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Sarah E. Jovan

Publications and source records attributed to Sarah E. Jovan.

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Global carbon investment in terrestrial biological nitrogen fixation

Biological nitrogen (N) fixation (BNF) provides the N needed to produce proteins and other biological building blocks, helping feed humanity and mitigate climate change. Due to its high energetic cost compared to other forms of N acquisition, biotic investment in BNF indicates N limitation. Globally gridded BNF flux data provide an opportunity to determine the energetic investment in BNF across ecosystems, which would help reconcile conflicting indicators of N limitation. Here, we use a new BNF synthesis to quantify the relative importance of BNF in different N-fixing niches, in different biomes, and across the globe by calculating the fraction of net primary productivity (NPP) invested in BNF and the fraction of plant N acquisition provided by BNF. Larger fractions of non-agricultural NPP were invested in BNF in less-productive, higher-latitude biomes. This pattern was driven by biocrusts and mosses. Similarly, non-agricultural symbiotic N-fixing plants invested relatively more of their own NPP in BNF in less-productive, higher-latitude biomes. This symbiotic plant pattern was driven by shrubs and herbs, overriding the opposite pattern in trees. Symbiotic plants also acquired a higher fraction of their N from BNF at higher latitudes and in less productive biomes. Investments in symbiotic BNF were 10× higher in agricultural (2.9% of NPP) versus natural (0.29%) biomes, providing 26% versus 3.1% of ecosystem-scale plant N acquisition. These results support the paradigm of strong N limitation at higher latitudes, help understand the rarity of N-fixing trees at higher latitudes, underscore the dominance of human activity, and inform terrestrial biosphere models.

Global Biogeochemical Cycles

Overestimated natural biological nitrogen fixation translates to an exaggerated CO2 fertilization effect in Earth System Models

CO 2 fertilization of the terrestrial biosphere is limited by nitrogen. Biological nitrogen fixation (BNF) is the dominant natural nitrogen source to the terrestrial biosphere and can alleviate nitrogen limitation but is poorly constrained in Earth system models (ESMs). Here, we compare terrestrial BNF from an ensemble of ESMs of the 6th Coupled Model Intercomparison Project to a new global synthesis of observations across natural and agricultural biomes. We find that compared to observations, ESMs underestimate agricultural BNF but overestimate natural BNF in the present day by over 50%. Natural BNF is overestimated in the most productive ecosystems that contribute most to the terrestrial carbon sink (forests and grasslands). ESMs with different BNF representations yield a range of BNF responses to CO 2 enrichment. Some ESMs with phenomenological representations of BNF predict a natural BNF increase in response to a doubling of CO 2 that aligns with a meta-analysis of CO 2 enrichment experiments (31% increase) but fail to account for the substantial carbon cost of BNF. In contrast, ESMs with mechanistic representations of BNF account for its carbon cost as well as its regulation by nitrogen limitation but overestimate the BNF response to a doubling of CO 2 (135% increase). Overall, all current BNF representations in ESMs fall short of fully capturing its response to rising atmospheric CO 2 . Finally, we find a positive correlation between modeled present-day natural BNF and the CO 2 fertilization effect across ESMs, suggesting that overestimated natural BNF translates to an exaggerated CO 2 fertilization effect of approximately 11% in ESMs.

PNAS

A global dataset of terrestrial biological nitrogen fixation

Biological nitrogen fixation (BNF) is the main natural source of new nitrogen inputs in terrestrial ecosystems, supporting terrestrial productivity, carbon uptake, and other Earth system processes. We assembled a comprehensive global dataset of field measurements of BNF in all major N-fixing niches across natural terrestrial biomes derived from the analysis of 376 BNF studies. The dataset comprises 32 variables, including site location, biome type, N-fixing niche, sampling year, quantification method, BNF rate (kg N ha −1 y −1 ), the percentage of nitrogen derived from the atmosphere (%N dfa ), N fixer or N-fixing substrate abundance, BNF rate per unit of N fixer abundance, and species identity. Overall, the dataset combines 1,207 BNF rates for trees, shrubs, herbs, soil, leaf litter, woody litter, dead wood, mosses, lichens, and biocrusts, 152 herb %N dfa values, 1,005 measurements of N fixer or N-fixing substrate abundance, and 762 BNF rates per unit of N fixer abundance for a total of 424 species across 66 countries. This dataset facilitates synthesis, meta-analysis, upscaling, and model benchmarking of BNF fluxes at multiple spatial scales.

Scientific Data

Global terrestrial nitrogen fixation and its modification by agriculture

Biological nitrogen fixation (BNF) is the largest natural source of new nitrogen (N) that supports terrestrial productivity 1,2 , yet estimates of global terrestrial BNF remain highly uncertain 3,4 . Here we show that this uncertainty is partly because of sampling bias, as field BNF measurements in natural terrestrial ecosystems occur where N fixers are 17 times more prevalent than their mean abundances worldwide. To correct this bias, we develop new estimates of global terrestrial BNF by upscaling field BNF measurements using spatially explicit abundances of all major biogeochemical N-fixing niches. We find that natural biomes sustain lower BNF, 65 (52–77) Tg N yr −1 , than previous empirical bottom-up estimates 3,4 , with most BNF occurring in tropical forests and drylands. We also find high agricultural BNF in croplands and cultivated pastures, 56 (54–58) Tg N yr −1 . Agricultural BNF has increased terrestrial BNF by 64% and total terrestrial N inputs from all sources by 60% over pre-industrial levels. Our results indicate that BNF may impose stronger constraints on the carbon sink in natural terrestrial biomes and represent a larger source of agricultural N than is generally considered in analyses of the global N cycle 5,6 , with implications for proposed safe operating limits for N use 7,8 .

Nature

Potential vulnerability of 348 herbaceous species to atmospheric deposition of nitrogen and sulfur in the United States

Atmospheric nitrogen and sulfur pollution increased over much of the United States during the twentieth century from fossil fuel combustion and industrial agriculture. Despite recent declines, nitrogen and sulfur deposition continue to affect many plant communities in the United States, although which species are at risk remains uncertain. We used species composition data from >14,000 survey sites across the contiguous United States to evaluate the association between nitrogen and sulfur deposition and the probability of occurrence for 348 herbaceous species. We found that the probability of occurrence for 70% of species was negatively associated with nitrogen or sulfur deposition somewhere in the contiguous United States (56% for N, 51% for S). Of the species, 15% and 51% potentially decreased at all nitrogen and sulfur deposition rates, respectively, suggesting thresholds below the minimum deposition they receive. Although more species potentially increased than decreased with nitrogen deposition, increasers tended to be introduced and decreasers tended to be higher-value native species. More vulnerable species tended to be shorter with lower tissue nitrogen and magnesium. These relationships constitute predictive equations to estimate critical loads. These results demonstrate that many herbaceous species may be at risk from atmospheric deposition and can inform improvements to air quality policies in the United States and globally.

Contiguous United States

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

Conditional vulnerability of plant diversity to atmospheric nitrogen deposition across the United States

Atmospheric nitrogen (N) deposition has been shown to decrease plant species richness along regional deposition gradients in Europe and in experimental manipulations. However, the general response of species richness to N deposition across different vegetation types, soil conditions, and climates remains largely unknown even though responses may be contingent on these environmental factors. We assessed the effect of N deposition on herbaceous richness for 15,136 forest, woodland, shrubland, and grassland sites across the continental United States, to address how edaphic and climatic conditions altered vulnerability to this stressor. In our dataset, with N deposition ranging from 1 to 19 kg N⋅ha−1⋅y−1, we found a unimodal relationship; richness increased at low deposition levels and decreased above 8.7 and 13.4 kg N⋅ha−1⋅y−1 in open and closed-canopy vegetation, respectively. N deposition exceeded critical loads for loss of plant species richness in 24% of 15,136 sites examined nationwide. There were negative relationships between species richness and N deposition in 36% of 44 community gradients. Vulnerability to N deposition was consistently higher in more acidic soils whereas the moderating roles of temperature and precipitation varied across scales. We demonstrate here that negative relationships between N deposition and species richness are common, albeit not universal, and that fine-scale processes can moderate vegetation responses to N deposition. Our results highlight the importance of contingent factors when estimating ecosystem vulnerability to N deposition and suggest that N deposition is affecting species richness in forested and nonforested systems across much of the continental United States.

PNAS

Indicators of climate impacts for forests: Recommendations for the U.S. National Climate Assessment Indicators system

The Third National Climate Assessment (NCA) process for the United States focused in part on developing a system of indicators to communicate key aspects of the physical climate, climate impacts, vulnerabilities, and preparedness to inform decisionmakers and the public. Initially, 13 active teams were formed to recommend indicators in a range of categories, including forest, agriculture, grassland, phenology, mitigation, and physical climate. This publication describes the work of the Forest Indicators Technical Team. We briefly describe the NCA indicator system effort, propose and explain our conceptual model for the forest system, present our methods, and discuss our recommendations. Climate is only one driver of changes in U.S. forests; other drivers include socioeconomic drivers such as population and culture, and other environmental drivers such as nutrients, light, and disturbance. We offer additional details of our work for transparency and to inform an NCA indicator Web portal. We recommend metrics for 11 indicators of climate impacts on forest, spanning the range of important aspects of forest as an ecological type and as a sector. Some indicators can be reported in a Web portal now; others need additional work for reporting in the near future. Indicators such as budburst, which are important to forest but more relevant to other NCA indicator teams, are identified. Potential indicators that need more research are also presented.

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