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Steven S. Perakis

Publications and source records attributed to Steven S. Perakis.

At least 37 records · Page 2Linked to original sources

A roadmap for sampling and scaling biological nitrogen fixation in terrestrial ecosystems

Accurately quantifying rates and patterns of biological nitrogen fixation (BNF) in terrestrial ecosystems is essential to characterize ecological and biogeochemical interactions, identify mechanistic controls, improve BNF representation in conceptual and numerical modelling, and forecast nitrogen limitation constraints on future carbon (C) cycling. While many resources address the technical advantages and limitations of different methods for measuring BNF, less systematic consideration has been given to the broader decisions involved in planning studies, interpreting data, and extrapolating results. Here, we present a conceptual and practical road map to study design, study execution, data analysis and scaling, outlining key considerations at each step. We address issues including defining N‐fixing niches of interest, identifying important sources of temporal and spatial heterogeneity, designing a sampling scheme (including method selection, measurement conditions, replication, and consideration of hotspots and hot moments), and approaches to analysing, scaling and reporting BNF. We also review the comparability of estimates derived using different approaches in the literature, and provide sample R code for simulating symbiotic BNF data frames and upscaling. Improving and standardizing study design at each of these stages will improve the accuracy and interpretability of data, define limits of extrapolation, and facilitate broader use of BNF data for downstream applications. We highlight aspects—such as quantifying scales of heterogeneity, statistical approaches for dealing with non‐normality, and consideration of rates versus ecological significance—that are ripe for further development.

Methods in Ecology and Evolution↗

Climate-mediated changes to linked terrestrial and marine ecosystems across the northeast Pacific coastal temperate rainforest margin

Coastal margins are important areas of materials flux that link terrestrial and marine ecosystems. Consequently, climate-mediated changes to coastal terrestrial ecosystems and hydrologic regimes have high potential to influence nearshore ocean chemistry and food web dynamics. Research from tightly coupled, high-flux coastal ecosystems can advance understanding of terrestrial–marine links and climate sensitivities more generally. In the present article, we use the northeast Pacific coastal temperate rainforest as a model system to evaluate such links. We focus on key above- and belowground production and hydrological transport processes that control the land-to-ocean flow of materials and their influence on nearshore marine ecosystems. We evaluate how these connections may be altered by global climate change and we identify knowledge gaps in our understanding of the source, transport, and fate of terrestrial materials along this coastal margin. Finally, we propose five priority research themes in this region that are relevant for understanding coastal ecosystem links more broadly.

BioScience↗

Decadal-scale decoupling of soil phosphorus and molybdenum cycles by temperate nitrogen-fixing trees

Symbiotic nitrogen- (N) fixing trees can influence multiple biogeochemical cycles by fixing atmospheric N, which drives net primary productivity and soil carbon (C) and N accumulation, as well as by mobilizing soil phosphorus (P) and other nutrients to support growth and metabolism. The soil micronutrient molybdenum (Mo) is essential to N-fixation, yet surprisingly little is known of whether N-fixing trees alter soil Mo cycling, and if changes to soil Mo are coupled to soil C, N, and P. We compared how symbiotic N-fixing red alder and non-N-fixing Douglas-fir trees modified surface soil C, N, P, and Mo across variation in climate and other site factors in the Pacific Northwest. We found that after two decades, N-fixing trees drove coupled increases in surface soil C, N, total P, and organic P. Consistent with contributions of N-fixing trees to soil organic matter, increased soil C and N were accompanied by lower δ13C in all sites, and lower δ15N in sites where non-fixer plots exhibited elevated soil δ15N. However, N-fixing trees did not affect surface soil Mo concentrations or fractions, suggesting that different factors control the cycling of P versus Mo over decadal timescales. Random forest analysis revealed that surface soil P was most strongly influenced by factors related to soil C accumulation, whereas surface soil Mo was related primarily to environmental factors, including potential differences in atmospheric Mo deposition across sites. Ratios of surface soil P:Mo were higher in extractable pools than in total soil digests, reinforcing the idea of stronger biotic cycling of P than Mo. Overall, our multi-site, multi-decadal field study found surprisingly small effects of N-fixing trees on soil Mo, despite rapid increases in soil organic C, N, and P. We hypothesize that, rather than direct effects of N-fixing vegetation, abiotic or indirect biotic factors such as soil sorption of atmospheric Mo inputs can link C–N–P–Mo cycles in terrestrial ecosystems on longer timescales.

British Columbia, Washington↗

A spatially explicit, empirical estimate of tree-based biological nitrogen fixation in forests of the United States

Quantifying human impacts on the nitrogen (N) cycle and investigating natural ecosystem N cycling depend on the magnitude of inputs from natural biological nitrogen fixation (BNF). Here, we present two bottom‐up approaches to quantify tree‐based symbiotic BNF based on forest inventory data across the coterminous United States and SE Alaska. For all major N‐fixing tree genera, we quantify BNF inputs using (1) ecosystem N accretion rates (kg N ha −1 yr −1 ) scaled with spatial data on tree abundance and (2) percent of N derived from fixation (%N dfa ) scaled with tree N demand (from tree growth rates and stoichiometry). We estimate that trees fix 0.30–0.88 Tg N yr −1 across the study area (1.4–3.4 kg N ha −1 yr −1 ). Tree‐based N fixation displays distinct spatial variation that is dominated by two genera, Robinia (64% of tree‐associated BNF) and Alnus (24%). The third most important genus, Prosopis , accounted for 5%. Compared to published estimates of other N fluxes, tree‐associated BNF accounted for 0.59 Tg N yr −1 , similar to asymbiotic (0.37 Tg N yr −1 ) and understory symbiotic BNF (0.48 Tg N yr −1 ), while N deposition contributed 1.68 Tg N yr −1 and rock weathering 0.37 Tg N yr −1 . Overall, our results reveal previously uncharacterized spatial patterns in tree BNF that can inform large‐scale N assessments and serve as a model for improving tree‐based BNF estimates worldwide. This updated, lower BNF estimate indicates a greater ratio of anthropogenic to natural N inputs, suggesting an even greater human impact on the N cycle.

Global Biogeochemical Cycles↗

Frequent burning causes large losses of carbon from deep soil layers in a temperate savanna

Fire activity is changing dramatically across the globe, with uncertain effects on ecosystem processes, especially below‐ground. Fire‐driven losses of soil carbon (C) are often assumed to occur primarily in the upper soil layers because the repeated combustion of above‐ground biomass limits organic matter inputs into surface soil. However, C losses from deeper soil may occur if frequent burning reduces root biomass inputs of C into deep soil layers or stimulates losses of C via leaching and priming. To assess the effects of fire on soil C, we sampled 12 plots in a 51‐year‐long fire frequency manipulation experiment in a temperate oak savanna, where variation in prescribed burning frequency has created a gradient in vegetation structure from closed‐canopy forest in unburned plots to open‐canopy savanna in frequently burned plots. Soil C stocks were nonlinearly related to fire frequency, with soil C peaking in savanna plots burned at an intermediate fire frequency and declining in the most frequently burned plots. Losses from deep soil pools were significant, with the absolute difference between intermediately burned plots versus most frequently burned plots more than doubling when the full 1 m sample was considered rather than the top 0–20 cm alone (losses of 98.5 Mg C/ha [−76%] and 42.3 Mg C/ha [−68%] in the full 1 m and 0–20 cm layers respectively). Compared to unburned forested plots, the most frequently burned plots had 65.8 Mg C/ha (−58%) less C in the full 1 m sample. Root biomass below the top 20 cm also declined by 39% with more frequent burning. Concurrent fire‐driven losses of nitrogen and gains in calcium and phosphorus suggest that burning may increase nitrogen limitation and play a key role in the calcium and phosphorus cycles in temperate savannas. Synthesis . Our results illustrate that fire‐driven losses in soil C and root biomass in deep soil layers may be critical factors regulating the net effect of shifting fire regimes on ecosystem C in forest‐savanna transitions. Projected changes in soil C with shifting fire frequencies in savannas may be 50% too low if they only consider changes in the topsoil.

Minnesota↗

Trait integration and functional differentiation among co-existing plant species

Premise Determining which traits characterize strategies of coexisting species is important to developing trait‐based models of plant communities. First, global dimensions may not exist locally. Second, the degree to which traits and trait spectra constitute independent dimensions of functional variation at various scales continues to be refined. Finally, traits may be associated with existing categorical groupings. Methods We assessed trait integration and differentiation across 57 forest understory plant species in Douglas‐fir forests of western Oregon, United States. We combined measurements for a range of traits with literature‐based estimates of seed mass and species groupings. We used network analysis and nonmetric multidimensional scaling ordination (NMS) to determine the degree of integration. Results We observed a strong leaf economics spectrum (LES) integrated with stem but not root traits. However, stem traits and intrinsic water‐use efficiency integrated LES and root traits. Network analyses indicated a modest grouping of a priori trait dimensions. NMS indicated that multivariate differences among species were related primarily to (1) rooting depth and plant height vs. specific root length, (2) the LES, and (3) leaf size vs. seed mass. These differences were related to species groupings associated with growth and life form, leaf lifespan and seed dispersal mechanisms. Conclusions The strategies of coexisting understory plant species could not be reduced to a single dimension. Yet, species can be characterized efficiently and effectively for trait‐based studies of plant communities by measuring four common traits: plant height, specific leaf area, leaf size, and seed mass.

Oregon↗

Riparian soil nitrogen cycling and isotopic enrichment in response to a long-term salmon carcass manipulation experiment

Pacific salmon acquire most of their biomass in the ocean before returning to spawn and die in coastal streams and lakes, thus providing subsidies of marine‐derived nitrogen (MDN) to freshwater and terrestrial ecosystems. Recent declines in salmon abundance have raised questions of whether managers should mitigate for losses of salmon MDN subsidies. To test the long‐term importance of salmon subsidies to riparian ecosystems, we measured soil nitrogen cycling in response to a 20‐yr manipulation where salmon carcasses were systematically removed from one bank and deposited on the opposite bank along a 2‐km stream in southwestern Alaska. Soil samples were taken at different distances from the stream bank along nine paired transects and measured for organic and inorganic nitrogen concentrations, and nitrogen transformation rates. Marine‐derived nitrogen was measured using 15 N/ 14 N for bulk soils, and NH 4 + "> 𝑁𝐻 + 4 NH4+ and NO 3 − "> 𝑁𝑂 − 3 NO3− soil pools. Stable isotope analyses confirmed 15 N/ 14 N was elevated on the salmon‐enhanced bank compared to the salmon‐depleted bank. However, 15 N/ 14 N values of plant‐available inorganic nitrogen exceeded the 15 N/ 14 N of salmon inputs, highlighting nitrogen isotope fractionation in soils that raises significant methodological issues with standard MDN assessments in riparian systems. Surprisingly, despite 20 yr of salmon supplementation, the presence of MDN did not cause a long‐term increase in soil nitrogen availability. This finding indicates the importance of MDN to ecosystem nitrogen biogeochemistry, and riparian vegetation may be overestimated for some systems. Given that essential nutrients can also be pollutants, we urge more critical analyses of the role of MDN to inform compensatory mitigation programs targeting salmon nutrient enhancement.

Ecosphere↗

Growth and survival relationships of 71 tree species with nitrogen and sulfur deposition across the conterminous U.S.

Atmospheric deposition of nitrogen (N) influences forest demographics and carbon (C) uptake through multiple mechanisms that vary among tree species. Prior studies have estimated the effects of atmospheric N deposition on temperate forests by leveraging forest inventory measurements across regional gradients in deposition. However, in the United States (U.S.), these previous studies were limited in the number of species and the spatial scale of analysis, and did not include sulfur (S) deposition as a potential covariate. Here, we present a comprehensive analysis of how tree growth and survival for 71 species vary with N and S deposition across the conterminous U.S. Our analysis of 1,423,455 trees from forest plots inventoried between 2000 and 2016 reveals that the growth and/or survival of the vast majority of species in the analysis (n = 66, or 93%) were significantly affected by atmospheric deposition. Species co-occurred across the conterminous U.S. that had decreasing and increasing relationships between growth (or survival) and N deposition, with just over half of species responding negatively in either growth or survival to increased N deposition somewhere in their range (42 out of 71). Averaged across species and conterminous U.S., however, we found that an increase in deposition above current rates of N deposition would coincide with a small net increase in tree growth (1.7% per Δ kg N ha -1 yr -1 ), and a small net decrease in tree survival (-0.22% per Δ kg N ha -1 yr -1 ), with substantial regional and among-species variation. Adding S as a predictor improved the overall model performance for 70% of the species in the analysis. Our findings have potential to help inform ecosystem management and air pollution policy across the conterminous U.S., and suggest that N and S deposition have likely altered forest demographics in the U.S.

PLoS ONE↗

Molybdenum isotope fractionation during adsorption to organic matter

Organic matter is of emerging interest as a control on molybdenum (Mo) biogeochemistry, and information on isotope fractionation during adsorption to organic matter can improve interpretations of Mo isotope variations in natural settings. Molybdenum isotope fractionation was investigated during adsorption onto insolubilized humic acid (IHA), a surrogate for organic matter, as a function of time (2–170 h) and pH (2–7). For the time series experiment performed at pH 4.2, the average Mo isotope fractionation between the solution and the IHA (Δ 98 Mo solution-IHA ) was 1.39‰ (± 0.16‰, 2σ, based on 98 Mo/ 95 Mo relative to the NIST 3134 standard) at steady state. For the pH series experiment, Mo adsorption decreased as pH increased from 2.0 to 6.9, and the Δ 98 Mo solution-IHA increased from 0.82‰ to 1.79‰. We also evaluated natural Mo isotope patterns in precipitation, foliage, organic horizon, surface mineral soil, and bedrock from 12 forested sites in the Oregon Coast Range. The average Mo isotope offset observed between precipitation and organic (O) horizon soil was 2.1‰, with light Mo isotopes adsorbing preferentially to organic matter. Fractionation during adsorption to organic matter is similar in magnitude and direction to prior observations of Mo fractionation during adsorption to Fe- and Mn- (oxyhydr)oxides. Our finding that organic matter influences Mo isotope composition has important implications for the role of organic matter as a driver of trace metal retention and isotopic fractionation.

Geochimica et Cosmochimica Acta↗

Centennial-scale reductions in nitrogen availability in temperate forests of the United States

Forests cover 30% of the terrestrial Earth surface and are a major component of the global carbon (C) cycle. Humans have doubled the amount of global reactive nitrogen (N), increasing deposition of N onto forests worldwide. However, other global changes—especially climate change and elevated atmospheric carbon dioxide concentrations—are increasing demand for N, the element limiting primary productivity in temperate forests, which could be reducing N availability. To determine the long-term, integrated effects of global changes on forest N cycling, we measured stable N isotopes in wood, a proxy for N supply relative to demand, on large spatial and temporal scales across the continental U.S.A. Here, we show that forest N availability has generally declined across much of the U.S. since at least 1850 C.E. with cool, wet forests demonstrating the greatest declines. Across sites, recent trajectories of N availability were independent of recent atmospheric N deposition rates, implying a minor role for modern N deposition on the trajectory of N status of North American forests. Our results demonstrate that current trends of global changes are likely to be consistent with forest oligotrophication into the foreseeable future, further constraining forest C fixation and potentially storage.

Scientific Reports↗

Intraspecific variability and reaction norms of forest understory plant species traits

Trait-based models of ecological communities typically assume intraspecific variation in functional traits is not important, though such variation can change species trait rankings along gradients in resources and environmental conditions, and thus influence community structure and function. We examined the degree of intraspecific relative to interspecific variation, and reaction norms of 11 functional traits for 57 forest understory plant species, including: intrinsic water-use efficiency (iWUE), Δ 15 N, 5 leaf traits, 2 stem traits and 2 root traits along gradients in light, nitrogen, moisture and understory cover. Our results indicate that interspecific trait variation exceeded intraspecific variation by at least 50% for most, but not all traits. Intraspecific variation in Δ 15 N, iWUE, leaf nitrogen content and root traits was high (47-70%) compared with most leaf traits and stem traits (13-38%). Δ 15 N varied primarily along gradients in abiotic conditions, while light and understory cover were relatively less important. iWUE was related primarily to light transmission, reflecting increases in photosynthesis relative to stomatal conductance. Leaf traits varied mainly as a function of light availability, with some reaction norms depending on understory cover. Plant height increased with understory cover, while stem specific density was related primarily to light. Resources, environmental conditions and understory cover did not contribute strongly to the observed variation in root traits. Gradients in resources, environmental conditions and competition all appear to control intraspecific variability in most traits to some extent. However, our results suggest that species cross-over (i.e., trait rank reversals) along the gradients measured here are generally not a concern. Intraspecific variability in understory plant species traits can be considerable. However, trait data collected under a narrow range of environmental conditions appears sufficient to establish species rankings and scale between community and ecosystem levels using trait-based models. Investigators may therefore focus on obtaining a sufficient sample size within a single set of conditions rather than characterizing trait variation across entire gradients in order to optimize sampling efforts.

Oregon↗

Nutrient feedbacks to soil heterotrophic nitrogen fixation in forests

Multiple nutrient cycles regulate biological nitrogen (N) fixation in forests, yet long-term feedbacks between N-fixation and coupled element cycles remain largely unexplored. We examined soil nutrients and heterotrophic N-fixation across a gradient of 24 temperate conifer forests shaped by legacies of symbiotic N-fixing trees. We observed positive relationships among mineral soil pools of N, carbon (C), organic molybdenum (Mo), and organic phosphorus (P) across sites, evidence that legacies of symbiotic N-fixing trees can increase the abundance of multiple elements important to heterotrophic N-fixation. Soil N accumulation lowered rates of heterotrophic N-fixation in organic horizons due to both N inhibition of nitrogenase enzymes and declines in soil organic matter quality. Experimental fertilization of organic horizon soil revealed widespread Mo limitation of heterotrophic N-fixation, especially at sites where soil Mo was scarce relative to C. Fertilization also revealed widespread absence of P limitation, consistent with high soil P:Mo ratios. Responses of heterotrophic N-fixation to added Mo (positive) and N (negative) were correlated across sites, evidence that multiple nutrient controls of heterotrophic N-fixation were more common than single-nutrient effects. We propose a conceptual model where symbiotic N-fixation promotes coupled N, C, P, and Mo accumulation in soil, leading to positive feedback that relaxes nutrient limitation of overall N-fixation, though heterotrophic N-fixation is primarily suppressed by strong negative feedback from long-term soil N accumulation.

Oregon Coast Range↗

Mechanisms of nitrogen deposition effects on temperate forest lichens and trees

We review the mechanisms of deleterious nitrogen (N) deposition impacts on temperate forests, with a particular focus on trees and lichens. Elevated anthropogenic N deposition to forests has varied effects on individual organisms depending on characteristics both of the N inputs (form, timing, amount) and of the organisms (ecology, physiology) involved. Improved mechanistic knowledge of these effects can aid in developing robust predictions of how organisms respond to either increases or decreases in N deposition. Rising N levels affect forests in micro- and macroscopic ways from physiological responses at the cellular, tissue, and organism levels to influencing individual species and entire communities and ecosystems. A synthesis of these processes forms the basis for the overarching themes of this paper, which focuses on N effects at different levels of biological organization in temperate forests. For lichens, the mechanisms of direct effects of N are relatively well known at cellular, organismal, and community levels, though interactions of N with other stressors merit further research. For trees, effects of N deposition are better understood for N as an acidifying agent than as a nutrient; in both cases, the impacts can reflect direct effects on short time scales and indirect effects mediated through long-term soil and belowground changes. There are many gaps on fundamental N use and cycling in ecosystems, and we highlight the most critical gaps for understanding potential deleterious effects of N deposition. For lichens, these gaps include both how N affects specific metabolic pathways and how N is metabolized. For trees, these gaps include understanding the direct effects of N deposition onto forest canopies, the sensitivity of different tree species and mycorrhizal symbionts to N, the influence of soil properties, and the reversibility of N and acidification effects on plants and soils. Continued study of how these N response mechanisms interact with one another, and with other dimensions of global change, remains essential for predicting ongoing changes in lichen and tree populations across North American temperate forests.

Ecosphere↗

Riparian soil development linked to forest succession above and below dams along the Elwha River, Washington, USA

Riparian forest soils can be highly dynamic, due to frequent fluvial disturbance, erosion, and sediment deposition, but effects of dams on riparian soils are poorly understood. We examined soils along toposequences within three river segments located upstream, between, and downstream of two dams on the Elwha River to evaluate relationships between riparian soil development and forest age, succession, and channel proximity, explore dam effects on riparian soils, and provide a baseline for the largest dam removal in history. We found that older, later-successional forests and geomorphic surfaces contained soils with finer texture and greater depth to cobble, supporting greater forest floor mass, mineral soil nutrient levels, and cation exchange. Forest stand age was a better predictor than channel proximity for many soil characteristics, though elevation and distance from the channel were often also important, highlighting how complex interactions between fluvial disturbance, sediment deposition, and biotic retention regulate soil development in this ecosystem. Soils between the dams, and to a lesser extent below the lower dam, had finer textures and higher mineral soil carbon, nitrogen, and cation exchange than above the dams. These results suggested that decreased fluvial disturbance below the dams, due to reduced sediment supply and channel stabilization, accelerated soil development. In addition, reduced sediment supply below the dams may have decreased soil phosphorus. Soil δ15N suggested that salmon exclusion by the dams had no discernable effect on nitrogen inputs to upstream soils. Recent dam removal may alter riparian soils further, with ongoing implications for riparian ecosystems.

Washington↗

Combined use of isotopic and hydrometric data to conceptualize ecohydrological processes in a high-elevation tropical ecosystem

Few high-elevation tropical catchments worldwide are gauged and even fewer are studied using combined hydrometric and isotopic data. Consequently, we lack information needed to understand processes governing rainfall-runoff dynamics and to predict their influence on downstream ecosystem functioning. To address this need, we present a combination of hydrometric and water stable isotopic observations in the wet Andean p&aacute;ramo ecosystem of the Zhurucay Ecohydrological Observatory (7.53 km2). The catchment is located in the Andes of south Ecuador between 3400 and 3900 m a.s.l. Water samples for stable isotopic analysis were collected during 2 years (May 2011 &ndash; May 2013), while rainfall and runoff measurements were continuously recorded since late 2010. The isotopic data reveal that Andosol soils predominantly situated on hillslopes drain laterally to Histosols (Andean p&aacute;ramo wetlands) mainly located at the valley bottom. Histosols, in turn, feed water to creeks and small rivers throughout the year, establishing hydrologic connectivity between wetlands and the drainage network. Runoff is primarily comprised of pre-event water stored in the Histosols, which is replenished by rainfall that infiltrates through the Andosols. Contributions from the mineral horizon and the top of the fractured bedrock are small and only seem to influence discharge in small catchments during low flow generation (non-exceedance flows < Q35). Variations in source contributions are controlled by antecedent soil moisture, rainfall intensity, and duration of rainy periods. Saturated hydraulic conductivity of the soils, higher than the year-round low precipitation intensity, indicates that Hortonian overland flow rarely occurs during high intensity precipitation events. Deep groundwater contributions to discharge seem to be minimal. These results suggest that, in this high-elevation tropical ecosystem: 1) subsurface flow is a dominant hydrological process and 2) (Histosols) wetlands are the major source of stream runoff. Our study highlights that detailed isotopic characterization during short time periods provides valuable information about ecohydrological processes in regions where very few basins are gauged.

Zhurucay River Ecohydrological Observatory↗

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.

Global Change Biology↗