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1. Data from: Rainfall events stimulate episodic associative nitrogen fixation in switchgrass

   https://doi.org/10.5061/dryad.s7h44j1h3  

   Vizza, Carmella; Belanger, Matthew; Jones, Jabarius; Murphy, Sean; Buckley, Daniel; Robertson, G Philip; Roley, Sarah (January 2025, Dryad)

   Associative N2 fixation (ANF) is widespread but poorly characterized, limiting our ability to estimate global inputs from N2 fixation. In some places, ANF rates are at or below detection most of the time, but occasionally and unpredictably spiking to very high rates. Here we test the hypothesis that plant phenology and rainfall events stimulate ANF episodes. We measured ANF in intact soil cores in switchgrass (Panicum virgatum L.) in Michigan, USA. We used rain exclusion shelters to impose three rainfall treatments with each receiving 60 mm of water over a 20-day period but at different frequencies. We concurrently established a treatment that received ambient rainfall, and all four treatments were replicated four times. To assess the effects of plant phenology, we measured ANF at key phenological stages in the ambient treatment. To assess the effects of rainfall, we measured ANF immediately before and immediately after each wetting event in each treatment involving rainfall manipulation. We found that the previous day’s rainfall could explain 29% of the variation in ANF rates within the ambient treatment alone, and that bulk soil C:N ratio was also positively correlated with ANF, explaining 18% of the variation alone. Wetting events increased ANF and the magnitude of response to wetting increased with the amount of water added and decreased with the amount of inorganic N added in water. ANF episodes thus appear to be driven primarily by wetting events. Wetting events likely increase C availability, promote microbial growth, and make rhizosphere conditions conducive to ANF. 

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2. Evidence for the Coupling of Plant Functional Diversity and Soil Biogeochemistry Under Climatic Stress in Pacific Northwest Grasslands

   https://doi.org/10.1029/2024JG008126  

   Bomfim, Barbara; Dawson, Hilary_R; Reed, Paul_B; Shek, Katherine_L; Bohannan, Brendan_J_M; Bridgham, Scott_D; Silva, Lucas_C_R (May 2025, Journal of Geophysical Research: Biogeosciences)

   Abstract Increasing warming and drought severity are projected for the Pacific Northwest (PNW) and are expected to negatively impact species composition and ecosystem function. In this study, we test the hypothesis that the impact of climatic stress (i.e., experimental warming and drought) on PNW grasslands are mediated by interactions between plant functional diversity and soil biogeochemical processes, including symbiotic nitrogen (N) fixation in legumes and free‐living asymbiotic nitrogen fixation (ANF) by soil microorganisms. To test this hypothesis, we measured the response of plants and soils to three years of warming (+2.5°C) and drought (−40% precipitation) in field experiments replicated at three different sites across a ∼520‐km latitudinal gradient. We observed interactive effects of warming and drought on functional diversity and soil biogeochemical properties, including both positive and negative changes in ANF. Although direct measurements of symbiotic nitrogen fixation (SNF) rates were not conducted, the observed variations in ANF, in conjunction with changes in legume cover, suggest a compensatory mechanism that may offset reductions in SNF. Generally, high ANF rates coincided with low legume cover, suggesting a connection between shifts in species composition and N cycling. Our ANF estimates were performed using isotopically labeled dinitrogen (¹⁵N₂) in tandem with soil carbon (C), phosphorus (P) and iron (Fe), pH, and moisture content. Along the latitudinal drought severity gradient, ANF rates were correlated with changes in species composition and soil N, P, moisture, and pH levels. These results highlight the importance of soil‐plant‐atmosphere interactions in understanding the impacts of climatic stress on ecosystem composition and function. 

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3. Effects of Altered Precipitation on Biological Soil Crusts, Fungi, and Soil Nitrogen Availability at the Monsoon Rainfall Manipulation Experiment (MRME) in the Sevilleta National Wildlife Refuge, New Mexico (2016)

   https://doi.org/10.6073/pasta/573848e6e65f686686884670de506465  

   Kwiecinski, Jarek V (January 2019, Environmental Data Initiative)

   Microbial activity in drylands is mediated by the magnitude and frequency of growing season rain events that will shift as climate change progresses. Nitrogen is often co-limiting with water availability to dryland plants, and thus we investigated how microbes important to the nitrogen (N) cycle and soil N availability varied temporally and spatially in the context of a long-term rainfall variability experiment in the northern Chihuahuan Desert. Specifically, we assessed biological soil crust (biocrust) chlorophyll content, fungal abundance, and inorganic N in soils adjacent to individuals of the grassland foundation species, Bouteloua eriopoda, and in the unvegetated interspace at multiple time points associated with an experimental monsoon rain treatment. Treatments included small weekly (5 mm) or large monthly (20 mm) rain events, which had been applied during the summer monsoon for nine years prior to our sampling. Additionally, we evaluated target plant C:N ratios and added 15 N-glutamate to biocrusts to determine potential for nutrient transport to B. eriopoda. Biocrust chlorophyll was up to 67% higher in the small weekly or large monthly rainfall regimes compared to ambient controls. Fungal biomass was 57% lower in soil interspaces than adjacent to plants but did not respond to rainfall regime treatments. Ammonium and nitrate concentrations near plants declined through the sampling period but varied little in soil interspaces. There was limited movement of 15 N from interspace biocrusts to leaves but high 15 N retention in the soils even after additional ambient and experimental rain events. Plant C:N ratio was unaffected by rainfall treatments. The long-term alteration in rainfall regime in this experiment did not change how short-term microbial abundance or N availability responded to the magnitude or frequency of events, suggesting a limited response of N availability to future climate change. 

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4. Belowground responses to altered precipitation regimes in two semi-arid grasslands

   https://doi.org/10.6073/pasta/2d233f4ba9af3a13b564a1c6364704c7  

   Holguin, Jennifer; Collins, Scott L; McLaren, Jennie R (January 2022, Environmental Data Initiative)

   Predicted climate change extremes, such as severe and prolonged drought, may profoundly impact biogeochemical processes like carbon and nitrogen cycling in water-limited ecosystems. To increase our understanding of how extreme climate events impact belowground ecosystem processes, we investigated the effects of five years of severe growing season drought and two-month delay in monsoon precipitation on belowground productivity and biogeochemical processes in two semi-arid grasslands. This experiment takes place during the fifth year of the Extreme Drought in Grassland Experiment (EDGE) at the Sevilleta National Wildlife Refuge (SNWR), a Long-Term Ecological Research in central New Mexico, USA. The two grassland sites a Chihuahuan Desert grassland dominated by Bouteloua eriopoda and Great Plains grassland dominated by B. gracilis are ~5km apart in the SWNR. The EDGE platform was established in the spring of 2012 (pre-treatment). Each site contains three treatments (ten replicates): ambient rainfall, extreme growing season drought, and delayed monsoon. The extreme drought treatment reduces growing season rainfall (April through September) each year by 66%, which equates to a 50% reduction of annual precipitation while maintaining natural precipitation patterns. There are 10 replicates per treatment within each site. All plots are 3 x 4 m in size and are paired spatially into blocks with treatments assigned randomly within a block. We measured an array of belowground and biogeochemical variables. Each variable was measured either once, twice, or three times (specific information on sampling scheme for each measured variable in methods section). Belowground net primary productivity, standing crop root biomass, total organic carbon, and total nitrogen were measured once. Extractable organic carbon, extractable total nitrogen, microbial biomass carbon, microbial biomass nitrogen and extracellular enzymes were measured twice. Available soil nitrate, available soil ammonium, and available soil phosphate were measured three times. 

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5. Deconstructing precipitation variability: Rainfall event size and timing uniquely alter ecosystem dynamics

   https://doi.org/10.1111/1365-2745.13724  

   Griffin‐Nolan, Robert J.; Slette, Ingrid J.; Knapp, Alan K. (July 2021, Journal of Ecology)

   Abstract Water‐limited ecosystems are highly sensitive to not only precipitation amount, but also precipitation pattern, particularly variability in the size and timing of growing season rainfall events. Both rainfall event size and timing are expected to be altered by climate change, but the relative responses of dryland ecosystems to changes in rainfall event size versus timing have not been resolved. Here, we disentangle the effects of these different aspects of precipitation pattern on ecosystem dynamics.We experimentally assessed how these two aspects of rainfall variability impacted a semi‐arid grassland ecosystem by altering an ambient precipitation pattern to eliminate variability in (a) rainfall event size (all events were made the same size), (b) rainfall event timing (all events were uniformly spaced in time) and (c) both. Total precipitation amount was constant for all treatments. We measured responses of soil moisture, ecosystem carbon flux (e.g. net primary production and soil CO₂flux), plant community composition and physiological responses of the dominant C₄grass,Bouteloua gracilis.Removing variability in rainfall event size altered ecosystem dynamics more than a pattern of uniform event timing, but the largest impact occurred when variability in both were removed. Notably, eliminating variability in both event size and timing increased above‐ground net primary productivity by 23%, consistent with reduced water stress in the dominant C₄grass, while also reducing seasonal variability in soil CO₂flux by 35%, reflecting lower seasonal variability in soil moisture.Synthesis. Unique responses to different aspects of precipitation variability highlight the complexity of predicting how dryland ecosystems will be affected by climate change‐induced shifts in rainfall patterns. Our results provide novel support for the key roles of rainfall event size and timing, in addition to total precipitation amount, as determinants of ecosystem function. 

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