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

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

Neutron radiography as a tool for revealing root development in soil: capabilities and limitations

Chronic atmospheric nitrogen deposition affects the cycling of carbon (C) and nitrogen (N) in forest ecosystems, and thereby alters the stable C isotopic abundance of plant and soil. Three successional stages, disturbed, rehabilitated and mature forests were studied for their responses to different nitrogen input levels. N-addition manipulative experiments were conducted at low, medium and high N levels. To study the responses of C cycling to N addition, the C concentration and ¹³C natural abundances for leaf, litter and soil were measured. Labile organic carbon fractions in mineral soils were measured to quantify the dynamics of soil organic C (SOC). Results showed that three-year continuous N addition did not significantly increase foliar C and N concentration, but decreased C/N ratio and enriched ¹³C in N-rich forests. In addition, N addition significantly decreased microbial biomass C, and increased water soluble organic C in surface soils of N-rich forests. This study suggests that N addition enhances the water consumption per unit C assimilation of dominant plant species, restricts SOC turnover in N-poor forests at early and medium successional stages (thus favored SOC sequestration), and vice versa for N-rich mature forests.     

Understanding the Influences of Spatial Patterns on N Availability Within the Brazilian Amazon Forest

Nitrogen variations at different spatial scales and integrated across functional groups were addressed for lowland tropical forests in the Brazilian Amazon as follows: (1) how does N availability vary across the region over different spatial scales (regional × landscape scale); (2) how are these variations in N availability integrated across plant functional groups (legume × non-legume trees). Leaf N, P, and Ca concentrations as well the leaf N isotope ratios (δ¹⁵N) from a large set of legume and non-legume tree species were measured. Legumes had higher foliar N/Ca ratios than non-legumes, consistent with the high energetic costs in plant growth associated with higher foliar P/Ca ratios found in legumes than in non-legumes. At the regional level, foliar δ¹⁵N decreased with increasing rainfall. At the landscape level, N availability was higher in the forests on clayey soils on the plateau than in forests on sandier soils. The isotope as well as the non-isotope data relationships here documented, explain to a large extent the variation in δ¹⁵N signatures across gradients of rainfall and soil. Although at the regional level, the precipitation regime is a major determinant of differences in N availability, at the landscape level, under the same precipitation regime, soil type seems to be a major factor influencing the availability of N in the Brazilian Amazon forest.     

Tree ring δ<Superscript>15</Superscript>N as validation of space-for-time substitution in disturbance studies of forest nitrogen status

Forest ecosystem nitrogen (N) response to disturbance has often been examined by space-for-time substitution, but there are few objective tests of the possible variation in disturbance type and intensity across chronosequence sites. We hypothesized that tree ring δ¹⁵N, as a record of ecosystem N status, could validate chronosequence assumptions and provide isotopic evidence to corroborate N trends. To test this we measured soil N availability, soil δ¹⁵N, and foliar N attributes of overstory Douglas-fir (Pseudotsuga menziesii) and understory western hemlock (Tsuga heterophylla) across three old-growth stands and nine second-growth plantations on southeast Vancouver Island, British Columbia (Canada). Increment cores for wood δ¹⁵N were retrieved from three co-dominant Douglas-fir per plot. Bulk soil δ¹⁵N was well aligned with both foliar and recent wood δ¹⁵N, demonstrating the utility of wood δ¹⁵N in monitoring ecosystem N status. Strongly contrasting trends in tree ring δ¹⁵N were evident among second-growth stands, with most trees from plantations older than 50 years exhibiting steep declines (3–4‰) in δ¹⁵N but with no temporal trends detected for younger plantations. The discrepancy in tree ring δ¹⁵N suggests disturbance history varied considerably among second-growth sites, likely because of greater slash loads and hotter broadcast burns on older cutblocks. As a consequence, the pattern of increased soil N availability and foliar N concentration with time since disturbance derived from the chronosequence could not be validated. Tree ring δ¹⁵N may provide insights into disturbance intensity, especially fire, and correlations with foliar N concentration could inform the extent of changes in stand nutrition.     

Use of foliar Ca/Sr discrimination and <Superscript>87</Superscript>Sr/<Superscript>86</Superscript>Sr ratios to determine soil Ca sources to sugar maple foliage in a northern hardwood forest

Calcium/strontium and ⁸⁷Sr/⁸⁶Sr ratios in foliage can be used to determine the relative importance of different soil sources of Ca to vegetation, if the discrimination of Ca/Sr by the plant between nutrient sources and foliage is known. We compared these tracers in the foliage of sugar maple (Acer saccharum) to the exchange fraction and acid leaches of soil horizons at six study sites in the White Mountains of New Hampshire, USA. In a previous study, sugar maple was shown to discriminate for Ca compared to Sr in foliage formation by a factor of 1.14 ± 0.12. After accounting for the predicted 14% shift in Ca/Sr, foliar Ca/Sr and ⁸⁷Sr/⁸⁶Sr ratios closely match the values in the Oie horizon at each study site across a 3.6-fold variation in foliar Ca/Sr ratios. Newly weathered cations, for which the Ca/Sr and ⁸⁷Sr/⁸⁶Sr ratios are estimated from acid leaches of soils, can be ruled out as a major Ca source to current foliage. Within sites, the ⁸⁷Sr/⁸⁶Sr ratio of the soil exchange pool in the Oa horizon and in the 0–10 cm and 10–20 cm increments of the mineral soil are similar to the Oie horizon and sugar maple foliar values, suggesting a common source of Sr in all of the actively cycling pools, but providing no help in distinguishing among them as sources to foliage. The Ca/Sr ratio in the soil exchange pool, however, decreases significantly with depth, and based on this variation, the exchange pool below the forest floor can be excluded as a major Ca source to the current sugar maple foliage. This study confirms that internal recycling of Ca between litter, organic soil horizons and vegetation dominate annual uptake of Ca in northern hardwood ecosystems. Refinement of our understanding of Ca and Sr uptake and allocation in trees allows improvement in the use of Ca/Sr and ⁸⁷Sr/⁸⁶Sr ratios to trace Ca sources to plants.     

Long-term records of strontium isotopic composition in tree rings suggest changes in forest calcium sources in the early 20th century

Many studies made in Europe and North America have shown an increasing depletion of exchangeable base cations that may cause tree nutritional deficiencies in sensitive soils. We use radial variation of strontium isotope in tree-rings (⁸⁷Sr/⁸⁶Sr ratio) to monitor possible changes in Ca sources for tree nutrition (Sr is used as an analog to Ca). The two main sources of Ca in forest stands are mineral weathering release and atmospheric inputs. Measurements in several forest stands in temperate regions show a steep decrease from pith to outer wood of the Sr isotope ratio from∼1870 to∼1920 except for stands developed on soils with a higher Ca status. This suggests a decrease of the weathering contribution (high ⁸⁷Sr/⁸⁶Sr ratio) when cations are displaced from the soil exchange complex by acid deposition at a rate faster than the replenishment of the cation pool by mineral weathering. This displacement enhances the atmospheric contribution, which is characterized by a low ⁸⁷Sr/⁸⁶Sr ratio. Tree-ring chronologies are an exceptional historic-timing record of chemical changes in the soil environment induced by atmospheric pollution. The reliability of the tree-ring recorder has been verified with a well-controlled nutritional perturbation in the context of a limed forest stand (with a known liming Sr isotopic signature). Our data suggest that forest ecosystems were affected by atmospheric inputs of strong acids earlier than previously thought.     

Nitrogen enrichment regulates calcium sources in forests

Nitrogen (N) is a key nutrient that shapes cycles of other essential elements in forests, including calcium (Ca). When N availability exceeds ecosystem demands, excess N can stimulate Ca leaching and deplete Ca from soils. Over the long term, these processes may alter the proportion of available Ca that is derived from atmospheric deposition vs. bedrock weathering, which has fundamental consequences for ecosystem properties and nutrient supply. We evaluated how landscape variation in soil N, reflecting long‐term legacies of biological N fixation, influenced plant and soil Ca availability and ecosystem Ca sources across 22 temperate forests in Oregon. We also examined interactions between soil N and bedrock Ca using soil N gradients on contrasting basaltic vs. sedimentary bedrock that differed 17‐fold in underlying Ca content. We found that low‐N forests on Ca‐rich basaltic bedrock relied strongly on Ca from weathering, but that soil N enrichment depleted readily weatherable mineral Ca and shifted forest reliance toward atmospheric Ca. Forests on Ca‐poor sedimentary bedrock relied more consistently on atmospheric Ca across all levels of soil N enrichment. The broad importance of atmospheric Ca was unexpected given active regional uplift and erosion that are thought to rejuvenate weathering supply of soil minerals. Despite different Ca sources to forests on basaltic vs. sedimentary bedrock, we observed consistent declines in plant and soil Ca availability with increasing N, regardless of the Ca content of underlying bedrock. Thus, traditional measures of Ca availability in foliage and soil exchangeable pools may poorly reflect long‐term Ca sources that sustain soil fertility. We conclude that long‐term soil N enrichment can deplete available Ca and cause forests to rely increasingly on Ca from atmospheric deposition, which may limit ecosystem Ca supply in an increasingly N‐rich world.     

Nitrogen deposition contributes to soil acidification in tropical ecosystems

Elevated anthropogenic nitrogen (N) deposition has greatly altered terrestrial ecosystem functioning, threatening ecosystem health via acidification and eutrophication in temperate and boreal forests across the northern hemisphere. However, response of forest soil acidification to N deposition has been less studied in humid tropics compared to other forest types. This study was designed to explore impacts of long‐term N deposition on soil acidification processes in tropical forests. We have established a long‐term N‐deposition experiment in an N‐rich lowland tropical forest of Southern China since 2002 with N addition as NH₄NO₃ of 0, 50, 100 and 150 kg N ha^?1 yr^?1. We measured soil acidification status and element leaching in soil drainage solution after 6‐year N addition. Results showed that our study site has been experiencing serious soil acidification and was quite acid‐sensitive showing high acidification (pH_(H2O)<4.0), negative water‐extracted acid neutralizing capacity (ANC) and low base saturation (BS,< 8%) throughout soil profiles. Long‐term N addition significantly accelerated soil acidification, leading to depleted base cations and decreased BS, and further lowered ANC. However, N addition did not alter exchangeable Al³⁺, but increased cation exchange capacity (CEC). Nitrogen addition‐induced increase in SOC is suggested to contribute to both higher CEC and lower pH. We further found that increased N addition greatly decreased soil solution pH at 20 cm depth, but not at 40 cm. Furthermore, there was no evidence that Al³⁺ was leaching out from the deeper soils. These unique responses in tropical climate likely resulted from: exchangeable H⁺ dominating changes of soil cation pool, an exhausted base cation pool, N‐addition stimulating SOC production, and N saturation. Our results suggest that long‐term N addition can contribute measurably to soil acidification, and that shortage of Ca and Mg should receive more attention than soil exchangeable Al in tropical forests with elevated N deposition in the future.     

Patterns of foliar δ15N and their control in Eastern Asian forests

Foliar δ¹⁵N has been used increasingly in research on ecosystem nitrogen (N) cycling, because it can serve as an integrator of ecosystem N cycling and thus has a potential to reveal temporal and spatial patterns of N cycling as well as how the N cycle is altered by disturbances. However, the current understanding on controls of foliar δ¹⁵N is based principally on studies from America, Europe, Australia and Africa. Here we compiled data from 65 forests at 33 sites across East Asia to explore regional patterns and what controls foliar δ¹⁵N by linking it to climate, species composition, soil depth, slope position, N deposition, and soil N availability. In East Asia, foliar δ¹⁵N ranged from ?7.1 to +2.7‰. Mean foliar δ¹⁵N values for tropical, subtropical and temperate forests were all ?3.1‰, which was unexpected. The patterns of foliar δ¹⁵N with precipitation, temperature and altitude were not clear. The variation in foliar δ¹⁵N among species and between different slope positions appeared to be small within a given forest. The δ¹⁵N for both bulk soil N and extractable inorganic N generally increased with soil depth as expected, strengthening the idea that deep-rooted trees may have access to ¹⁵N-enriched N. Different from the positive correlations reported across America and Europe, in East Asia we found that foliar δ¹⁵N decreased with increasing N deposition and did not relate to soil N availability. These discrepancies deserve more research to elucidate the mechanisms by which foliar δ¹⁵N is affected by ecosystem N availability at a regional scale.     

The effects of acid deposition on the biogeochemical cycles of major nutrients in miniature red spruce ecosystems

As part of an experimental study of air pollution effects on tree growth and health, we combined process studies with an ecosystem approach to evaluate the effects of acidic deposition on soil acidification, nutrient cycling and proton fluxes in miniature red spruce ecosystems. Ninety red spruce saplings were transplanted into 1-m diameter pots containing reconstructed soil profiles and exposed to simulated acid rain treatments of pH 3.1, 4.1 and 5.1 for four consecutive growing seasons. All the principal fluxes of the major elements were measured. During the first year of treatments, the disturbance associated with the transplanting of the experimental trees masked any treatment effects by stimulating N mineralization rates and consequent high N0₃ ⁻ cation, and H⁺ flux through the soil profile. In subsequent years, leaching of base cations and labile Al was accelerated in the most intensive acid treatment and corresponding declines in soil pH and exchangeable pools of Ca and Mg and increases in exchangeable Al concentrations were observed in the organic horizon. Leaching of Ca²⁺ and Mg²⁺ also was significantly higher in the pH 4.1 than in the pH 5.1 treatment. Flux of Ca from foliage and soil was increased in response to strong acid loading and root uptake increased to compensate for foliar Ca losses. In contrast, K cycling was dominated by root uptake and internal cycling and was relatively insensitive to strong acid inputs. Cation leaching induced by acidic deposition was responsible for the majority of H⁺ flux in the pH 3.1 treatment in the organic soil horizon whereas root uptake accounted for most of the H⁺ flux in the pH 4.1 and 5.1 treatments. Although no measurable effects on tree nutrition or health were observed, base cation leaching was significantly accelerated by acidic deposition, even at levels below that observed in the eastern U.S., warranting continued concern about acid deposition effects on the soil base status of forested ecosystems.     

Evaluation of the mobility and discrimination of Ca, Sr and Ba in forest ecosystems: consequence on the use of alkaline-earth element ratios as tracers of Ca

A comprehensive understanding of Ca cycling in an ecosystem is desirable because of the role of this element in tree mineral nutrition and its status as a major base cation on the soil exchange complex. The determination of the origin of Ca in forests is particularly indicated in regard of important changes linked to acid inputs and intensive logging. Natural strontium isotopes are increasingly used as tracers of Ca in forest ecosystems for qualitative and quantitative assessments. Nevertheless this method is limited to relatively simple systems with two sources of nutrients. Some recent studies coupled Sr/Ca or Sr/Ba ratios to Sr isotopic measurements in order to solve more complex systems. Such method has however associated with it some uncertainties: this approach assumed that Ca, Sr and Ba behave similarly throughout the ecosystem and does not take into account the Ca biopurification processes occurring in some tree’s organs which can alter element ratio. The present work focuses on two deciduous species covering large areas in Europe: European beech (Fagus sylvatica L.) and pedunculate oak (Quercus robur L.). In order to test the similarity of behaviour between Ca, Sr and Ba, their concentrations were measured extensively in the major compartments of two forest ecosystems. In parallel, the discrimination process inside tree organs was studied in 23 stands for beech and 10 stands for oak. We found that Sr and Ca behave similarly in all soil and tree compartments. By contrast, Ba and Ca appear to have contrasting behaviours, especially in streams, soil solution and soil exchange complex (no correlations between element concentrations). Sr/Ba and Ba/Ca ratios must therefore be used with care as tracer of Ca. The Ca biopurification is absent in roots and slight in bole wood but is large in bark, twigs and leaves. The discrimination factors (DF) between wood and leaves are characteristic of the two species studied and do not change significantly as a function of the soil Ca status (acidic or calcareous soils). Therefore, strontium–calcium DF can be used as a correction factor of the Sr/Ca ratio of leaves when this ratio is used in connection with Sr isotopic ratios. This correction allows to solve systems of tree nutrition with more than two sources of Ca.     

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

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

Changes in nutrient distribution in forests and soils of Walker Branch watershed,Tennessee, over an eleven-year period

Changes in vegetation, litter, and soil nutrient content were measured in selected plots on Walker Branch watershed, Tennessee, from 1972–73 to 1982. The watershed has been allowed to revert to forest since 1942, before which it consisted of small subsistence farms and woodland pastures. Changes in Ca status were of particular interest because initial nutrient cycling characterizations indicated that net Ca accumulation in vegetation could have caused large decreases in soil exchangeable Ca²⁺ within 20 years.Decreases in forest floor and subsoil (45–60 cm) N, exchangeable Ca²⁺, and Mg²⁺ content were noted in several plots from 1972 to 1982. Surface soils (0–15 cm) showed either no change or, in some cases (e.g., N and exchangeable K⁺ in certain plots), increases over the 11-year period. Reductions in forest floor and subsoil exchangeable Ca²⁺ and exchangeable Mg²⁺ on cherty, upper slope oak-hickory and chestnut oak forests were most striking. The changes in Ca²⁺ are thought to be due primarily to high rates of Ca²⁺ incorporation into woody tissues of oak and hickory species. Reductions in forest floor and subsoil exchangeable Mg²⁺ could not be accounted for by woody increment; leaching may have played a major role in causing these decreases. Changes in P and exchangeable K⁺ were variable, with both increases and decreases.There were significant increases in exchangeable Al³⁺ in both subsoils and surface soils of certain plots, but these were not accompanied by decreases in exchangeable base cations or consistent decreases in pH. Dissolution of interlayer Al from 2:1 clays may be the cause of the exchangeable Al³⁺ increases.These results suggest a general decline in fertility, especially with regard to Ca and Mg in those forests with low soil Ca and Mg supplies. Monitoring of further changes (if any) in these ecosystems will continue as the currently aggrading forests approach steady state.     

Weathering versus atmospheric sources of strontium in ecosystems on young volcanic soils

We used isotopes of Sr to quantify weathering versus atmospheric sources of foliar Sr in 34 Hawaiian forests on young volcanic soils. The forests varied widely in climate, and in lava flow age and texture. Weathering supplied most of the Sr in most of the sites, but atmospheric deposition contributed 30–50% of foliar Sr in the wettest rainforests. A stepwise multiple regression using annual precipitation, distance from the ocean, and texture of the underlying lava explained 76% of the variation in Sr isotope ratios across the sites. Substrate age did not contribute significantly to variation in Sr isotope ratios in the range of ages evaluated here (11–3000 years), although atmospheric sources eventually dominate pools of biologically available Sr in Hawaiian rainforests in older substrates (≥150,000 years). Received: 25 January 1999 / Accepted: 23 June 1999     

Impact of atmospheric nitrogen deposition on soil properties and herb-layer diversity in remnant forests along an urban–rural gradient in Guangzhou, southern China

Atmospheric nitrogen (N) deposition in subtropical metropolitan regions has increased greatly because of rapid urbanization, and such increase could lead to N-related changes in soil properties and plant diversity in remnant forests of urban ecosystems. To investigate the pattern of atmospheric N deposition along an urban?Crural gradient in metropolitan Guangzhou, southern China, and to assess the potential influence of N deposition on soil properties and understory plant diversity in remnant forests, precipitation, and soil samples were collected and vegetation was surveyed from four forest sites between March 2010 and March 2011. The atmospheric inorganic nitrogen deposition (DIN) decreased with increasing distance from the urban center: DIN inputs were 43.3, 41.2, 35.2, and 30.1?kg?N?ha^?1?year^?1 in two urban sites, a suburban site and a rural site, respectively. However, forest soil N status (NH₄ ⁺-N, NO₃ ^?-N, and total nitrogen) showed the opposite pattern. Understory herb-layer diversity was negatively correlated to DIN input and positively correlated to soil calcium (Ca) and potassium (K) concentrations and pH; with highest herb-layer diversity found in the rural site receiving the lowest amount of DIN input. These results indicated that higher DIN along with soil acidification and leaching of base cations (Ca and K) might change the current N status and increase nutrients leaching and thereby cause reductions in understory plant diversity. A regional policy linking atmospheric pollution and land protection is needed to protect the most N-sensitive herb species (e.g., forbs and ferns) in these remnant forests.     

Assessment of soil calcium status in red spruce forests in the northeastern United States

Long-term changes in concentrations of available Ca in soils of redspruce forests have been documented, but remaining questions aboutthe magnitude and regional extent of these changes have precluded anassessment of the current and future status of soil Ca. To addressthis problem, soil samples were collected in 1992—93from 12 sites in New York, Vermont, New Hampshire, and Maine toprovide additional data necessary to synthesize all availableresearch results on soil Ca in red spruce forests. Sites werechosen to encompass the range of environmental conditionsexperienced by red spruce. Concentrations of exchangeableCa ranged from 2.13 to 21.6 cmol_ckg^–1 in the Oa horizon, and from 0.11 to 0.68cmol_c kg^–1 in the upper 10 cm of theB horizon. These measurements expanded the range of exchangeable Ca reported in the literature for both horizons in northeastern redspruce forests. Exchangeable Ca was the largest Ca fraction in theforest floor at most sites (92% ofacid-extractableCa), but mineral Ca was the largest fraction at the three sites that also had the highest mineral-matter concentrations. Theprimary factor causing variability in Ca concentrations among siteswas the mineralogy of parent material, but exchangeable concentrationsin the B horizon of all sites were probably reduced by acidicdeposition. Because the majority of Ca in the forest floor isin a readily leachable form, and Ca inputs to the forest floor from the mineral soil and atmospheric deposition have beendecreasing in recent decades, the previously documented decreasesin Ca concentrations in the forest floor over previous decades mayextend into the future.     

Interactive Effects of Vegetation Type and Topographic Position on Nitrogen Availability and Loss in a Temperate Montane Ecosystem

Determining the fate of deposited nitrogen (N) in natural ecosystems remains a challenge. Heterogeneity of vegetation types and resulting plant–soil feedbacks interact with topo-hydrologic gradients to mediate spatial patterns of N availability and loss, yet net effects of variation in these two factors together across complex terrain remain unclear. Here we measured a suite of N-cycle pools and fluxes in sites that differed factorially in vegetation type (mixed forest vs. herbaceous) and topographic position (upslope vs. downslope) in a protected montane watershed near Salt Lake City, UT. Vegetation type was associated with large variation in N availability—herbaceous sites had larger NO₃ ^? pools, higher NO₃ ^?:NH₄ ⁺ ratios, higher nitrification potentials, lower soil C:N values, enriched δ¹⁵N values, and lower microbial biomass compared to forests, especially those upslope. Downslope sites tended to exhibit higher N availability and indicators of N-cycle openness, but patterns were moderated by vegetation type. In downslope forest, soil NO₃ ^? depth profiles and higher foliar N content suggested trees were accessing deep soil N and transferring it to the surface via litterfall, while more deep soil NO₃ ^? but no change in surface or foliar N suggested herbaceous cover was not N limited or deeper N pools were not accessible. Soil NO₃ ^? leaching from below the rooting zone closely tracked N availability, revealing a link between N status and hydrologic loss as well as an important role for roots in N retention. NO₃ ^? isotopes did not reveal a similar link for gaseous losses (that is, denitrification), instead reflecting nitrification and/or transport dynamics. Together, these results suggest a coupled ecological, topo-hydrologic perspective can help assess the fate of N in complex landscapes.     

Erosion and landscape development affect plant nutrient status in the Hawaiian Islands

We quantified variation in plant nutrient concentrations and provenance along catenas in landscapes of three different ages (0.15, 1.4, and 4.1 ma) in the Hawaiian Islands. Strontium (Sr) isotopes demonstrate that erosion provides a renewed source of rock-derived nutrients to slopes in landscapes of all ages, in some cases reversing a million years of ecosystem development in a distance of 100 m. However the effects of this input vary with landscape age. Plants on uneroded surfaces in a 0.15-ma landscape derive ~20% of their Sr from local bedrock (foliar ⁸⁷Sr/⁸⁶Sr~0.7085), while on adjacent slopes this increases to ~80% (foliar ⁸⁷Sr/⁸⁶Sr~0.7045). Despite this shift in provenance, foliar N and P do not vary systematically with slope position. Conversely, eroded slopes in a 4.1-ma landscape show smaller increases in rock-derived cations relative to stable uplands (foliar ⁸⁷Sr/⁸⁶Sr~0.7075 vs 0.7090), but have >50% higher foliar N and P. These results demonstrate both that erosion can greatly increase nutrient availability in older landscapes, and that the ecological effects of erosion vary with landscape age. In addition, there can be as much biogeochemical variation on fine spatial scales in eroding landscapes as there is across millions of years of ecosystem development on stable surfaces.     

Variations in nitrogen-15 natural abundance of plant and soil systems in four remote tropical rainforests, southern China

The foliar stable N isotope ratio (δ¹⁵N) can provide integrated information on ecosystem N cycling. Here we present the δ¹⁵N of plant and soil in four remote typical tropical rainforests (one primary and three secondary) of southern China. We aimed to examine if (1) foliar δ¹⁵N in the study forests is negative, as observed in other tropical and subtropical sites in eastern Asia; (2) variation in δ¹⁵N among different species is smaller compared to that in many N-limited temperate and boreal ecosystems; and (3) the primary forest is more N rich than the younger secondary forests and therefore is more ¹⁵N enriched. Our results show that foliar δ¹⁵N ranged from ?5.1 to 1.3 ‰ for 39 collected plant species with different growth strategies and mycorrhizal types, and that for 35 species it was negative. Soil NO₃ ^? had low δ¹⁵N (?11.4 to ?3.2 ‰) and plant NO₃ ^? uptake could not explain the negative foliar δ¹⁵N values (NH₄ ⁺ was dominant in the soil inorganic-N fraction). We suggest that negative values might be caused by isotope fractionation during soil NH₄ ⁺ uptake and mycorrhizal N transfer, and by direct uptake of atmospheric NH₃/NH₄ ⁺. The variation in foliar δ¹⁵N among species (by about 6 ‰) was smaller than in many N-limited ecosystems, which is typically about or over 10 ‰. The primary forest had a larger N capital in plants than the secondary forests. Foliar δ¹⁵N and the enrichment factor (foliar δ¹⁵N minus soil δ¹⁵N) were higher in the primary forest than in the secondary forests, albeit differences were small, while there was no consistent pattern in soil δ¹⁵N between primary and secondary forests.     

Nitrogen stable isotopes indicate differences in nitrogen cycling between two contrasting Jamaican montane forests

Background and aims

The aim of this study is to enhance our knowledge of nitrogen (N) cycling and N acquisition in tropical montane forests through analysis of stable N isotopes (δ¹⁵N).

Methods

Leaves from eight common tree species, leaf litter, soils from three depths and roots were sampled from two contrasting montane forest types in Jamaica (mull ridge and mor ridge) and were analysed for δ¹⁵N.

Results

All foliar δ¹⁵N values were negative and varied among the tree species but were significantly more negative in the mor ridge forest (by about 2 ‰). δ¹⁵N of soils and roots were also more negative in mor ridge forests by about 3 ‰. Foliar δ¹⁵N values were closer to that of soil ammonium than soil nitrate suggesting that trees in these forests may have a preference for ammonium; this may explain the high losses of nitrate from similar tropical montane forests. There was no correlation between the rankings of foliar δ¹⁵N in the two forest types suggesting a changing uptake ratio of different N forms between forest types.

Conclusions

These results indicate that N is found at low concentrations in this ecosystem and that there is a tighter N cycle in the mor ridge forest, confirmed by reduced nitrogen availability and lower rates of nitrification. Overall, soil or root δ¹⁵N values are more useful in assessing ecosystem N cycling patterns as different tree species showed differences in foliar δ¹⁵N between the two forest types.