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Steven S. Perakis

Publications and source records attributed to Steven S. Perakis.

At least 55 records · Page 3Linked to original sources

Decomposition drives convergence of forest litter nutrient stoichiometry following phosphorus addition

Background and aims Nutrient levels in decomposing detritus and soil can influence decomposition rates and detrital nutrient dynamics in differing ways among various detrital components of forests. We assessed whether increased phosphorus (P) levels in litter and soil influenced decomposition rates and litter nutrient dynamics of foliage, fine roots, and twigs in nitrogen (N)-rich Douglas-fir ( Pseudotsuga menziesii (Mirb.) Franco) forests in the Oregon Coast Range. Methods We decomposed fresh foliage, fine root, and twig litter from Douglas-fir seedlings at three sites for two years. Half of the seedlings and half of the plots at each of the sites were fertilized with P resulting in a factorial design with the following treatments: control (no P fertilization), plant P (P-fertilized litter), soil P (P-fertilized soil), and plant P × soil P. Results Soil P fertilization slightly decreased foliage decomposition rates. Fertilization of seedlings increased litter P concentrations by an average of 250 % relative to controls, but did not alter litter decomposition rates. Litter fertilized with P mineralized P rapidly and early in the decomposition process compared to N. Litter P concentrations decreased over the 2 years for all treatments, whereas N concentrations increased. Decomposition rates and loss of N and P were strongly related to initial litter chemistry. Despite different initial litter C:N:P ratios in P fertilized seedlings, ratios of C:N, C:P and N:P converged to similar values across treatments within a given litter type over 2 years. Conclusions We conclude that litter P concentrations and to some extent soil P may influence litter nutrient dynamics during decomposition, resulting in a convergence of element ratios that reflect the balance of substrate decomposition and microbial nutrient stoichiometry.

Oregon Coast Range↗

A framework to assess biogeochemical response to ecosystem disturbance using nutrient partitioning ratios

Disturbances affect almost all terrestrial ecosystems, but it has been difficult to identify general principles regarding these influences. To improve our understanding of the long-term consequences of disturbance on terrestrial ecosystems, we present a conceptual framework that analyzes disturbances by their biogeochemical impacts. We posit that the ratio of soil and plant nutrient stocks in mature ecosystems represents a characteristic site property. Focusing on nitrogen (N), we hypothesize that this partitioning ratio (soil N: plant N) will undergo a predictable trajectory after disturbance. We investigate the nature of this partitioning ratio with three approaches: (1) nutrient stock data from forested ecosystems in North America, (2) a process-based ecosystem model, and (3) conceptual shifts in site nutrient availability with altered disturbance frequency. Partitioning ratios could be applied to a variety of ecosystems and successional states, allowing for improved temporal scaling of disturbance events. The generally short-term empirical evidence for recovery trajectories of nutrient stocks and partitioning ratios suggests two areas for future research. First, we need to recognize and quantify how disturbance effects can be accreting or depleting, depending on whether their net effect is to increase or decrease ecosystem nutrient stocks. Second, we need to test how altered disturbance frequencies from the present state may be constructive or destructive in their effects on biogeochemical cycling and nutrient availability. Long-term studies, with repeated sampling of soils and vegetation, will be essential in further developing this framework of biogeochemical response to disturbance.

Ecosystems↗

Response of the nitrogen-fixing lichen Lobaria pulmonaria to phosphorus, molybdenum, and vanadium

Nitrogen-fixing lichens (cyanolichens) are an important source of nitrogen (N) in Pacific Northwest forests, but limitation of lichen growth by elements essential for N fixation is poorly understood. To investigate how nutrient limitation may affect cyanolichen growth rates, we fertilized a tripartite cyanobacterial lichen ( Lobaria pulmonaria ) and a green algal non-nitrogen fixing lichen ( Usnea longissima ) with the micronutrients molybdenum (Mo) and vanadium (V), both known cofactors for enzymes involved in N fixation, and the macronutrient phosphorus (P). We then grew treated lichens in the field for one year in western Oregon, USA. Lichen growth was very rapid for both species and did not differ across treatments, despite a previous demonstration of P-limitation in L. pulmonaria at a nearby location. To reconcile these disparate findings, we analyzed P, Mo, and V concentrations, natural abundance δ 15 N isotopes, %N and change in thallus N in Lobaria pulmonaria from both growth experiments. Nitrogen levels in deposition and in lichens could not explain the large difference in growth or P limitation observed between the two studies. Instead, we provide evidence that local differences in P availability may have caused site-specific responses of Lobaria to P fertilization. In the previous experiment, Lobaria had low background levels of P, and treatment with P more than doubled growth. In contrast, Lobaria from the current experiment had much higher background P concentrations, similar to P-treated lichens in the previous experiment, consistent with the idea that ambient variation in P availability influences the degree of P limitation in cyanolichens. We conclude that insufficient P, Mo, and V did not limit the growth of either cyanolichens or chlorolichens at the site of the current experiment. Our findings point to the need to understand landscape-scale variation in P availability to cyanolichens, and its effect on spatial patterns of cyanolichen nutrient limitation and N fixation.

Oregon↗

Soil organic matter regulates molybdenum storage and mobility in forests

The trace element molybdenum (Mo) is essential to a suite of nitrogen (N) cycling processes in ecosystems, but there is limited information on its distribution within soils and relationship to plant and bedrock pools. We examined soil, bedrock, and plant Mo variation across 24 forests spanning wide soil pH gradients on both basaltic and sedimentary lithologies in the Oregon Coast Range. We found that the oxidizable organic fraction of surface mineral soil accounted for an average of 33 %of bulk soil Mo across all sites, followed by 1.4 % associated with reducible Fe, Al, and Mn-oxides, and 1.4 % in exchangeable ion form. Exchangeable Mo was greatest at low pH, and its positive correlation with soil carbon (C) suggests organic matter as the source of readily exchangeable Mo. Molybdenum accumulation integrated over soil profiles to 1 m depth ( τMo Nb ) increased with soil C, indicating that soil organic matter regulates long-term Mo retention and loss from soil. Foliar Mo concentrations displayed no relationship with bulk soil Mo, and were not correlated with organic horizon Mo or soil extractable Mo, suggesting active plant regulation of Mo uptake and/or poor fidelity of extractable pools to bioavailability. We estimate from precipitation sampling that atmospheric deposition supplies, on average, over 10 times more Mo annually than does litterfall to soil. In contrast, bedrock lithology had negligible effects on foliar and soil Mo concentrations and on Mo distribution among soil fractions. We conclude that atmospheric inputs may be a significant source of Mo to forest ecosystems, and that strong Mo retention by soil organic matter limits ecosystem Mo loss via dissolution and leaching pathways.

Oregon↗

Effects and empirical critical loads of Nitrogen for ecoregions of the United States

Human activity in the last century has increased nitrogen (N) deposition to a level that has caused or is likely to cause alterations to the structure and function of many ecosystems across the United States. We synthesized current research relating atmospheric N deposition to effects on terrestrial and freshwater ecosystems in the United States, and estimated associated empirical critical loads of N for several receptors: freshwater diatoms, mycorrhizal fungi, lichens, bryophytes, herbaceous plants, shrubs, and trees. Biogeochemical responses included increased N mineralization and nitrification, increased gaseous N losses, and increased N leaching. Individual species, population, and community responses included increased tissue N, physiological and nutrient imbalances, increased growth, altered root-shoot ratios, increased susceptibility to secondary stresses, altered fire regime, shifts in competitive interactions and community composition, changes in species richness and other measures of biodiversity, and increases in invasive species. The range of critical loads of nutrient N reported for U.S. ecoregions, inland surface waters, and freshwater wetlands is 1–39 kg N ha−1 yr−1, spanning the range of N deposition observed over most of the country. The empirical critical loads of N tend to increase in the following sequence: diatoms, lichens and bryophytes, mycorrhizal fungi, herbaceous plants and shrubs, trees.

Book chapter↗

Disturbance and topography shape nitrogen availability and δ15 N over long-term forest succession

Forest disturbance and long-term succession towards old-growth are thought to increase nitrogen (N) availability and N loss, which should increase soil δ15N values. We examined soil and foliar patterns in N and δ15N, and soil N mineralization, across 800 years of forest succession in a topographically complex montane landscape influenced by human logging and wildfire. In contrast to expectations, we found that disturbance caused declines in surface mineral soil δ15N values, both in logged forests measured 40–50 years after disturbance, and in unlogged forests disturbed by severe wildfire within the last 200 years. Both symbiotic N fixation and N transfers from disturbed vegetation and detritus could lower soil δ15N values after disturbance. A more important role for symbiotic N fixation is suggested by lower soil δ15N values in slow-successional sites with slow canopy closure, which favors early-successional N fixers. Soil δ15N values increased only marginally throughout 800 years of succession, reflecting soil N uptake by vegetation and strong overall N retention. Although post-disturbance N inputs lowered surface soil δ15N values, steady-state mass balance calculations suggest that wildfire combustion of vegetation and detritus can dominate long-term N loss and increase whole-ecosystem δ15N. On steeper topography, declining soil δ15N values highlight erosion and accelerated soil turnover as an additional abiotic control on N balances. We conclude for N-limited montane forests that soil δ15N and N availability are less influenced by nitrate leaching and denitrification loss than by interactions between disturbance, N fixation, and erosion.

Ecosystems↗

Convergence of soil nitrogen isotopes across global climate gradients

Quantifying global patterns of terrestrial nitrogen (N) cycling is central to predicting future patterns of primary productivity, carbon sequestration, nutrient fluxes to aquatic systems, and climate forcing. With limited direct measures of soil N cycling at the global scale, syntheses of the 15 N: 14 N ratio of soil organic matter across climate gradients provide key insights into understanding global patterns of N cycling. In synthesizing data from over 6000 soil samples, we show strong global relationships among soil N isotopes, mean annual temperature (MAT), mean annual precipitation (MAP), and the concentrations of organic carbon and clay in soil. In both hot ecosystems and dry ecosystems, soil organic matter was more enriched in 15 N than in corresponding cold ecosystems or wet ecosystems. Below a MAT of 9.8°C, soil δ 15 N was invariant with MAT. At the global scale, soil organic C concentrations also declined with increasing MAT and decreasing MAP. After standardizing for variation among mineral soils in soil C and clay concentrations, soil δ 15 N showed no consistent trends across global climate and latitudinal gradients. Our analyses could place new constraints on interpretations of patterns of ecosystem N cycling and global budgets of gaseous N loss.

Scientific Reports↗

Reconstructing disturbances and their biogeochemical consequences over multiple timescales

Ongoing changes in disturbance regimes are predicted to cause acute changes in ecosystem structure and function in the coming decades, but many aspects of these predictions are uncertain. A key challenge is to improve the predictability of postdisturbance biogeochemical trajectories at the ecosystem level. Ecosystem ecologists and paleoecologists have generated complementary data sets about disturbance (type, severity, frequency) and ecosystem response (net primary productivity, nutrient cycling) spanning decadal to millennial timescales. Here, we take the first steps toward a full integration of these data sets by reviewing how disturbances are reconstructed using dendrochronological and sedimentary archives and by summarizing the conceptual frameworks for carbon, nitrogen, and hydrologic responses to disturbances. Key research priorities include further development of paleoecological techniques that reconstruct both disturbances and terrestrial ecosystem dynamics. In addition, mechanistic detail from disturbance experiments, long-term observations, and chronosequences can help increase the understanding of ecosystem resilience.

BioScience↗

Three-year growth response of young Douglas-fir to nitrogen, calcium, phosphorus, and blended fertilizers in Oregon and Washington

Studies of nutrient limitation in Douglas-fir forests of the Pacific Northwest focus predominantly on nitrogen, yet many stands demonstrate negligible or even negative growth response to nitrogen fertilization. To understand what nutrients other than nitrogen may limit forest productivity in this region, we tested six fertilizer treatments for their ability to increase stem volume growth response of dominant and co-dominant trees in young Douglas-fir plantations across a range of foliar and soil chemistry in western Oregon and Washington. We evaluated responses to single applications of urea, lime, calcium chloride, or monosodium phosphate at 16 sites, and to two site-specific nutrients blends at 12 of these sites. Across sites, the average stem volume growth increased marginally with urea, lime, and phosphorus fertilization. Fertilization responses generally aligned with plant and soil indicators of nutrient limitation. Response to nitrogen addition was greatest on soils with low total nitrogen and high exchangeable calcium concentrations. Responses to lime and calcium chloride additions were greatest at sites with low foliar calcium and low soil pH. Response to phosphorus addition was greatest on sites with low foliar phosphorus and high soil pH. Blended fertilizers yielded only marginal growth increases at one site, with no consistent effect across sites. Overall, our results highlight that calcium and phosphorus can be important growth limiting nutrients on specific sites in nitrogen-rich Douglas-fir forests of the Pacific Northwest.

Oregon, Washington↗

Calcium oxalate contribution to calcium cycling in forests of contrasting nutrient status

Calcium oxalate (Ca oxalate) is an insoluble biomineral that forms in plants and fungi, and occurs in soils across many types of ecosystems. Assessing how Ca oxalate may shape ecosystem Ca cycling requires information on the distribution of Ca oxalate among plant biomass, detritus, and mineral soil, and how it varies with ecosystem Ca status. We compared two Douglas-fir forests of contrasting ecosystem Ca availability, and found that Ca oxalate was partitioned similarly among plant biomass, detritus and mineral soil major ecosystem compartments at both sites, and total pools of Ca oxalate were greater in the high-Ca forest. However, the proportional importance of Ca oxalate was greater in the low-Ca than high-Ca forest (18% versus 4% of actively cycling ecosystem Ca, respectively). And calcium oxalate in mineral soil, which is of particular interest as a potential long-term Ca reservoir, was a larger portion of total available Ca (exchangeable Ca plus Ca oxalate Ca) in the low-Ca site than the high-Ca site (9% versus 1% of available soil Ca, respectively). Calcium oxalate was the dominant form of Ca returned from plants to soil as leaf litterfall at the high-Ca site, yet calcium oxalate disappeared rapidly from decomposing litter (0.28 yr−1 or faster) at both sites. We conclude that accumulation of Ca oxalate in forest ecosystems appears most closely related to overall Ca supply for live biomass pools, and that the accumulation of Ca oxalate in forest floor and mineral soil is limited by rapid microbial degradation of putatively unavailable Ca oxalate.

Forest Ecology and Management↗

Soil fluxes of methane, nitrous oxide, and nitric oxide from aggrading forests in coastal Oregon

Soil exchanges of greenhouse and other gases are poorly known for Pacific Northwest forests where gradients in nutrient availability and soil moisture may contribute to large variations in fluxes. Here we report fluxes of methane (CH 4 ), nitrous oxide (N 2 O), and nitric oxide (NO) over multiple seasons from three naturally N-rich, aggrading forests of coastal Oregon, USA. Mean methane uptake rates (3.2 mg CH 4 m −2 d −1 ) were high compared with forests globally, negatively related to water-filled pore space (WFPS), but unrelated to N availability or temperature. Emissions of NO (6.0 μg NO–N m −2 h −1 ) exceeded N 2 O (1.4 μg N 2 O–N m −2 h −1 ), except when WFPS surpassed 55%. Spatial variation in NO fluxes correlated positively with soil nitrate concentrations (which generally exceeded ammonium concentrations, indicating the overall high N status for the sites) and negatively with soil pH, and at one site increased with basal area of N 2 -fixing red alder. Combined NO and N 2 O emissions were greatest from the site with highest annual net N mineralization and lowest needle litterfall C/N. Our findings of high CH 4 uptake and NO/N 2 O ratios generally >1 most likely reflect the high porosity of the andic soils underlying the widespread regenerating forests in this seasonally wet region.

Oregon↗

Thresholds for protecting Pacific Northwest ecosystems from atmospheric deposition of nitrogen: state of knowledge report

The National Park Service and U.S. Forest Service manage areas in the states of Idaho, Oregon, and Washington – collectively referred to in this report as the Pacific Northwest - that contain significant natural resources and provide many recreational opportunities. The agencies are mandated to protect the air quality and air pollution-sensitive resources on these federal lands. Human activity has greatly increased the amount of nitrogen emitted to the atmosphere, resulting in elevated amounts of nitrogen being deposited in park and forest ecosystems. There is limited information in the Pacific Northwest about the levels of nitrogen that negatively affect natural systems, i.e., the critical loads. The National Park Service and U.S. Forest Service, with scientific input from the U.S. Geological Survey, have developed an approach for accumulating additional nitrogen critical loads information in the Pacific Northwest and using the data in planning and regulatory arenas. As a first step in that process, this report summarizes the current state of knowledge about nitrogen deposition, effects, and critical loads in the region. It also describes ongoing research efforts and identifies and prioritizes additional data needs.

Idaho;Oregon;Washington↗

Nutrient limitation of native and invasive N 2 -fixing plants in northwest prairies

Nutrient rich conditions often promote plant invasions, yet additions of non-nitrogen (N) nutrients may provide a novel approach for conserving native symbiotic N-fixing plants in otherwise N-limited ecosystems. Lupinus oreganus is a threatened N-fixing plant endemic to prairies in western Oregon and southwest Washington (USA). We tested the effect of non-N fertilizers on the growth, reproduction, tissue N content, and stable isotope δ 15 N composition of Lupinus at three sites that differed in soil phosphorus (P) and N availability. We also examined changes in other Fabaceae (primarily Vicia sativa and V. hirsuta) and cover of all plant species. Variation in background soil P and N availability shaped patterns of nutrient limitation across sites. Where soil P and N were low, P additions increased Lupinus tissue N and altered foliar δ 15 N, suggesting P limitation of N fixation. Where soil P was low but N was high, P addition stimulated growth and reproduction in Lupinus. At a third site, with higher soil P, only micro- and macronutrient fertilization without N and P increased Lupinus growth and tissue N. Lupinus foliar δ 15 N averaged −0.010‰ across all treatments and varied little with tissue N, suggesting consistent use of fixed N. In contrast, foliar δ 15 N of Vicia spp. shifted towards 0‰ as tissue N increased, suggesting that conditions fostering N fixation may benefit these exotic species. Fertilization increased cover, N fixation, and tissue N of non-target, exotic Fabaceae, but overall plant community structure shifted at only one site, and only after the dominant Lupinus was excluded from analyses. Our finding that non-N fertilization increased the performance of Lupinus with few community effects suggests a potential strategy to aid populations of threatened legume species. The increase in exotic Fabaceae species that occurred with fertilization further suggests that monitoring and adaptive management should accompany any large scale applications.

Oregon↗

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

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

Oecologia↗

Nitrate in watersheds: straight from soils to streams?

Human activities are rapidly increasing the global supply of reactive N and substantially altering the structure and hydrologic connectivity of managed ecosystems. There is long-standing recognition that N must be removed along hydrologic flowpaths from uplands to streams, yet it has proven difficult to assess the generality of this removal across ecosystem types, and whether these patterns are influenced by land-use change. To assess how well upland nitrate (NO 3 - ) loss is reflected in stream export, we gathered information from >50 watershed biogeochemical studies that reported nitrate concentrations ([NO 3 - ]) for stream water and for either upslope soil solution or groundwater NO 3 - to examine whether stream export of NO3- accurately reflects upland NO 3 - losses. In this dataset, soil solution and streamwater [NO 3 - ] were correlated across 40 undisturbed forest watersheds, with streamwater [NO 3 - ] typically half (median = 50%) soil solution [NO 3 - ]. A similar relationship was seen in 10 disturbed forest watersheds. However, for 12 watersheds with significant agricultural or urban development, the intercept and slope were both significantly higher than the relationship seen in forest watersheds. Differences in concentration between soil solution or groundwater and stream water may be attributed to biological uptake, microbial processes including denitrification, and/or preferential flow routing. The results of this synthesis are consistent with the hypotheses that undisturbed watersheds have a significant capacity to remove nitrate after it passes below the rooting zone and that land use changes tend to alter the efficiency or the length of watershed flowpaths, leading to reductions in nitrate removal and increased stream nitrate concentrations.

Journal of Geophysical Research G: Biogeosciences↗

Contribution of calcium oxalate to soil-exchangeable calcium

Acid deposition and repeated biomass harvest have decreased soil calcium (Ca) availability in many temperate forests worldwide, yet existing methods for assessing available soil Ca do not fully characterize soil Ca forms. To account for discrepancies in ecosystem Ca budgets, it has been hypothesized that the highly insoluble biomineral Ca oxalate might represent an additional soil Ca pool that is not detected in standard measures of soil-exchangeable Ca. We asked whether several standard method extractants for soil-exchangeable Ca could also access Ca held in Ca oxalate crystals using spike recovery tests in both pure solutions and soil extractions. In solutions of the extractants ammonium chloride, ammonium acetate, and barium chloride, we observed 2% to 104% dissolution of Ca oxalate crystals, with dissolution increasing with both solution molarity and ionic potential of cation extractant. In spike recovery tests using a low-Ca soil, we estimate that 1 M ammonium acetate extraction dissolved sufficient Ca oxalate to contribute an additional 52% to standard measurements of soil-exchangeable Ca. However, in a high-Ca soil, the amount of Ca oxalate spike that would dissolve in 1 M ammonium acetate extraction was difficult to detect against the large pool of exchangeable Ca. We conclude that Ca oxalate can contribute substantially to standard estimates of soil-exchangeable Ca in acid forest soils with low soil-exchangeable Ca. Consequently, measures of exchangeable Ca are unlikely to fully resolve discrepancies in ecosystem Ca mass balance unless the contribution of Ca oxalate to exchangeable Ca is also assessed.

Soil Science↗

Regional constraints to biological nitrogen fixation in post-fire forest communities

Biological nitrogen fixation (BNF) is a key ecological process that can restore nitrogen (N) lost in wildfire and shape the pace and pattern of post-fire forest recovery. To date, there is limited information on how climate and soil fertility interact to influence different pathways of BNF in early forest succession. We studied asymbiotic (forest floor and soil) and symbiotic (the shrub Ceanothus integerrimus) BNF rates across six sites in the Klamath National Forest, California, USA. We used combined gradient and experimental phosphorus (P) fertilization studies to explore cross-site variation in BNF rates and then related these rates to abiotic and biotic variables. We estimate that our measured BNF rates 22 years after wildfire (6.1–12.1 kg N·ha -1 ·yr -1 ) are unlikely to fully replace wildfire N losses. We found that asymbiotic BNF is P limited, although this is not the case for symbiotic BNF in Ceanothus. In contrast, Ceanothus BNF is largely driven by competition from other vegetation: in high-productivity sites with high potential evapotranspiration (Et), shrub biomass is suppressed as tree biomass increases. Because shrub biomass governed cross-site variation in Ceanothus BNF, this competitive interaction led to lower BNF in sites with high productivity and Et. Overall, these results suggest that the effects of nutrients play a larger role in driving asymbiotic than symbiotic fixation across our post-fire sites. However, because symbiotic BNF is 8–90 x greater than asymbiotic BNF, it is interspecific plant competition that governs overall BNF inputs in these forests.

California↗

Forest calcium depletion and biotic retention along a soil nitrogen gradient

High nitrogen (N) accumulation in terrestrial ecosystems can shift patterns of nutrient limitation and deficiency beyond N toward other nutrients, most notably phosphorus (P) and base cations (calcium [Ca], magnesium [Mg], and potassium [K]). We examined how naturally high N accumulation from a legacy of symbiotic N fixation shaped P and base cation cycling across a gradient of nine temperate conifer forests in the Oregon Coast Range. We were particularly interested in whether long-term legacies of symbiotic N fixation promoted coupled N and organic P accumulation in soils, and whether biotic demands by non-fixing vegetation could conserve ecosystem base cations as N accumulated. Total soil N (0–100 cm) pools increased nearly threefold across the N gradient, leading to increased nitrate leaching, declines in soil pH from 5.8 to 4.2, 10-fold declines in soil exchangeable Ca, Mg, and K, and increased mobilization of aluminum. These results suggest that long-term N enrichment had acidified soils and depleted much of the readily weatherable base cation pool. Soil organic P increased with both soil N and C across the gradient, but soil inorganic P, biomass P, and P leaching loss did not vary with N, implying that historic symbiotic N fixation promoted soil organic P accumulation and P sufficiency for non-fixers. Even though soil pools of Ca, Mg, and K all declined as soil N increased, only Ca declined in biomass pools, suggesting the emergence of Ca deficiency at high N. Biotic conservation and tight recycling of Ca increased in response to whole-ecosystem Ca depletion, as indicated by preferential accumulation of Ca in biomass and surface soil. Our findings support a hierarchical model of coupled N–Ca cycling under long-term soil N enrichment, whereby ecosystem-level N saturation and nitrate leaching deplete readily available soil Ca, stimulating biotic Ca conservation as overall supply diminishes. We conclude that a legacy of biological N fixation can increase N and P accumulation in soil organic matter to the point that neither nutrient is limiting to subsequent non-fixers, while also resulting in natural N saturation that intensifies base cation depletion and deficiency.

Oregon↗