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Long-term reactive nitrogen loading alters soil carbon and microbial community properties in a subalpine forest ecosystem

Elevated nitrogen (N) deposition due to increased fossil fuel combustion and agricultural practices has altered global carbon (C) cycling. Additions of reactive N to N-limited environments are typically accompanied by increases in plant biomass. Soil C dynamics, however, have shown a range of different responses to the addition of reactive N that seem to be ecosystem dependent. We evaluated the effect of N amendments on biogeochemical characteristics and microbial responses of subalpine forest organic soils in order to develop a mechanistic understanding of how soils are affected by N amendments in subalpine ecosystems. We measured a suite of responses across three years (2011–2013) during two seasons (spring and fall). Following 17 years of N amendments, fertilized soils were more acidic (control mean 5.09, fertilized mean 4.68), and had lower %C (control mean 33.7% C, fertilized mean 29.8% C) and microbial biomass C by 22% relative to control plots. Shifts in biogeochemical properties in fertilized plots were associated with an altered microbial community driven by reduced arbuscular mycorrhizal (control mean 3.2 mol%, fertilized mean 2.5 mol%) and saprotrophic fungal groups (control mean 17.0 mol%, fertilized mean 15.2 mol%), as well as a decrease in N degrading microbial enzyme activity. Our results suggest that decreases in soil C in subalpine forests were in part driven by increased microbial degradation of soil organic matter and reduced inputs to soil organic matter in the form of microbial biomass.

Soil Biology and Biochemistry↗

Ammonia and aquatic ecosystems – A review of global sources, biogeochemical cycling, and effects on fish

The purpose of this review is to better understand the full life cycle and influence of ammonia from an aquatic biology perspective. While ammonia has toxic properties in water and air, it also plays a central role in the biogeochemical nitrogen (N) cycle and regulates mechanisms of normal and abnormal fish physiology. Additionally, as the second most synthesized chemical on Earth, ammonia contributes economic value to many sectors, particularly fertilizers, energy storage, explosives, refrigerants, and plastics. But, with so many uses, industrial N 2 -fixation effectively doubles natural reactive N concentrations in the environment. The consequence is global, with excess fixed nitrogen driving degradation of soils, water, and air; intensifying eutrophication, biodiversity loss, and climate change; and creating health risks for humans, wildlife, and fisheries. Thus, the need for ammonia research in aquatic systems is growing. In response, we prepared this review to better understand the complexities and connectedness of environmental ammonia. Even the term “ammonia” has multiple meanings. So, we have clarified the nomenclature, identified units of measurement, and summarized methods to measure ammonia in water. We then discuss ammonia in the context of the N-cycle, review its role in fish physiology and mechanisms of toxicity, and integrate the effects of human N-fixation, which continuously expands ammonia's sources and uses. Ammonia is being developed as a carbon-free energy carrier with potential to increase reactive nitrogen in the environment. With this in mind, we review the global impacts of excess reactive nitrogen and consider the current monitoring and regulatory frameworks for ammonia. The presented synthesis illustrates the complex and interactive dynamics of ammonia as a plant nutrient, energy molecule, feedstock, waste product, contaminant, N-cycle participant, regulator of animal physiology, toxicant, and agent of environmental change. Few molecules are as influential as ammonia in the management and resilience of Earth's resources.

Science of the Total Environment↗

Characterization of water quality and biological communities, Fish Creek, Teton County, Wyoming, 2007-2011

Fish Creek, an approximately 25-kilometer-long tributary to Snake River, is located in Teton County in western Wyoming near the town of Wilson. Fish Creek is an important water body because it is used for irrigation, fishing, and recreation and adds scenic value to the Jackson Hole properties it runs through. Public concern about nuisance growths of aquatic plants in Fish Creek has been increasing since the early 2000s. To address these concerns, the U.S. Geological Survey conducted a study in cooperation with the Teton Conservation District to characterize the hydrology, water quality, and biologic communities of Fish Creek during 2007–11. The hydrology of Fish Creek is strongly affected by groundwater contributions from the area known as the Snake River west bank, which lies east of Fish Creek and west of Snake River. Because of this continuous groundwater discharge to the creek, land-use activities in the west bank area can affect the groundwater quality. Evaluation of nitrate isotopes and dissolved-nitrate concentrations in groundwater during the study indicated that nitrate was entering Fish Creek from groundwater, and that the source of nitrate was commonly a septic/sewage effluent or manure source, or multiple sources, potentially including artificial nitrogen fertilizers, natural soil organic matter, and mixtures of sources. Concentrations of dissolved nitrate and orthophosphate, which are key nutrients for growth of aquatic plants, generally were low in Fish Creek and occasionally were less than reporting levels (not detected). One potential reason for the low nutrient concentrations is that nutrients were being consumed by aquatic plant life that increases during the summer growing season, as a result of the seasonal increase in temperature and larger number of daylight hours. Several aspects of Fish Creek’s hydrology contribute to higher productivity and biovolume of aquatic plants in Fish Creek than typically observed in streams of its size in Wyoming. Especially in the winter, the proportionately large, continuous gain of groundwater into Fish Creek in the perennial section keeps most of the creek free of ice. Because sunlight can still reach the streambed in Fish Creek and the water is still flowing, aquatic plants continue to photosynthesize in the winter, albeit at a lower level of productivity. Additionally, the cobble and large gravel substrate in Fish Creek provides excellent attachment points for aquatic plants, and when combined with Fish Creek’s channel stability allows rapid growth of aquatic plants once conditions allow during the spring. The aquatic plant community of Fish Creek was different than most streams in Wyoming in that it contains many different macrophytes—including macroalgae such as long streamers of Cladophora , aquatic vascular plants, and moss; most other streams in the state contain predominantly algae. From the banks of Fish Creek, the bottom of the stream sometimes appeared to be a solid green carpet. A shift was observed from higher amounts of microalgae in April/May to higher amounts macrophytes in August and October, and differences in the relative abundance of microalgae and macrophytes were statistically significant between seasons. Differences in dissolved-nitrate concentrations and in the nitrogen-to-phosphorus ratio were significantly different between seasons, as concentrations of dissolved nitrate decreased from April/May to August and October. It is likely that dissolved-nitrate concentrations in Fish Creek were lower in August and October because macrophytes were quickly utilizing the nutrient, and a negative correlation between macro-phytes and nitrate was found. Macroinvertebrates also were sampled because of their role as indicators of water quality and their documented responses to perturbation such as degradation of water quality and habitat. Statistically significant seasonal differences were noted in the macroinvertebrate community. Taxa richness and relative abundance of Ephemeroptera, Plecoptera, and Trichoptera, which tend to be intolerant of water-quality degradation, decreased from April/May to August; the same time period saw a corresponding increase in Diptera and noninsects, particularly Oligochaeta (worms) that are more tolerant. Seasonal changes in macroinvertebrate functional feeding groups were significantly different. The relative abundance of gatherer-collector and scraper feeding groups decreased from April/May to August, accompanied by an increase in filterer-collector and shredders feeding groups. Seasonal changes in feeding groups might be due to the seasonal shift in aquatic plant communities, as indicated by comparison with other streams in the area that had fewer aquatic macrophytes than Fish Creek. Statistical tests of macroinvertebrate metrics indicated few differences between years or biological sampling sites on Fish Creek, although the site farthest upstream sometimes was different not only in terms of macroinvertebrates but also in streamflow, water quality, and aquatic plants. Potential effects of contributions of additional nutrients to the Fish Creek ecosystem beyond the conditions sampled during the study period are not known. However, because virtually all of the detectable dissolved nitrate commonly was consumed by aquatic plants in August (leaving dissolved nitrate less than the reporting level in water samples), it is possible that increased nutrient contributions could cause increased growth of aquatic plants. Additional long-term monitoring of the stream, with concurrent data analysis and interpretation would be needed to determine the effects of additional nutrients on the aquatic plant community and on higher levels of the food chain.

Wyoming↗

Hydrologic regime controls soil phosphorus fluxes in restoration and undisturbed wetlands

Many wetland restoration projects occur on former agricultural soils that have a history of disturbance and fertilization, making them prone to phosphorus (P) release upon flooding. To study the relationship between P release and hydrologic regime, we collected soil cores from three restoration wetlands and three undisturbed wetlands around Upper Klamath Lake in southern Oregon, U.S.A. Soil cores were subjected to one of three hydrologic regimes - flooded, moist, and dry - for 7.5 weeks, and P fluxes were measured upon reflooding. Soils from restoration wetlands released P upon reflooding regardless of the hydrologic regime, with the greatest releases coming from soils that had been flooded or dried. Undisturbed wetland soils released P only after drying. Patterns in P release can be explained by a combination of physical and biological processes, including the release of iron-bound P due to anoxia in the flooded treatment and the mineralization of organic P under aerobic conditions in the dry treatment. Higher rates of soil P release from restoration wetland soils, particularly under flooded conditions, were associated with higher total P concentrations compared with undisturbed wetland soils. We conclude that maintaining moist soil is the means to minimize P release from recently flooded wetland soils. Alternatively, prolonged flooding provides a means of liberating excess labile P from former agricultural soils while minimizing continued organic P mineralization and soil subsidence. ?? 2005 Society for Ecological Restoration International.

Restoration Ecology↗

Patterns and controls on nitrogen cycling of biological soil crusts

Biocrusts play a significant role in the nitrogen [N ] cycle within arid and semi-arid ecosystems, as they contribute major N inputs via biological fixation and dust capture, harbor internal N transformation processes, and direct N losses via N dissolved, gaseous and erosional loss processes (Fig. 1). Because soil N availability in arid and semi-arid ecosystems is generally low and may limit net primary production (NPP), especially during periods when adequate water is available, understanding the mechanisms and controls of N input and loss pathways in biocrusts is critically important to our broader understanding of N cycling in dryland environments. In particular, N cycling by biocrusts likely regulates short-term soil N availability to support vascular plant growth, as well as long-term N accumulation and maintenance of soil fertility. In this chapter, we review the influence of biocrust nutrient input, internal cycling, and loss pathways across a range of biomes. We examine linkages between N fixation capabilities of biocrust organisms and spatio-temporal patterns of soil N availability that may influence the longer-term productivity of dryland ecosystems. Lastly, biocrust influence on N loss pathways such as N gas loss, leakage of N compounds from biocrusts, and transfer in wind and water erosion are important to understand the maintenance of dryland soil fertility over longer time scales. Although great strides have been made in understanding the influence of biocrusts on ecosystem N cycling, there are important knowledge gaps in our understanding of the influence of biocrusts on ecosystem N cycling that should be the focus of future studies. Because work on the interaction of N cycling and biocrusts was reviewed in Belnap and Lange (2003), this chapter will focus primarily on research findings that have emerged over the last 15 years (2000-2015).

Book chapter↗

Spatial and temporal variation of stream chemistry associated with contrasting geology and land-use patterns in the Chesapeake Bay watershed—Summary of results from Smith Creek, Virginia; Upper Chester River, Maryland; Conewago Creek, Pennsylvania; and Difficult Run, Virginia, 2010–2013

Despite widespread and ongoing implementation of conservation practices throughout the Chesapeake Bay watershed, water quality continues to be degraded by excess sediment and nutrient inputs. While the Chesapeake Bay Program has developed and maintains a large-scale and long-term monitoring network to detect improvements in water quality throughout the watershed, fewer resources have been allocated for monitoring smaller watersheds, even though water-quality improvements that may result from the implementation of conservation practices are likely to be first detected at smaller watershed scales. In 2010, the U.S. Geological Survey partnered with the U.S. Environmental Protection Agency and the U.S. Department of Agriculture to initiate water-quality monitoring in four selected small watersheds that were targeted for increased implementation of conservation practices. Smith Creek watershed is an agricultural watershed in the Shenandoah Valley of Virginia that is dominated by cattle and poultry production, and the Upper Chester River watershed is an agricultural watershed on the Eastern Shore of Maryland that is dominated by row-cropping activities. The Conewago Creek watershed is an agricultural watershed in southeastern Pennsylvania that is characterized by mixed agricultural activities. The fourth watershed, Difficult Run, is a suburban watershed in northern Virginia that is dominated by medium density residential development. The objective of this study was to investigate spatial and temporal variations in water chemistry and suspended sediment in these four relatively small watersheds that represent a range of land-use patterns and underlying geology to (1) characterize current water-quality conditions in these watersheds, and (2) identify the dominant sources, sinks, and transport processes in each watershed. The general study design involved two components. The first included intensive routine water-quality monitoring at an existing streamgage within each study area (including continuous water-quality monitoring as well as discrete water-quality sampling) to develop a detailed understanding of the temporal and hydrologic variability in stream chemistry and sediment transport in each watershed. The second component involved extensive water-quality monitoring at various sites throughout each watershed to develop a detailed understanding of spatial patterns. Both components were used to improve understanding of sources and transport processes affecting stream chemistry, including nutrients and suspended sediments, and their implications for detecting long-term trends related to best management practices. This report summarizes the results of monitoring that was performed from April 2010 through September 2013. Individual Small Watershed Summaries Summaries for each of the four small watersheds are presented below. Each watershed has a more descriptive and detailed section in the report, but these summaries may be particularly useful for some watershed managers and stakeholders desiring slightly less technical detail. Smith Creek Smith Creek is a 105.39-mi 2 watershed within the Shenandoah Valley that drains to the North Fork Shenandoah River. The long-term Smith Creek base-flow index is 72.3 percent, indicating that on average, approximately 72 percent of Smith Creek flow was base flow, which suggests that Smith Creek streamflow is dominated by groundwater discharge rather than stormwater runoff. A series of cluster and principal components analyses demonstrated that the majority of the variability in Smith Creek water quality could be attributed to hydrologic and seasonal variability. Statistically significant positive correlations with flow were observed for turbidity, suspended sediments, total nitrogen, ammonium, orthophosphate, iron, total phosphorus, and the ratio of calcium to magnesium. Statistically significant inverse correlations with flow were observed for specific conductance, magnesium, δ 15 N of nitrate, pH, bicarbonate, calcium, and δ 18 O of nitrate. Of particular note, flow and nitrate were not statistically significantly correlated, likely because of the relatively complex concentration-discharge relationship observed in continuous and discrete datasets. Statistically significant seasonal patterns were observed for numerous water-quality constituents: water temperature, turbidity, orthophosphate, total phosphorus, suspended-sediment concentration, and silica were higher during the warm season, but pH, dissolved oxygen, and sulfate were higher during the cool season. Surrogate regression models were developed to compute sediment and nutrient loads in Smith Creek using the continuous water-quality monitors. The mean Smith Creek in-stream sediment load was approximately 6,900 tons per year, with nearly 90 percent of the sediment load over the 3-year study period contributed during the eight largest storm events during that period. The Smith Creek total phosphorus load was approximately 21,000 pounds of phosphorus per year, with the majority of the load contributed during stormflow periods, although a substantial phosphorus load still occurs during base-flow conditions. The Smith Creek total nitrogen load was approximately 400,000 pounds per year, with total nitrogen accumulation less dominated by stormflow contributions (as was the case for sediment and total phosphorus) and strongly affected by base-flow export of nitrogen from the basin. Extensive water-quality monitoring throughout the Smith Creek watershed revealed how the complex geology and hydrology interacted to result in variable water chemistry. During relatively dry and low base-flow periods, much of the discharge in Smith Creek was contributed by a single dominant spring—Lacey Spring. During wetter base-flow periods, the flows in Smith Creek were largely generated by a mixture of headwater springs and forested mountain tributaries with very different geochemical composition. The headwater springs generally issued from limestone bedrock and were characterized as having relatively high nitrate, specific conductance, calcium, and magnesium, as well as relatively low concentrations of phosphorus, ammonium, iron, and manganese. The undeveloped, high-gradient, forested mountain sites were generally characterized by low ionic strength waters with low nutrient concentrations. Nitrate isotope data from the limestone springs generally were consistent with manure-derived nitrogen sources (such as cattle and poultry), although the possibility of other mixed sources cannot be excluded. Nitrate isotope data from the undeveloped, high-gradient forested mountain sites were more consistent with nitrogen from undisturbed soils, atmospheric deposition, or nitrogen fixation. Regardless of the nitrogen source, oxygen isotope data indicate that the nitrate was largely a result of nitrification. Land-use data indicate that manure sources of nitrogen dominated watershed nitrogen inputs. Phosphorus sources were less well studied. The presence of a single point-source discharge near the town of New Market contributed the majority of the phosphorus to Smith Creek under base-flow conditions, but nonpoint sources of phosphorus dominated the loading to Smith Creek during stormflow periods. Implementation of conservation practices increased in the Smith Creek watershed during the study period, and even though a broad range of practice types was implemented, the most common practices included stream fencing (for cattle exclusion), the development of nutrient management plans, conservation crop rotation, and the planting of cover crops. While the implementation of these conservation practices is encouraging, results indicate small increases in nitrate concentrations at the streamgage over the last 29 years, concurrent with small decreases in nitrate fluxes. It will likely be years before the cumulative effect of these practices can be detected in the Smith Creek water quality, and the magnitude of the effect of these conservation practices detected in Smith Creek will depend largely on whether nutrient loading (of manure and commercial fertilizer) is reduced over time. Upper Chester River The Upper Chester River watershed includes the 36-square-mile (mi 2 ) watershed area around several nontidal tributaries that drain into the tidal Chester River. The streamgage is on Chesterville Branch, the largest nontidal tributary (approximately 6.12 mi 2 ) and is the site for continuous water-quality monitoring for this project. The base-flow index at Chesterville Branch is about 72 percent and indicates that, as in most of the Coastal Plain, groundwater is the greatest contributor to streamflow. As such, more than 90 percent of the nitrogen in the stream is in the form of nitrate from groundwater. Continuous and discrete data collected at Chesterville Branch show the effects of streamflow and season on water quality. Significantly positive correlations with flow were observed for ammonium, dissolved and total phosphorus, sediment, and turbidity as runoff carried these constituents from the land surface into Chesterville Branch. Other constituents that increased significantly with flow include potassium, sulfate, iron, and manganese, which are likely contributed from near-stream areas and ponds with high organic-matter content. Total nitrogen, pH, and specific conductance, along with chemical constituents associated with groundwater inputs including nitrate, calcium, ratio of calcium to magnesium, silica, bicarbonate, and sodium, were negatively correlated with flow because concentrations of these constituents were diluted by runoff. Seasonal differences in water chemistry, which are most likely related to increased biologic effects on the uptake and release of chemicals in the stream and near-stream areas, also were observed. Water temperature, orthophosphate, δ 15 N of nitrate, bicarbonate, sodium, and the ratio of sodium to chloride were higher during the warm season, and dissolved oxygen, total nitrogen, nitrate, magnesium, sulfate, and manganese were higher during the cool season. Surrogate-regression models developed by using continuous water-quality data showed that the annual sediment load for the 2013 water year was about 2,600 tons, with more than 90 percent of this sediment contributed during two storms. The total phosphorus load in 2013 was about 13,000 pounds with more than 90 percent contributed during the same two storms as sediment. The load of total nitrogen, 140,000 pounds, accumulated steadily throughout the 2013 water year as nitrate in groundwater continuously discharged into the stream. The same two large storms that contributed 90 percent of the suspended-sediment and total phosphorus load only contributed about 20 percent of the annual total nitrogen load. Extensive water-quality monitoring of stream base flow throughout the Upper Chester River watershed identified how differences in land use and hydrogeology affected water chemistry. In parts of the watershed with well-drained soil and thick sandy aquifer sediments, concentrations of nitrate and other chemicals associated with fertilizer and lime application increased in streams as agricultural land use increased. More than 90 percent of the nitrogen in streams from these areas was in the form of nitrate, and concentrations ranged from about 5 milligrams per liter (mg/L) to 8 mg/L as nitrogen in the two largest tributaries. Stream nitrate concentrations were about 1 mg/L as nitrogen where soils were more poorly drained, the surficial aquifer sediments were thinner, and forests and wetlands were more widespread than agriculture. Nitrate isotope data were consistent with inorganic fertilizers ± atmospheric deposition and N 2 fixation as sources of nitrogen, and with nitrification as the dominant nitrate-forming process. Nitrate reduction was indicated by elevated δ 15 N and δ 18 O values in some samples from streams draining watersheds with poorly drained soils. An analysis of land-use data and SPARROW modeling input data attributed almost 90 percent of the nitrogen sources in the Upper Chester River watershed to inorganic fertilizer and fixation of atmospheric nitrogen by legumes, which is in agreement with the isotopic characteristics of nitrate in this watershed. Local sources of manure are limited in this area. Total phosphorus concentrations during base flow ranged from below detection to about 0.2 mg/L. Stream phosphorus concentrations during base flow were generally lower than those measured during storms because most phosphorus transport likely occurs as phosphorus attached to sediment particles during runoff. Because manure is not widely used in this area, the major source of phosphorus is likely fertilizer. The implementation of conservation practices in the Upper Chester River watershed increased substantially during the study period, with a total implementation of 1,194 U.S. Department of Agriculture-compliant practices. The most frequently used practices were oriented towards nutrient and sediment control, including cover crops, nutrient management planning, conservation crop rotation, conservation tillage, and irrigation management. The current Chesapeake Bay model for this area predicts that implementation of best management practices should result in a 13-percent decrease in overall delivery of nitrogen to the Upper Chester River. Because most nitrogen travels through the groundwater system for years to decades before being discharged to streams, the time period of monitoring was not sufficient to see the effects of these practices on water quality. The magnitude of the effect that may eventually be detected will depend on the degree to which nitrate leaching into the groundwater system is reduced over time. Loadings of phosphorus and sediment are primarily transported during large runoff events and are difficult to control and analyze for trends because of their timing and episodic nature. Conewago Creek Conewago Creek has two primary monitoring locations—one near the middle of the 47-mi 2 watershed and the other near the outlet just upstream of the Susquehanna River. The base-flow index was 47.3 percent for 2012–2013, indicating that on average, approximately 53 percent of the streamflow in Conewago Creek exited the watershed as surface flow, which suggests that the stormwater runoff was somewhat greater than groundwater discharge (base flow). A series of cluster and principal components analyses demonstrated that the majority of the variability in the Conewago Creek water quality could be attributed to hydrologic and seasonal variability. Statistically significant positive correlations with flow were observed at both monitoring sites for ammonium, total phosphorus, orthophosphate, iron, and manganese; additionally, at the upstream monitoring station, total nitrogen demonstrated a statistically significant positive correlation with flow. Statistically significant inverse correlations with flow were observed at both sites for water temperature, specific conductance (at the downstream site only), sulfate, chloride, calcium, and magnesium. Statistically significant seasonal patterns were observed for several water-quality constituents. Water temperature, phosphorus (upstream site only), and orthophosphate were higher during the warm season, and nitrate and total nitrogen (upstream site only) were higher during the cool season. Surrogate regression models were developed to compute sediment and nutrient load in Conewago Creek by using the continuous water-quality monitors and water-quality samples. Conewago Creek sediment load was approximately 9,900 tons in 2012 and approximately 18,900 tons in 2013, with nearly 80 percent of the sediment load in 2013 contributed by the three largest storm events. Annual total nitrogen loads could not be estimated due to poor model performance. The addition of continued monitoring or a continuously recording nitrate sensor could improve estimates of total nitrogen loads. During 2012 and 2013, phosphorus loads in Conewago Creek were approximately 50,000 pounds in each year. Combining data from one high-flow synoptic sampling with the data from routine sampling revealed how the geology and hydrology interact to result in variable water chemistry throughout the Conewago Creek watershed. The areas above the upstream gage in the headwaters are generally underlain by forested non-carbonate bedrock and are characterized by relatively low nitrate, specific conductance, calcium, and magnesium, as well as relatively low concentrations of phosphorus, ammonium, iron, and manganese. The more developed, agricultural areas below the upstream site were generally characterized by higher ionic strength waters with higher nutrient and metal concentrations. An analysis of land-use data and SPAtially Referenced Regressions On Watershed (SPARROW) modeling data indicates that manure sources of nitrogen dominate the input of nitrogen to the watershed. Implementation of conservation practices increased in the Conewago Creek watershed during the study period, and while a broad range of practice types were implemented, the most common practices included residue and tillage management, cover crops, nutrient management, terracing, and stream fencing (for animal exclusion or bank restoration). While the implementation of these conservation practices is encouraging, the cumulative effects of these practices probably will not be detected in Conewago Creek water quality for several years. The magnitude of the effects of these conservation practices on water quality in Conewago Creek will depend largely on the extent to which nutrient loading (septic, manure, and commercial fertilizer) and sediment-producing activities are reduced over time. Difficult Run The Difficult Run watershed is a 57.82-mi 2 watershed that drains to the Potomac River. The long-term Difficult Run base-flow index (from 1936 to 2010) was 57.9, indicating that approximately 58 percent of streamflow exited the watershed as base flow and 42 percent as stormflow; however, with continued development and urbanization of the watershed, the base-flow index has decreased to 50 percent during the last 20 years. This base-flow index was less than those of the other watersheds evaluated in this study, likely because the Difficult Run watershed largely is underlain by crystalline piedmont metamorphic rocks and has a greater proportion of impervious urban land cover. A series of cluster and principal components analyses indicated that most of the variability in Difficult Run water quality could be attributed to hydrologic variability and seasonality. Statistically significant positive correlations with flow were observed for turbidity, dissolved oxygen, suspended sediments, ammonium, orthophosphate, iron, and total phosphorus. Statistically significant inverse correlations with flow were observed for water temperature, pH, specific conductance, bicarbonate, calcium, magnesium, nitrate, δ 15 N of nitrate, and silica. Statistically significant seasonal patterns were observed for numerous water-quality constituents: water temperature, ammonium, orthophosphate, and δ 15 N of nitrate were higher during the warm season, and dissolved oxygen, nitrate, and manganese were higher during the cool season. Surrogate regression models were developed to compute sediment and nutrient loading rates. The Difficult Run sediment load was approximately 8,000 tons per year, with greater than 95 percent of the sediment load in the 2013 water year contributed by the seven largest storm events. The total phosphorus load in Difficult Run was approximately 14,000 pounds of phosphorus per year, with the majority of the load contributed during stormflow periods. The total nitrogen load in Difficult Run is estimated to have been approximately 140,000 pounds per year, with total nitrogen accumulation less dominated by stormflow contributions than that of phosphorus and strongly affected by base-flow export of nitrogen from the basin. Extensive water-quality monitoring throughout the Difficult Run watershed revealed relatively uniform generation of flow per unit of watershed area, as well as spatial variation in water quality that is strongly related to land-use activities. Elevated nitrate concentrations were observed in a subset of monitoring sites that are inversely correlated with population density and positively correlated to the septic system density within each subwatershed. The majority of the elevated nitrate concentrations for these sites are hypothesized to be caused by nitrate leaching from septic systems, more so than homeowner fertilizer usage among these subwatersheds that have lower population densities than other parts of the watershed. Nitrate isotope data, temporal patterns in the water-quality data, mass-balance computations, and a separate land-use analysis all generally indicate that leachate from septic systems was the likely source of the elevated nitrate. Another group of water-quality sites have relatively low nitrogen concentrations, are located in areas that are served by city sewer lines, and have experienced stream restoration activities. A final group of sites drained the areas with the highest imperviousness and had strongly elevated specific conductance, chloride, and sodium, which were likely caused by a combination of road salting and other anthropogenic sources draining these urbanized areas in the watershed. A fourth group of sites represents a mixture of water sources and had water quality similar to that at the Difficult Run streamgage. Analysis of the nitrate isotope data generally indicates a broad range of composition indicative of mixed natural and anthropogenic nitrogen sources. Implementation of conservation practices increased in the Difficult Run watershed during the study period, and while a broad range of practice types was implemented, the most common practices included stream restoration. While the implementation of these conservation practices is encouraging, the cumulative effect of these practices probably will not be detected in Difficult Run water quality for several years.

Maryland, Pennsylvania, Virginia↗

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↗

Multiple mechanisms determine the effect of warming on plant litter decomposition in a dryland

In drylands, where soil fertility is typically low, plant litter decomposition provides particularly critical carbon and nitrogen inputs into soil. Although climate change is projected to increase the already large global extent of drylands, it is unknown how warmer temperatures will affect core ecosystem processes, such as plant litter decomposition, in these systems. To address this key unknown, we conducted a litterbag study in a long-term dryland warming experiment in southeastern Utah, USA. Unexpectedly, we did not find an overall effect of warming on leaf litter mass loss over time. Instead, our results indicated both positive and negative effects of warming on mass loss which offset one another. In particular, our findings suggested that a warming-induced degradation of biological soil crusts (soil surface community of mosses, lichens, and/or cyanobacteria that live in drylands worldwide) increased soil-litter mixing, thereby accelerating decomposition. Results also suggested that warming-induced decreases in litter moisture slowed decomposition. In addition to assessing mass loss, we found that warming lowered the carbon-to-nitrogen ratio of the decomposing litter. These results showed that warming did not alter the total litter mass-loss rates in this ecosystem, but that decomposition patterns were affected through more nuanced changes to both the biological and physical environment of dryland soils.

Utah↗

Production of greenhouse-grown biocrust mosses and associated cyanobacteria to rehabilitate dryland soil function

Mosses are an often-overlooked component of dryland ecosystems, yet they are common members of biological soil crust communities (biocrusts) and provide key ecosystem services, including soil stabilization, water retention, carbon fixation, and housing of N 2 fixing cyanobacteria. Mosses are able to survive long dry periods, respond rapidly to precipitation, and reproduce vegetatively. With these qualities, dryland mosses have the potential to be an excellent dryland restoration material. Unfortunately, dryland mosses are often slow growing in nature, and ex situ cultivation methods are needed to enhance their utility. Our goal was to determine how to rapidly produce, vegetatively, Syntrichia caninervis and S. ruralis , common and abundant moss species in drylands of North America and elsewhere, in a greenhouse. We manipulated the length of hydration on a weekly schedule (5, 4, 3, or 2 days continuous hydration per week), crossed with fertilization (once at the beginning, monthly, biweekly, or not at all). Moss biomass increased sixfold for both species in 4 months, an increase that would require years under dryland field conditions. Both moss species preferred short hydration and monthly fertilizer. Remarkably, we also unintentionally cultured a variety of other important biocrust organisms, including cyanobacteria and lichens. In only 6 months, we produced functionally mature biocrusts, as evidenced by high productivity and ecosystem-relevant levels of N 2 fixation. Our results suggest that biocrust mosses might be the ideal candidate for biocrust cultivation for restoration purposes. With optimization, these methods are the first step in developing a moss-based biocrust rehabilitation technology.

Restoration Ecology↗

Treatment effects on performance of N-fixing lichens in disturbed crusts of the Colorado Plateau

Biological soil crusts arrest soil erosion and supply nitrogen to arid ecosystems. To understand their recovery from disturbance, we studied performances of Collema spp. lichens relative to four experimental treatments plus microtopography of soil pedicels, oriented north-northwest to south-southeast in crusts. At sites in Needles (NDLS) and Island in the Sky (ISKY) districts of Canyonlands National Park, lichens were transplanted to NNW, SSE, ENE, WSW, and TOP pedicel faces and exposed to a full-factorial, randomized block experiment with four treatments: nutrient addition (P and K), soil stabilization with polyacrylamide resin (PAM), added cyanobacterial fiber, and biweekly watering. After 14.5 mo (NDLS) and 24 mo (ISKY), both visual rankings of lichen condition and measures of chlorophyll fluorescence were generally higher at ISKY than on more fertile but less stable soils at NDLS. On ENE and NNW pedicel faces, both these values and nitrogenase activity (NDLS only) exceeded corresponding values on WSW and SSE faces. Treatment effects were site specific and largely negative at NDLS; both nutrient and cyanobacterial addition led to poorer lichen condition, and added nutrients led to reduced fluorescence. Responses to nutrients may have been mediated partly by disturbance of unstable soils and by competition with cyanobacteria. In a separate experiment investigating recruitment responses to adding fungal spores or Nostoc cells, rates of Collema establishment responded significantly to the former but not the latter. Low establishment overall suggests that natural recruitment may occur mainly from iscidia or thallus fragments, not spores. Measured simultaneously on artificial soil pedicels at NDLS and ISKY, both deposition and erosion declined at NDLS across the four pedicel microaspects as: WSW > SSE > NNW > ENE (or ENE > NNW), during fall and spring trials. Patterns were similar at ISKY, but WSW ≈ SSE for spring deposition, and deposition did not differ by microaspect in fall. Greater deposition at ISKY, despite higher abundance of cyanobacteria, may be explained by stronger wind velocities. Together, microtopographic differences in erosion, microclimate, and nutrient regimes help explain variable lichen performance, but microtopography influenced lichen performance more consistently than did any treatment. Demonstrated effects of pedicel development in crust recovery concur with prior surveys showing greatest microbial biomass and/or cover on ENE and NNW exposures at various spatial scales.

Colorado Plateau↗

Transport of biologically important nutrients by wind in an eroding cold desert

Wind erosion following fire is an important landscape process that can result in the redistribution of ecologically important soil resources. In this study we evaluated the potential for a fire patch in a desert shrubland to serve as a source of biologically important nutrients to the adjacent, downwind, unburned ecosystem. We analyzed nutrient concentrations (P, K, Ca, Mg, Cu, Fe, Mn, Al) in wind-transported sediments, and soils from burned and adjacent unburned surfaces, collected during the first to second growing seasons after a wildfire that burned in 2007 in Idaho, USA in sagebrush steppe; a type of cold desert shrubland. We also evaluated the timing of potential wind erosion events and weather conditions that might have contributed to nutrient availability in downwind shrubland. Findings indicated that post-fire wind erosion resulted in an important, but transient, addition of nutrients on the downwind shrubland. Aeolian sediments from the burned area were enriched relative to both the up- and down-wind soil and indicated the potential for a fertilization effect through the deposition of the nutrient-enriched sediment during the first, but not second, summer after wildfire. Weather conditions that could have produced nutrient transport events might have provided increased soil moisture necessary to make nutrients accessible for plants in the desert environment. Wind transport of nutrients following fire is likely important in the sagebrush steppe as it could contribute to pulses of resource availability that might, for example, affect plant species differently depending on their phenology, and nutrient- and water-use requirements.

Idaho↗

Controls on sediment production in two U.S. deserts

Much of the world’s airborne sediment originates from dryland regions. Soil surface disturbances in these regions are ever-increasing due to human activities such as energy and mineral exploration and development, recreation, suburbanization, livestock grazing and cropping. Sediment production can have significant impacts to human health with particles potentially carrying viruses such as Valley Fever or causing asthma or other respiratory diseases. Dust storms can cause decreased visibility at the ground level, resulting in highway accidents, and reduced visual quality in park and wildland airsheds. Sediment production and deposition is also detrimental to ecosystem health, as production reduces soil fertility at its source and can bury plants and other organisms where it is deposited. Therefore, it is important to understand how we can predict what areas are prone to producing sediment emissions both before and after soil surface disturbance. We visited 87 sites in two deserts of the western U.S. that represented a range of soil texture and surface cover types. We used a portable wind tunnel to estimate the threshold friction velocity (TFV) required to initiate sediment transport and the amount of sediment produced by the tunnel at a set wind speed. Wind tunnel runs were done before and after soil surface disturbance with a four-wheel drive vehicle. Results show that most undisturbed desert soils are very stable, especially if covered by rocks or well-developed biological soil crusts, which make them virtually wind-erosion proof. Particles at disturbed sites, in contrast, moved at relatively low wind speeds and produced high amounts of sediment. Silt was an important predictor of TFV and sediment production across all sites, whereas the influence of rock cover and biological soil crusts was site-dependent. Understanding the vulnerability of a site after disturbance is important information for land managers as they plan land use activities and attempt to mitigate the harmful effects that sediment production can have on both human and ecosystem health.

Aeolian Research↗

Long-term tillage and crop rotation effects on residual nitrate in the crop root zone and nitrate accumulation in the intermediate vadose zone

Tillage influences the physical and biological environment of soil. Rotation of crops with a legume affects the soil N status. A furrow irrigated site was investigated for long-term tillage and crop rotation effects on leaching of nitrate from the root zone and accumulation in the intermediate vadose zone (IVZ). The investigated tillage systems were disk-plant (DP), ridge-till (RT) and slot-plant (SP). These tillage treatments have been maintained on the Hastings silt loam (Udic Argiustoll) and Crete silt loam (Pachic Argiustoll) soils since 1976. Continuous corn (CC) and corn soybean (CS) rotations were the subtreatments. Since 1984, soybeans have been grown in CS plots in even calendar years. All tillage treatments received the same N rate. The N rate varied annually depending on the root zone residual N. Soybeans were not fertilized with N-fertilizer. Samples for residual nitrate in the root zone were taken in 8 of the 15 year study while the IVZ was only sampled at the end of the study. In seven of eight years, root zone residual soil nitrate-N levels were greater with DP than RT and SP. Residual nitrate-N amounts were similar in RT and SP in all years. Despite high residual nitrate-N with DP and the same N application rate, crop yields were higher in RT and SP except when DP had an extremely high root zone nitrate level. By applying the same N rates on all tillage treatments, DP may have been fertilized in excess of crop need. Higher residual nitrate-N in DP was most likely due to a combination of increased mineralization with tillage and lower yield compared to RT and SP. Because of higher nitrate availability with DP, the potential for nitrate leaching from the root zone was greater with DP as compared to the RT and SP tillage systems. Spring residual nitrate-N contents of DP were larger than RT and SP in both crop rotations. Ridge till and SP systems had greater nitrate-N with CS than CC rotations. Nitrate accumulation in IVZ at the upstream end of the field was twice as high with DP compared to RT and SP. At the downstream end, it was 2.4 and 1.6 times greater with DP than RT and SP, respectively. Nitrate concentration was greater in the IVZ of DP compared to RT and SP tillage systems. Nitrate accumulations in IVZ of RT and SP were not different. Continuous corn had slightly higher nitrate levels in IVZ than CS. The depth of nitrate penetration at the upstream end was greater than that of the downstream end. Estimated rates of nitrate movement ranged from 0.87 to 0.92 m yr-1 at the upstream end and 0.73 to 0.78 m yr-1 at the downstream end.

Transactions of the American Society of Agricultur↗

A hierarchical perspective of plant diversity

Predictive models of plant diversity have typically focused on either a landscapea??s capacity for richness (equilibrium models), or on the processes that regulate competitive exclusion, and thus allow species to coexist (nonequilibrium models). Here, we review the concepts and purposes of a hierarchical, multiscale model of the controls of plant diversity that incorporates the equilibrium model of climatic favorability at macroscales, nonequilibrium models of competition at microscales, and a mixed model emphasizing environmental heterogeneity at mesoscales. We evaluate the conceptual model using published data from three spatially nested datasets: (1) a macroscale analysis of ecoregions in the continental and western U.S.; (2) a mesoscale study in California; and (3) a microscale study in the Siskiyou Mountains of Oregon and California. At the macroscale (areas from 3889 km2 to 638,300 km2), climate (actual evaporation) was a strong predictor of tree diversity (R2 = 0.80), as predicted by the conceptual model, but area was a better predictor for vascular plant diversity overall (R2 = 0.38), which suggests different types of plants differ in their sensitivity to climatic controls. At mesoscales (areas from 1111 km2 to 15,833 km2 ), climate was still an important predictor of richness (R2 = 0.52), but, as expected, topographic heterogeneity explained an important share of the variance (R2 = 0.19), showed positive correlations with diversity of trees, shrubs, and annual and perennial herbs, and was the primary predictor of shrub and annual plant species richness. At microscales (0.1 ha plots), spatial patterns of diversity showed a clear unimodal pattern along a climatea??driven productivity gradient and a negative relationship with soil fertility. The strong decline in understory and total diversity at the most productive sites suggests that competitive controls, as predicted, can override climatic controls at this scale. We conclude that this hierarchical, multiscale model provides a sound basis to understand and analyze plant species diversity. Specifically, future research should employ the principles in this paper to explore climatic controls on species richness of different life forms, better quantify environmental heterogeneity in landscapes, and analyze how these largea??scale factors interact with local nonequilibrium dynamics to maintain plant diversity.

The Quarterly Review of Biology↗

Muted responses to chronic experimental nitrogen deposition on the Colorado Plateau

Anthropogenic nitrogen (N) deposition is significantly altering both community structure and ecosystem processes in terrestrial ecosystems across the globe. However, our understanding of the consequences of N deposition in dryland systems remains relatively poor, despite evidence that drylands may be particularly vulnerable to increasing N inputs. In this study, we investigated the influence of 7 years of multiple levels of simulated N deposition (0, 2, 5, and 8 kg N ha −1 year −1 ) on plant community structure and biological soil crust (biocrust) cover at three semi-arid grassland sites spanning a soil texture gradient. Biocrusts are a surface community of mosses, lichens, cyanobacteria, and/or algae, and have been shown to be sensitive to N inputs. We hypothesized that N additions would decrease plant diversity, increase abundance of the invasive annual grass Bromus tectorum, and decrease biocrust cover. Contrary to our expectations, we found that N additions did not affect plant diversity or B. tectorum abundance. In partial support of our hypotheses, N additions negatively affected biocrust cover in some years, perhaps driven in part by inter-annual differences in precipitation. Soil inorganic N concentrations showed rapid but ephemeral responses to N additions and plant foliar N concentrations showed no response, indicating that the magnitude of plant and biocrust responses to N fertilization may be buffered by endogenous N cycling. More work is needed to determine N critical load thresholds for plant community and biocrust dynamics in semi-arid systems and the factors that determine the fate of N inputs.

Arizona, Colorado, New Mexico, Utah↗

Germination and growth of native and invasive plants on soil associated with biological control of tamarisk (Tamarix spp.)

Introductions of biocontrol beetles (tamarisk beetles) are causing dieback of exotic tamarisk in riparian zones across the western United States, yet factors that determine plant communities that follow tamarisk dieback are poorly understood. Tamarisk-dominated soils are generally higher in nutrients, organic matter, and salts than nearby soils, and these soil attributes might influence the trajectory of community change. To assess physical and chemical drivers of plant colonization after beetle-induced tamarisk dieback, we conducted separate germination and growth experiments using soil and litter collected beneath defoliated tamarisk trees. Focal species were two common native (red threeawn, sand dropseed) and two common invasive exotic plants (Russian knapweed, downy brome), planted alone and in combination. Nutrient, salinity, wood chip, and litter manipulations examined how tamarisk litter affects the growth of other species in a context of riparian zone management. Tamarisk litter, tamarisk litter leachate, and fertilization with inorganic nutrients increased growth in all species, but the effect was larger on the exotic plants. Salinity of 4 dS m −1 benefitted Russian knapweed, which also showed the largest positive responses to added nutrients. Litter and wood chips generally delayed and decreased germination; however, a thinner layer of wood chips increased growth slightly. Time to germination was lengthened by most treatments for natives, was not affected in exotic Russian knapweed, and was sometimes decreased in downy brome. Because natives showed only small positive responses to litter and fertilization and large negative responses to competition, Russian knapweed and downy brome are likely to perform better than these two native species following tamarisk dieback.

Invasive Plant Science and Management↗

Water-quality assessment of the Albemarle-Pamlico drainage basin, North Carolina and Virginia environmental setting and water-quality issues

The Albemarle-Pamlico drainage study unit is one of 60 units of the U.S. Geological Survey's National Water-Quality Assessment Program, and includes the large river basins which drain into the Albemarle and Pamlico Sounds-the Chowan, Roanoke, Tar-Pamlico, and Neuse River Basins. The study unit includes about 28,000 square miles and has an interrelated set of environmental characteristics which strongly influence water quality. The chemical and physical nature of these characteristics are the dominant controls on baseline water quality in the study area. About 50 percent of the study area is forested, slightly more than 30 percent is agricultural, about 15 percent is wetlands, and less than 5 percent is developed. Three million people live in the study area, and activities related to agriculture and development have caused increased concentrations of constituents such as nutrients, pesticides, and suspended sediment. About two-thirds of the 36 to 52 inches of precipitation in the area reenters the atmosphere by evapotranspiration. About one-third of the remaining precipitation reaches streams by overland runoff; the remainder recharges the water table aquifer, where much of the water eventually discharges to streams as ground water. Thus, ground-water quality has a substantial influence on surface-water quality, particularly during dry weather. In 1990, about 152,900 tons of elemental nitrogen and 10,500 tons of elemental phosphorus either were applied to crops as fertilizer or fixed by biological processes, and in 1987, about 43,500 tons of nitrogen and 12,200 tons of phosphorus were produced as animal wastes. In addition, about 1,300 tons of selected herbicides and 400 tons of selected insecticides were applied to crops in 1990. Some 249 permitted point sources discharged 410 million gallons per day, containing an annual load of 5,800 tons of nitrogen and 1,800 tons of phosphorus, to the study area in 1990. Data from 1970-79 indicate that mean annual suspended-sediment yields for selected forested, agricultural, and developed urban basins in North Carolina are 50, 250, and 550 tons per square mile, respectively. In order to facilitate comparisons, much of the data were compiled by hydrologic unit. Homogeneous areas, or strata, representing the most prevalent combinations of environmental factors, such as land use, soils, and geology, were defined. Future data collection and analyses will be designed to answer objective-related concerns about the relations between important water-quality conditions and these study-unit strata.

North Carolina, Virginia↗

Effects of elevated CO2 on fine root dynamics in a Mojave Desert community: A FACE study

Fine roots (??? 1mm diameter) are critical in plant water and nutrient absorption, and it is important to understand how rising atmospheric CO2 will affect them as part of terrestrial ecosystem responses to global change. This study's objective was to determine effects of elevated CO2 on production, mortality, and standing crops of fine root length over 2 years in a free-air CO2 enrichment (FACE) facility in the Mojave Desert of southern Nevada, USA. Three replicate 25m diameter FACE rings were maintained at ambient (??? 370 ??mol mol-1) and elevated CO2 (??? 550 ??mol mol-1) atmospheric concentrations. Twenty-eight minirhizotron tubes were placed in each ring to sample three microsite locations: evergreen Larrea shrubs, drought-deciduous Ambrosia shrubs, and along systematic community transects (primarily in shrub interspaces which account for ??? 85% of the area). Seasonal dynamics were similar for ambient and elevated CO2: fine root production peaked in April-June, with peak standing crop occurring about 1 month later, and peak mortality occurring during the hot summer months, with higher values for all three measures in a wet year compared with a dry year. Fine root standing crop, production, and mortality were not significantly different between treatments except standing crop along community transects, where fine root length was significantly lower in elevated CO2. Fine root turnover (annual cumulative mortality/mean standing crop) ranged from 2.33 to 3.17 year-1, and was not significantly different among CO2 treatments, except for community transect tubes where it was significantly lower for elevated CO2. There were no differences in fine root responses to CO2 between evergreen (Larrea) and drought-deciduous (Ambrosia) shrubs. Combined with observations of increased leaf-level water-use efficiency and lack of soil moisture differences, these results suggest that under elevated CO2 conditions, reduced root systems (compared with ambient CO2) appear sufficient to provide resources for modest aboveground production increases across the community, but in more fertile shrub microsites, fine root systems of comparable size with those in ambient CO2 were required to support the greater aboveground production increases. For community transects, development of the difference in fine root standing crops occurred primarily through lower stimulation of fine root production in the elevated CO2 treatment during periods of high water availability. ?? 2005 Blackwell Publishing Ltd.

Global Change Biology↗