Denitrification in a semi-arid grazing ecosystem

The effect of large herbivores on gaseous N loss from grasslands, particularly via denitrification, is poorly understood. In this study, we examined the influence of native migratory ungulates on denitrification in grasslands of Yellowstone National Park in two ways, by (1) examining the effect of artificial urine application on denitrification, and (2) comparing rates inside and outside long-term exclosures at topographically diverse locations. Artificial urine did not influence denitrification 3 and 12 days after application at hilltop, mid-slope, and slope-bottom sites. Likewise, grazers had no effect on community-level denitrification at dry exclosure sites, where rates were low. At mesic sites, however, ungulates enhanced denitrification by as much as 4 kg N ha⁻¹ year⁻¹, which was double atmospheric N inputs to this ecosystem. Denitrification enzyme activity (DEA, a measure of denitrification potential) was positively associated with soil moisture at exclosure sites, and herbivores stimulated DEA when accounting for the soil moisture effect. Glucose additons to soils increased denitrification and nitrate additions had no influence, suggesting that denitrification was limited by the amount of labile soil carbon, which previously has been shown to be enhanced by ungulates in Yellowstone. These results indicate that denitrification can be an ecologically important flux in portions of semi-arid landscapes, and that there is a previously unsuspected regulation of this process by herbivores. Received: 6 March 1998 / Accepted: 28 August 1998     

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.     

The biogeochemistry of a north-temperate grassland with native ungulates: Nitrogen dynamics in Yellowstone National Park

Nutrient dynamics of large grassland ecosystems possessing abundant migratory grazers are poorly understood. We examined N cycling on the northern winter range of Yellowstone National Park, home for large herds of free-roaming elk (Cervus elaphus) and bison (Bison bison). Plant and soil N, net N mineralization, and the deposition of ungulate fecal-N were measured at five sites, a ridgetop, mid-slope bench, steep slope, valley-bottom bench, and riparian area, within a watershed from May, 1991 to April, 1992.Results indicated similarities between biogeochemical properties of Yellowstone grassland and other grassland ecosystems: (1) landscape position and soil water affected nutrient dynamics, (2) annual mineralization was positively related to soil N content, and (3) the proportion of soil N mineralized during the year was negatively related to soil C/N.Grazers were a particularly important component of the N budget of this grassland. Estimated rates of N flow from ungulates to the soil ranged from 8.1 to 45.6 kg/ha/yr at the sites (average = 27.0 kg/ha/yr), approximately 4.5 times the amount of N in senescent plants. Rates of nitrogen mineralization for Yellowstone northern range grassland were higher than those measured in other temperate grassland ecosystems, possibly due to grazers promoting N cycling in Yellowstone.     

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.     

Nitrogen dynamics in alpine ecosystems of the northern Caucasus

Net N mineralization, nitrification, microbial biomass N and ¹⁵N natural abundance were studied in a toposequence of representative soils and plant communities in the alpine zone of the northern Caucasus. The toposequence was represented by (1) low-productive alpine lichen heath (ALH) of wind-exposed ridge and upper slope; (2) more productive Festuca varia grassland (FG) of middle slope; (3) most productive Geranium gymnocaulon/Hedusarum caucasicummeadow (GHM) of lower slope; (4) low-productive snowbed community (SBC) of the slope bottom. N availability, net N mineralization and nitrification were higher in soils of alpine grassland and meadow of the middle part of the toposequence compared with soils of lichen heath and snowbed community of extreme habitats in the alpine zone. There was no correlation between intensities of N transformation processes and favorable (low soil acidity, low C/N ratio, long vegetation period, relatively high temperature, absence of hydromorphic features) and unfavorable (opposite) factors, indicating that the intensity of N mineralization and nitrification in the alpine soils is controlled by a complex combination of these factors. Potential net N mineralization and nitrification in alpine soils determined in the short-term laboratory incubation were considerably higher than those determined in the long-term field incubation. The differences of potential nitrification between soils of various plant communities did not correspond to the field determined pattern indicating the importance of on-site climatic conditions for control of nitrification in high mountains. The result of comparison of N transformation potentials in incubated and native soils indicated that nitrification potential was significantly increased after long-term soil incubation. It means that net nitrification determined in the field was probably overestimated, especially in the meadow soils. A soil translocation experiment indicated that low temperature was an important factor limiting net N mineralization and nitrification in alpine soils: net N mineralization and especially nitrification increased when alpine soils were translocated into the subalpine zone and mean annual temperature increased by about 3°C. Additional N input increased N availability (NH₄ ⁺-N) and potential nitrification in soils of the lower part of the toposequense (GHM and SBC), and potential net N mineralization in two soils of extreme habitats (ALH and SBC). A positive correlation was found between soil ¹⁵N and net N mineralization and nitrification; the relative ¹⁵N enrichment was characteristic of grassland and meadow ecosystems. ¹⁵N of total soil N pool increased during the field mineralization experiment; there was a positive tendency between the change in ¹⁵N and net N mineralization and nitrification, however the relationship was not significant. Foliar ¹⁵N of dominant plant species varied widely within community, however, a tendency of higher foliar ¹⁵N for species growing on the soils with higher net N mineralization, nitrification and ¹⁵N was observed.     

The interactive effects of grazing ungulates and aboveground production on grassland diversity

The variable and nonlinear relationships between plant species richness (SR) and aboveground production (NAP) among terrestrial ecosystems indicate that the energetic capacity of ecosystems interacts with other environmental factors to control diversity. One contributing factor determining plant diversity is herbivory; but few studies have effectively examined the interaction of herbivores and NAP on SR. The objective of this study was to investigate how NAP and herds of native migrating ungulates determine plant SR in grasslands of Yellowstone National Park. Plant SR at peak aboveground biomass was compared inside and outside ungulate exclosures at two spatial scales, 1.0 m² (“local”) and 100 m² (“community”), in ten variable grasslands. NAP also was determined inside and outside exclosures. The relationship between SR and NAP was unimodal for grazed and ungrazed grassland at both spatial scales. Grazers increased local SR, independent of NAP. In contrast, herbivore effects on community SR ranged from no effect among low-productive grassland to an increasingly positive influence as NAP increased. In addition, ungulates reduced beta diversity (the contribution to community SR attributed to variability among local patches) at dry, low-productive and wet, high-productive sites. These results suggest that the size of the pool of species available to colonize grassland is an important factor controlling the response of grassland SR to herbivory, particularly from low- to intermediate-productive grassland.     

Plant and soil natural abundance <Emphasis Type="Italic">δ</Emphasis><Superscript>15</Superscript>N: indicators of relative rates of nitrogen cycling in temperate forest ecosystems

Watersheds within the Catskill Mountains, New York, receive among the highest rates of nitrogen (N) deposition in the northeastern United States and are beginning to show signs of N saturation. Despite similar amounts of N deposition across watersheds within the Catskill Mountains, rates of soil N cycling and N retention vary significantly among stands of different tree species. We examined the potential use of δ ¹⁵N of plants and soils as an indicator of relative forest soil N cycling rates. We analyzed the δ ¹⁵N of foliage, litterfall, bole wood, surface litter layer, fine roots and organic soil from single-species stands of American beech (Fagus grandifolia), eastern hemlock (Tsuga canadensis), red oak (Quercus rubra), and sugar maple (Acer saccharum). Fine root and organic soil δ ¹⁵N values were highest within sugar maple stands, which correlated significantly with higher rates of net mineralization and nitrification. Results from this study suggest that fine root and organic soil δ ¹⁵N can be used as an indicator of relative rates of soil N cycling. Although not statistically significant, δ ¹⁵N was highest within foliage, wood and litterfall of beech stands, a tree species associated with intermediate levels of soil N cycling rates and forest N retention. Our results show that belowground δ ¹⁵N values are a better indicator of relative rates of soil N cycling than are aboveground δ ¹⁵N values.     

Variations in soil N cycling and trace gas emissions in wet tropical forests

We used a previously described precipitation gradient in a tropical montane ecosystem of Hawai’i to evaluate how changes in mean annual precipitation (MAP) affect the processes resulting in the loss of N via trace gases. We evaluated three Hawaiian forests ranging from 2200 to 4050 mm year⁻¹ MAP with constant temperature, parent material, ecosystem age, and vegetation. In situ fluxes of N₂O and NO, soil inorganic nitrogen pools (NH₄⁺ and NO₃⁻), net nitrification, and net mineralization were quantified four times over 2 years. In addition, we performed ¹⁵N-labeling experiments to partition sources of N₂O between nitrification and denitrification, along with assays of nitrification potential and denitrification enzyme activity (DEA). Mean NO and N₂O emissions were highest at the mesic end of the gradient (8.7±4.6 and 1.1±0.3 ng N cm⁻² h⁻¹, respectively) and total oxidized N emitted decreased with increased MAP. At the wettest site, mean trace gas fluxes were at or below detection limit (≤0.2 ng N cm⁻² h⁻¹). Isotopic labeling showed that with increasing MAP, the source of N₂O changed from predominately nitrification to predominately denitrification. There was an increase in extractible NH₄⁺ and decline in NO₃⁻, while mean net mineralization and nitrification did not change from the mesic to intermediate sites but decreased dramatically at the wettest site. Nitrification potential and DEA were highest at the mesic site and lowest at the wet site. MAP exerts strong control N cycling processes and the magnitude and source of N trace gas flux from soil through soil redox conditions and the supply of electron donors and acceptors.     

Nitrate sources and watershed denitrification inferred from nitrate dual isotopes in the Beijiang River,south China

The great spatial and temporal variability of nitrogen (N) processing introduces large uncertainties for quantifying N cycles in large scales, e.g. a watershed scale, and hence challenges the present techniques in measuring ecosystem N mass balance. The dual isotopes of nitrate (δ¹⁸O and δ¹⁵N) integrate signals for both nitrate sources and N processing, making them promising for studies on large scale N cycling. Here, the dual isotopes, as well as some ion tracers, from a subtropical river in south China were reported to identify the main nitrate sources and to assess the possible occurrence and degree of denitrification in the context of monsoon climate. Our results indicated that nitrification of reduced fertilizer N in soil zones was the main nitrate source, with sewage and manure as another important source in dry winter. Seasonal changes of denitrification was apparent by the ~1:2 enrichment of ¹⁸O and ¹⁵N from April to August, and suggested to occur over the watershed rather than in the river. The lowest denitrification (10%) occurred in April, when the fertilizer application was strongest and the monsoon rainfall abruptly increased, causing enhancement of leaching. The highest denitrification (48%) took place in August due to the high soil temperature and moisture. In December, denitrification was significant (26%) perhaps due to the high enough temperature for microbial activities, whereas the low soil moisture appeared to limit the degree of denitrification. This study suggests that the seasonal variations in denitrification should be taken into account when estimating regional N mass balance.     

The pattern between nitrogen mineralization and grazing intensities in an Inner Mongolian typical steppe

Ungulate grazing is known to play a crucial role in regulating energy flow and nutrient cycling in grassland ecosystems. However, previous studies of the effect of grazing on soil N dynamics have showed controversial results. Some studies indicate that grazing stimulates N mineralization while others report that grazing suppresses N mineralization. In order to reconcile these contrasting results, we investigated the response pattern of nitrogen transformation to multiple grazing intensities in an Inner Mongolian steppe. In our study, we measured net nitrogen mineralization rates and nitrification rates during a whole growing season in a 17-year field experiment that had five grazing intensities (0.00, 1.33, 2.67, 4.00 and 5.33 sheep ha⁻¹). Primarily because of changes in temperature and moisture conditions, net N mineralization rates varied substantially during the growing season with higher values occurring in late July. No consistent differences in net N mineralization rates were observed between grazing intensity treatments at the monthly time scale. Compared to mineralization rates, net nitrification rates were generally low with slightly higher values occurring in late July and late August. Ungulate grazing stimulated the cumulative net N transformations (mineralization, nitrification and ammonification) at the annual time scale, and the most stimulation occurred at a moderate grazing intensity of 4.00 sheep ha⁻¹, whereas the highest grazing intensity of 5.33 sheep ha⁻¹ and the lighter grazing intensity of 1.33 sheep ha⁻¹ stimulated less. The general response of net N mineralization to grazing intensity gradient is roughly in the form of a normal distribution at the annual time scale. Our study demonstrated that grazing intensity in concert with soil moisture and temperature conditions imposed significant controls on soil N transformation and availability in this Inner Mongolian steppe.     

Denitrification and N mineralization from hairy vetch (Vicia villosa Roth) and rye (Secale cereale L.) cover crop monocultures and bicultures

N mineralization, N immobilization and denitrification were determined for vetch, rye and rye-vetch cover crops using large packed soil cores. Plants were grown to maturity from seed in cores. Cores were periodically leached, allowing for quantification of NO₃ ⁻ and NH₄ ⁺ production, and denitrification incubations were conducted before and after cover crop kill. Gas permeable tubing was buried at two depths in cores allowing for quantification of N₂O in the soil profile. Cover crops assimilated most soil N prior to kill. After kill, relative rates of N mineralization were vetch > rye-vetch mixture > fallow > rye. After correcting for N mineralization from fallow cores, net N mineralization was observed in vetch and rye-vetch cores, while net N immobilization was observed in rye cores. Denitrification incubations were conducted 5, 15 and 55 days after kill, with adjustment of cores to 75% water filled pore space (WFPS). The highest denitrification was observed in vetch cores 5 days after kill, when soil NO₃ ⁻ and respiration rates were high. Substantially lower denitrification was observed on subsequent measurement dates and in other treatments probably due to either limited NO₃ ⁻ or organic carbon in the soil. On day 5, 3%, 23%, 31% and 31% of the N₂O was recovered in the headspace of fallow, vetch, rye and rye-vetch cores, respectively. The rest was stored in the soil profile. In a field study using intact soil cores, denitrification rates also peaked 1 week after cover crop kill and decreased significantly thereafter. Results suggest greater potential N losses from vetch than rye or rye-vetch cover crops due to rapid N-mineralization in conjunction with denitrification and potential leaching, prior to significant crop N-assimilation. This revised version was published online in June 2006 with corrections to the Cover Date.     

Correlations between foliar δ15N and nitrogen concentrations may indicate plant-mycorrhizal interactions

Nitrogen isotope measurements may provide insights into changing interactions among plants, mycorrhizal fungi, and soil processes across environmental gradients. Here, we report changes in δ¹⁵N signatures due to shifts in species composition and nitrogen (N) dynamics. These changes were assessed by measuring fine root biomass, net N mineralization, and N concentrations and δ¹⁵N of foliage, fine roots, soil, and mineral N across six sites representing different post-deglaciation ages at Glacier Bay, Alaska. Foliar δ¹⁵N varied widely, between 0 and –2‰ for nitrogen-fixing species, between 0 and –7‰ for deciduous non-fixing species, and between 0 and –11‰ for coniferous species. Relatively constant δ¹⁵N values for ammonium and generally low levels of soil nitrate suggested that differences in ammonium or nitrate use were not important influences on plant δ¹⁵N differences among species at individual sites. In fact, the largest variation among plant δ¹⁵N values were observed at the youngest and oldest sites, where soil nitrate concentrations were low. Low mineral N concentrations and low N mineralization at these sites indicated low N availability. The most plausible mechanism to explain low δ¹⁵N values in plant foliage was a large isotopic fractionation during transfer of nitrogen from mycorrhizal fungi to plants. Except for N-fixing plants, the foliar δ¹⁵N signatures of individual species were generally lower at sites of low N availability, suggesting either an increased fraction of N obtained from mycorrhizal uptake (f), or a reduced proportion of mycorrhizal N transferred to vegetation (T _r). Foliar and fine root nitrogen concentrations were also lower at these sites. Foliar N concentrations were significantly correlated with δ¹⁵N in foliage of Populus, Salix, Picea, and Tsuga heterophylla, and also in fine roots. The correlation between δ¹⁵N and N concentration may reflect strong underlying relationships among N availability, the relative allocation of carbon to mycorrhizal fungi, and shifts in either f or T _r. Received: 14 December 1998 / Accepted: 16 August 1999     

Net mineralization and nitrification rates in a clay soil measured and predicted in permanent grassland from soil temperature and moisture content

Summary Net mineralization of N and net nitrification in field-moist clay soils (Evesham-Kingston series) from arable and grassland sites were measured in laboratory incubation experiments at 4, 10 and 20°C. Three depth fractions to 30 cm were used. Nitrate accumulated at all temperatures except when the soil was very dry (=0.13 cm³ cm^–3). Exchangeable NH₄-ions declined during the first 24 h and thereafter remained low. Net mineralization and net nitrification approximated to zero-order reactions after 24 h, with Q₁₀ values generally <1.6. The effect of temperature on both processes was linear although some results conformed to an Arrhenius-type relationship. The dependence of net mineralization and net nitrification in the field soil on soil temperature (10 cm depth) and moisture (0–15, 15–25, 25–35 cm depths) was modelled using the laboratory incubation data. An annual net mineralization of 350 kg N ha^–1 and net nitrification of 346 kg N ha^–1 were predicted between September 1980 and August 1981. The model probably overstressed the effect of soil moisture relative to soil temperature.     

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.     

The effect of single tree species on soil microbial activities related to C and N cycling in the Siberian artificial afforestation experiment

The effects of grassland conversion to forest vegetation and of individual tree species on microbial activity in Siberia are largely unstudied. Here, we examined the effects of the six most commonly dominant tree species in Siberian forests (Scots pine, spruce, Arolla pine, larch, aspen and birch) on soil C and N mineralization, N₂O-reduction and N₂O production during denitrification 30 years after planting. We also documented the effect of grassland conversion to different tree species on microbial activities at different soil depths and their relationships to soil chemical properties. The effects of tree species and grassland conversion were more pronounced on N than on C transformations. Tree species and grassland conversion did significantly alter substrate-induced respiration (SIR) and basal respiration, but the differences were not as large as those observed for N transformations. Variances in SIR and basal respiration within species were markedly lower than those in N transformations. Net N mineralization, net nitrification, and denitrification potential were highest under Arolla pine and larch, intermediate under deciduous aspen and birch, and lowest beneath spruce and Scots pine. Tree species caused similar effects on denitrification potential, net N mineralization, and net nitrification, but effects on N₂O reduction rate were idiosyncratic, indicating a decoupling of N₂O production and reduction. We predict that deciduous species should produce more N₂O in the field than conifers, and that Siberian forests will produce more N₂O if global climate change alters tree species composition. Basal respiration and SIR showed inverse responses to tree species: when basal respiration increased in response to a given tree species, SIR declined. SIR may have been controlled by NH₄ ⁺ availability and related therefore to N mineralization, which was negatively affected by grassland conversion. Basal respiration appeared to be less limited by NH₄ ⁺ and controlled mostly by readily available organic C (DOC), which was higher in concentration under forests than in grassland and therefore basal respiration was higher in forested soils. We conclude that in the Siberian artificial afforestation experiment, soil C mineralization was not limited by N.     

Net nitrogen mineralization and net nitrification along a tropical forest-to-pasture chronosequence

Soil inorganic nitrogen pools, net mineralization and net nitrification rates were compared during the dry season along a chronosequence of upland (terra firme) forest, 3-, 9- and 20-year-old pastures in the western Brazilian Amazon Basin state of Rondônia to investigate the influence of forest conversion to pasture on soil nitrogen cycles. Surface soil (0 to 10 cm) from forest had larger extractable inorganic nitrogen pools than pasture soils. In the forest, NO ₃ ^– pools equaled or exceeded NH ₄ ⁺ pools, while pasture inorganic N pools consisted almost exclusively of NH ₄ ⁺ . Rates of net N mineralization and net nitrification in seven -day laboratory incubations were higher in the seven - day forest than in the pastures. Net N mineralization rates did not differ significantly among different-aged pastures, but net nitrification rates were significantly lower in the 20-year-old pasture. Higher net N mineralization and net nitrification rates were measured in laboratory and in situ incubations of sieved soil, compared with in situ incubations of intact soil cores. Rates calculated in seven-day incubations were higher than determined by longer incubations. Sieving may increase N mineralization and/or decrease N immobilization compared with intact cores. We concluded that 7-day laboratory incubation of sieved soil was the most useful index for comparing N availability across the chronosequence of forest and pasture sites. High net nitrification rates in forest soils suggest a potential for NO ₃ ^– losses either through leaching or gaseous emissions.     

Imprint of oaks on nitrogen availability and δ<Superscript>15</Superscript>N in California grassland-savanna: a case of enhanced N inputs?

Woody vegetation is distributed patchily in many arid and semi-arid ecosystems, where it is often associated with elevated nitrogen (N) pools and availability in islands of fertility. We measured N availability and δ¹⁵N in paired blue-oak versus annual grass dominated patches to characterize the causes and consequences of spatial variation in N dynamics of grassland-savanna in Sequoia-Kings Canyon National Park. We found significantly greater surface soil N pools (0–20 cm) in oak patches compared to adjacent grass areas across a 700 m elevation gradient from foothills to the savanna-forest boundary. N accumulation under oaks was associated with a 0.6‰ depletion in soil δ¹⁵N relative to grass patches. Results from a simple δ¹⁵N mass balance simulation model, constrained by surface soil N and δ¹⁵N measured in the field, suggest that the development of islands of N fertility under oaks can be traced primarily to enhanced N inputs. Net N mineralization and percent nitrification in laboratory incubations were consistently higher under oaks across a range of experimental soil moisture regimes, suggesting a scenario whereby greater N inputs to oak patches result in net N accumulation and enhanced N cycling, with a potential for greater nitrate loss as well. N concentrations of three common herbaceous annual plants were nearly 50% greater under oak than in adjacent grass patches, with community composition shifted towards more N-demanding species under oaks. We find that oaks imprint distinct N-rich islands of fertility that foster local feedback between soil N cycling, plant N uptake, and herbaceous community composition. Such patch-scale differences in N inputs and plant–soil interactions increase biogeochemical heterogeneity in grassland-savanna ecosystems and may shape watershed-level responses to chronic N deposition.     

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.     

Comparison of soil nitrogen mineralization and nitrification in a mixed grassland and forested ecosystem in central Taiwan

We used the buried bag incubation method to study temporal patterns of net N mineralization and net nitrification in soils at Ta-Ta-Chia forest in central Taiwan. The site included a grassland zone, (dominant vegetation consists of Yushania niitakayamensis and Miscanthus transmorrisonensis Hayata) and a forest zone (Tsuga chinensis var. formosana and Yushania niitakamensis). In the grassland, soil concentration NH₄ ⁺ in the organic horizon (0.1–0.2 m) ranged from 1.0 to 12.4 mg N kg^–1 soil and that of NO₃ ^– varied from 0.2 to 2.1 mg N kg^–1 soil. In the forest zone, NH₄ ⁺ concentration was between 2.8 and 25.0 mg N kg^–1 soil and NO₃ ^–varied from 0.2 to 1.3 mg N kg^–1 soil. There were lower soil NH₄ ⁺ concentrations during the summer than other seasons. Net N mineralization was higher during the summer while net nitrification rates did not show a distinct seasonal pattern. In the grassland, net N mineralization and net nitrification rates were between –0.1 and 0.24 and from –0.04 to 0.04 mg N kg^–1 soil day^–1, respectively. In the forest zone, net N mineralization rates were between –0.03 and 0.45 mg N kg^–1 soil day^–1 and net nitrification rates were between –0.01 and 0.03 mg N kg^–1 soil day^–1. These differences likely result from differing vegetation communities (C₃ versus C₄ plant type) and soil characteristics.     

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.