrient concentration was assessed in soil and three dominant plant species (Stipa krylovii, Leymus chinensis, and Cleistogenes squarrosa). Assessed quantities included four macroelements (carbon (C), nitrogen (N), phosphorus (P), and magnesium (Mg)) and four microelements (copper (Cu), iron (Fe), manganese (Mn), and zinc (Zn)). Soil total C, total N, total P, available N, and available P concentrations significantly increased with grazing intensity but soil Mg, Cu, Fe, Mn, Zn concentrations had no significant response. Plant C concentration decreased but plant N, P, Mg, Cu, Fe, Mn, and Zn concentrations significantly increased with grazing intensity. In soil, macroelement dynamics (i.e., C, N, and P) exhibited higher sensitivity with grazing intensity, conversely in plants, microelements were more sensitive. This result indicates macroelements and microelements in soil and plants had asymmetric responses with grazing intensity. The slopes of nutrient linear regression in C. squarrosa were higher than that of S. krylovii and L. chinensis, indicating that C. squarrosa had higher nutrient acquisition capacity and responded more rapidly to heavy grazing. These findings indicate that short‐term heavy grazing accelerates nutrient cycling of the soil–plant system in grassland ecosystems, elucidate the multiple nutrient dynamics of soil and plants with grazing intensity, and emphasize the important function of microelements in plant adaptation in grazing management.     

Effects of an African grass invasion on Hawaiian shrubland nitrogen biogeochemistry

African perennial C₄ grasses are highly successful invaders in Hawaiian ecosystems. We examined the effects of African molasses grass (Melinis minutiflora Beauv.) on Hawaiian shrubland nitrogen (N) dynamics without the influence of fire disturbance. Vegetation tissue carbon and nitrogen chemistry, soil inorganic N pools, net N mineralization rates, and total soil N were studied in three adjacent areas: a monospecificMelinis grassland, a mixed grass/shrubland mosaic, and an un-invaded shrubland.Melinis plots within the mosaic area exhibited the largest inorganic N pools and fastest net N mineralization rates, but were temporally variable with grass phenology. Un-invaded shrubland plots contained the smallest inorganic N pools and lowest net N mineralization rates. Grass foliar C:N and litter C:N were lower than those of common shrubland species, providing one possible link between species and ecosystem N dynamics at this site. The combined effects of N cycle modification, successful light competition, and fire-cycle enhancement make the invasion ofMelinis a significant perturbation to Hawaiian shrubland ecosystem function and successional dynamics. ei]Section editor H Lambers     

The effects of warming and nitrogen addition on soil nitrogen cycling in a temperate grassland, northeastern China

Background

Both climate warming and atmospheric nitrogen (N) deposition are predicted to affect soil N cycling in terrestrial biomes over the next century. However, the interactive effects of warming and N deposition on soil N mineralization in temperate grasslands are poorly understood.

Methodology/Principal Findings

A field manipulation experiment was conducted to examine the effects of warming and N addition on soil N cycling in a temperate grassland of northeastern China from 2007 to 2009. Soil samples were incubated at a constant temperature and moisture, from samples collected in the field. The results showed that both warming and N addition significantly stimulated soil net N mineralization rate and net nitrification rate. Combined warming and N addition caused an interactive effect on N mineralization, which could be explained by the relative shift of soil microbial community structure because of fungal biomass increase and strong plant uptake of added N due to warming. Irrespective of strong intra- and inter-annual variations in soil N mineralization, the responses of N mineralization to warming and N addition did not change during the three growing seasons, suggesting independence of warming and N responses of N mineralization from precipitation variations in the temperate grassland.

Conclusions/Significance

Interactions between climate warming and N deposition on soil N cycling were significant. These findings will improve our understanding on the response of soil N cycling to the simultaneous climate change drivers in temperate grassland ecosystem.