Small effects of livestock grazing intensification on diversity, abundance, and composition in a dryland plant community.

Livestock grazing is a globally important land use and has the potential to significantly influence plant community structure and ecosystem function, yet several critical knowledge gaps remain on the direction and magnitude of grazing impacts. Furthermore, much of our understanding of the long-term effects on plant community composition and structure are based on grazer exclusion experiments, which explicitly avoid characterizing effects along grazing intensity gradients. We sampled big sagebrush plant communities using 68 plots located along grazing intensity gradients to determine how grazing intensity influences multiple aspects of plant community structure over time. This was accomplished by sampling plant communities at different distances from 17 artificial watering sources, using distance from water and cow dung density as proxies for grazing intensity at individual plots. Total vegetation cover and total grass cover were negatively related to grazing intensity, and cover of annual forbs, exotic cover, and exotic richness were positively related to grazing intensity. In contrast, species richness and composition, bunchgrass biomass, shrub density and size, percentage cover of bare ground, litter, and biological soil crusts did not vary along our grazing intensity gradients, in spite of our expectations to the contrary. Our results suggest that the effects of livestock grazing over multiple decades (mean = 46 years) in our sites are relatively small, especially for native perennial species, and that the big sagebrush plant communities we sampled are somewhat resistant to livestock grazing. Collectively, our findings are consistent with existing evidence that indicates the stability of the big sagebrush plant functional type composition under current grazing management regimes.

Soil and stand structure explain shrub mortality patterns following global change-type drought and extreme precipitation.

The probability of extreme weather events is increasing, with the potential for widespread impacts to plants, plant communities, and ecosystems. Reports of drought-related tree mortality are becoming more frequent, and there is increasing evidence that drought accompanied by high temperatures is especially detrimental. Simultaneously, extreme large precipitation events have become more frequent over the past century. Water-limited ecosystems may be more vulnerable to these extreme events than other ecosystems, especially when pushed outside of their historical range of variability. However, drought-related mortality of shrubs-an important component of dryland vegetation-remains understudied relative to tree mortality. In 2014, a landscape-scale die-off of the widespread shrub, big sagebrush (Artemisia tridentata Nutt.), was reported in southwest Wyoming, following extreme hot and dry conditions in 2012 and extremely high precipitation in September of 2013. Here we examine how severe drought, extreme precipitation, soil texture and salinity, and shrub-stand characteristics contributed to this die-off event. At 98 plots within and around the die-off, we quantified big sagebrush mortality, characterized soil texture and salinity, and simulated soil-water conditions from 1916 to 2016 using an ecosystem water-balance model. We found that the extreme weather conditions alone did not explain patterns of big sagebrush mortality and did not result in extreme (historically unprecedented) soil-water conditions during the drought. Instead, plots with chronically dry soil conditions experienced greatest mortality following the global change-type (hot) drought in 2012. Furthermore, mortality was greater in locations with high potential run-on and low potential run-off where saturated soil conditions were simulated in September 2013, suggesting that extreme precipitation also played an important role in the die-off in these locations. In locations where drought alone contributed to mortality, stem density negatively impacted big sagebrush. In locations that may have been affected by both drought and saturation, however, mortality was greatest where stem density was lowest, suggesting that these locations may have already been less favorable to big sagebrush. Paradoxically, vulnerability to both extreme events (drought and saturation) was associated with finer-textured soils, and our results highlight the importance of soils in determining local variation of the vulnerability of dryland plants to extreme events.

Soil texture and precipitation seasonality influence plant community structure in North American temperate shrub steppe.

In drylands, the coexistence of grasses and woody plants has been attributed to soil-water resource partitioning. Soil texture and precipitation seasonality can influence the amount and distribution of water in the soil, and their interaction may play an important role in determining the relative importance of grasses and woody plants. We investigated the influence of this interaction on plant functional types across a broad range of precipitation regimes and soil textures in western North America by analyzing plant-cover data collected at 2,084 plots that included the widespread shrub big sagebrush (Artemisia tridentata Nutt.). We characterized how the significance of the inverse-texture effect varies across soil conditions by quantifying relationships between precipitation and foliar cover on finer- vs. coarser-textured soils across a range of potential texture divisions represented by sand content. We found evidence of the inverse-texture effect for every plant functional type (except for cheatgrass) that we examined with at least one component of precipitation (annual, warm, or cold season), and provide the first evidence for this effect in locations with cold-season-dominated precipitation regimes. The texture and precipitation combinations that exhibited the inverse-texture effect varied with plant functional type, presumably because of effects of soil texture on water availability at different soil depths with season. Furthermore, we found an inverse-texture effect that was remarkably similar for shrub cover with cold-season precipitation and grass cover with warm-season precipitation. These results provide new insight into how the inverse-texture effect interacts with precipitation seasonality to influence plant functional type composition in drylands, and further suggest that quantifying the soil-texture division at which the inverse-texture effect is relevant under a given set of environmental conditions may provide support for the effect across dryland plant communities.

Nitrogen addition pulse has minimal effect in big sagebrush (Artemisia tridentata) communities on the Pinedale Anticline, Wyoming (USA).

Nitrogen additions are known to elicit variable responses in semi-arid ecosystems, with responses increasing with precipitation. The response of semi-arid ecosystems to nitrogen are important to understand due to their large spatial extent worldwide and the global trend of increasingly available nitrogen. In this study, we evaluated the impact of a single nitrogen addition pulse on a semi-arid big sagebrush (Artemisia tridentata) ecosystem in western Wyoming. This is important given that sagebrush ecosystems are poorly understood, despite their prevalence in the western US. In addition, large-scale nitrogen additions have begun on sagebrush landscapes in Wyoming in order to mitigate population declines in mule deer (Odocoileus hemionus). The study objectives were (1) to evaluate the effectiveness of a nitrogen fertilization pulse in increasing sagebrush biomass and forage quality, and (2) to assess effects of nitrogen addition on soil biogeochemistry and vegetation community structure. We fertilized 15 plots across 5 locations in western Wyoming using a single pulse of urea (5.5g N m-2). In addition, we immobilized available nitrogen through surface hay treatments (250g hay/m2). Nitrogen additions failed to increase growth of sagebrush, alter nitrogen content of sagebrush leaders, or alter greenhouse gas efflux from soils. The plant community also remained unchanged; total cover, species richness, and community composition were all unaffected by our treatment application. Over the two years of this study, we did not find indications of nitrogen limitation of ecosystem processes, despite a wet growing season in 2014. Thus, we have found a general lack of response to nitrogen in sagebrush ecosystems and no treatment effect of a single pulse of N to sagebrush biomass or forage quality.

Inorganic nitrogen supply and dissolved organic nitrogen abundance across the US Great Plains.

Across US Great Plains grasslands, a gradient of increasing mean annual precipitation from west to east corresponds to increasing aboveground net primary productivity (ANPP) and increasing N-limitation. Previous work has shown that there is no increase in net N mineralization rates across this gradient, leading to the question of where eastern prairie grasses obtain the nitrogen to support production. One as-yet unexamined source is soil organic N, despite abundant literature from other ecosystems showing that plants take up dissolved soil organic N. This study measured KCl-extractable dissolved organic N (DON) in surface soils across the grassland productivity gradient. We found that KCl-extractable DON pools increased from west to east. If available to and used by plants, this DON may help explain the high ANPP in the eastern Great Plains. These results suggest a need for future research to determine whether, in what quantities, and in what forms prairie grasses use organic N to support primary production.

Is bacterial moisture niche a good predictor of shifts in community composition under long-term drought?

Both biogeographical and rainfall manipulation studies show that soil water content can be a strong driver of microbial community composition. However, we do not yet know if these patterns emerge because certain bacterial taxa are better able to survive at dry soil moisture regimes or if they are due to other drought-sensitive ecosystem properties indirectly affecting microbial community composition. In this study, we evaluated (1) whether bacterial community composition changed under an 11-year drought manipulation and (2) whether shifts under drought could be explained by variation in the moisture sensitivity of growth among bacterial taxa (moisture niche partitioning). Using 454 pyrosequencing of 16S rRNA, we observed shifts in bacterial community composition under drought, coincident with changes in other soil properties. We wet-up dry soils from drought plots to five moisture levels, and measured respiration and the composition of actively growing communities using bromodeoxyuridine (BrdU) labeling of DNA. The field drought experiment affected the composition of the active community when incubated at different moisture levels in the laboratory, as well as short-term (36-hour) respiration rates. Independent of history, bacterial communities also displayed strong niche partitioning across the wet-up moisture gradient. Although this indicates that moisture has the potential to drive bacterial community composition under long-term drought, species distributions predicted by response to moisture did not reflect the community composition of plots that were subjected to long-term drought. Bacterial community structure was likely more strongly driven by other environmental factors that changed under long-term drought, or not shaped by response to water level upon wet-up. The approach that we present here for linking niches to community composition could be adapted for other environmental variables to aid in predicting microbial species distributions and community responses to environmental change.

Grazing and no-till cropping impacts on nitrogen retention in dryland agroecosystems.

As the world's population increases, marginal lands such as drylands are likely to become more important for food production. One proven strategy for improving crop production in drylands involves shifting from conventional tillage to no-till to increase water use efficiency, especially when this shift is coupled with more intensive crop rotations. Practices such as no-till that reduce soil disturbance and increase crop residues may promote C and N storage in soil organic matter, thus promoting N retention and reducing N losses. By sampling soils 15 yr after a N tracer addition, this study compared long-term soil N retention across several agricultural management strategies in current and converted shortgrass steppe ecosystems: grazed and ungrazed native grassland, occasionally mowed planted perennial grassland, and three cropping intensities of no-till dryland cropping. We also examined effects of the environmental variables site location and topography on N retention. Overall, the long-term soil N retention of >18% in these managed semiarid ecosystems was high compared with published values for other cropped or grassland ecosystems. Cropping practices strongly influenced long-term N retention, with planted perennial grass systems retaining >90% of N in soil compared with 30% for croplands. Grazing management, topography, and site location had smaller effects on long-term N retention. Estimated 15-yr N losses were low for intact and cropped systems. This work suggests that semiarid perennial grass ecosystems are highly N retentive and that increased intensity of semiarid land management can increase the amount of protein harvested without increasing N losses.

Conservation of nitrogen increases with precipitation across a major grassland gradient in the Central Great Plains of North America.

Regional analyses and biogeochemical models predict that ecosystem N pools and N cycling rates must increase from the semi-arid shortgrass steppe to the sub-humid tallgrass prairie of the Central Great Plains, yet few field data exist to evaluate these predictions. In this paper, we measured rates of net N mineralization, N in above- and belowground primary production, total soil organic matter N pools, soil inorganic N pools and capture in resin bags, decomposition rates, foliar (15)N, and N use efficiency (NUE) across a precipitation gradient. We found that net N mineralization did not increase across the gradient, despite more N generally being found in plant production, suggesting higher N uptake, in the wetter areas. NUE of plants increased with precipitation, and delta(15)N foliar values and resin-captured N in soils decreased, all of which are consistent with the hypothesis that N cycling is tighter at the wet end of the gradient. Litter decomposition appeared to play a role in maintaining this regional N cycling trend: litter decomposed more slowly and released less N at the wet end of the gradient. These results suggest that immobilization of N within the plant-soil system increases from semi-arid shortgrass steppe to sub-humid tallgrass prairie. Despite the fact that N pools increase along a bio-climatic gradient from shortgrass steppe to mixed grass and tallgrass prairie, this element becomes relatively more limiting and is therefore more tightly conserved at the wettest end of the gradient. Similar to findings from forested systems, our results suggest that grassland N cycling becomes more open to N loss with increasing aridity.

Global-scale similarities in nitrogen release patterns during long-term decomposition.

Litter decomposition provides the primary source of mineral nitrogen (N) for biological activity in most terrestrial ecosystems. A 10-year decomposition experiment in 21 sites from seven biomes found that net N release from leaf litter is dominantly driven by the initial tissue N concentration and mass remaining regardless of climate, edaphic conditions, or biota. Arid grasslands exposed to high ultraviolet radiation were an exception, where net N release was insensitive to initial N. Roots released N linearly with decomposition and exhibited little net N immobilization. We suggest that fundamental constraints on decomposer physiologies lead to predictable global-scale patterns in net N release during decomposition.

Plant effects on nitrogen retention in shortgrass steppe 2 years after (15)N addition.

Ecosystems where plant-available nitrogen (N) is limited by constraints on decomposition may be quite capable of retaining additional N. However, there are many factors that will control the quantity of N retained, with potential implications for system carbon and nitrogen storage. We examined the retention and allocation of (15)N 2 years after labeling a semiarid, shortgrass steppe ecosystem in northeastern Colorado. The plant community of the study area is a patchy mixture of C3 (cool-season) and C4 (warm-season) graminoids; we hypothesized that differences in allocation patterns between the two plant types would lead to differing rates of N retention in this grazed system. We found that after three growing seasons (just over 2 years), an average of 28.3% of the original (15)N was retained in our plots, with nearly all of this N in soils (24.9%) rather than plants. Plots dominated by C3 plants had significantly less (15)N retained after 2 years than mixed C3-C4 plots. A high initial rate of retention by C3 plants, combined with a propensity for allocation to shoots rather than roots, likely led to this result in a system that typically has much of its aboveground tissue removed by grazers. In comparing our retention patterns to those of other studies in the shortgrass steppe, we found that the distribution of added (15)N to various ecosystem compartments (plants, mineral soil, soil organic matter) was similar across studies regardless of the experimental conditions, duration of study, and amount of (15)Nretained. We additionally observed the beginning of the formation of "resource islands," with (15)N being physically and biologically redistributed under plants from between plants.

Ecosystem consequences of plant life form changes at three sites in the semiarid United States.

Many semiarid rangelands have recently experienced changes in dominant plant life form. Both woody plant expansion into grasslands and the invasion of annual grasses into shrublands have potential influence on regional carbon cycling. Soil carbon content, chemistry, and distribution may change following shifts in dominant plant life form because plant life forms differ in litter chemistry and patterns of detrital input. This study assesses the amount, quality, and distribution of soil C below woody vegetation and grasses at three rangelands in Texas, New Mexico, and Utah. At each of these sites there has been a well-documented shift in dominant plant life form. In Texas and New Mexico, woody plants have increased in grasslands, while grasses have invaded into former shrublands in Utah. We measured total soil carbon, particulate organic matter (POM) C, and the carbon isotopic composition of soil carbon beneath woody plants and grasses at each of these three sites. At the La Copita Research Area in south-central Texas there was significantly more soil C found beneath Prosopis glandulosa, the dominant woody plant, than was found beneath grasses. Mean soil C content to 1 m was 7.2 kg C m-2 beneath P. glandulosa and 6.0 kg C m-2 beneath grasses. There was also significantly more POM C beneath P. glandulosa than beneath grasses. Stable carbon isotopic composition indicated that the expansion of P. glandulosa in savannas in Texas first influences carbon cycling in surface soils, then deep soil C, and finally throughout the soil profile. At the Sevilleta National Wildlife Refuge in central New Mexico, we found that there was significantly more soil C in the upper 10 cm of the soil profile beneath Larrea tridentata than was found beneath Bouteloua spp. Stable carbon isotopic composition indicated that the expansion of L. tridentata influenced C cycling throughout the soil profile. At Curlew Valley in northern Utah, we found no significant differences in total profile soil C beneath different plant life forms. However, there was significantly more soil C found at the soil surface beneath woody plants than was observed beneath annual grasses. There was significantly less POM C beneath annual grasses than was found beneath woody plants or perennial grasses. Based on stable carbon isotopic analyses, we concluded that the invasion of grasses into shrublands influenced only the upper 30 cm of the soil profile. We determined that following changes in plant life form dominance, the most consistent change in soil C was an alteration in content and distribution of POM C, a slowly cycling pool of soil C. While we failed to find a consistent change in total profile soil C with plant life form across our sites, the change in soil C chemistry may have important implications for long-term soil C storage in semiarid systems where there have been shifts in plant life form.

Contingent effects of plant species on soils along a regional moisture gradient in the Great Plains.

The central grassland region of the United States encompasses major gradients in temperature and precipitation that determine the distribution of plant life forms, which in turn may influence key ecosystem processes such as nutrient cycling and soil organic matter dynamics. One such gradient is the threefold increase in precipitation from the eastern Colorado shortgrass-steppe, in the rain shadow of the Rocky Mountains, to the tallgrass prairie in eastern Kansas. We investigated the relative roles of plant species and plant cover in influencing soil C and N cycling in three sites along this gradient. Plant cover (i.e., the presence or absence of an individual plant) was relatively more important than plant species in explaining variability in soil properties at the dry site, the Central Plains Experimental Range in␣northeastern Colorado. However, plant species explained relatively more of the variability in soil properties than did plant cover at the two wetter sites, Hays and Konza, in central and eastern Kansas. The wetter sites had more continuous plant cover, resulting in less plant-cover-induced variation in soil C and N, than did the dry site, which had distinct patches of bare ground. Plant species at the wetter sites had higher and more variable levels of tissue C:N than plant species at the dry site, due to both within species changes and changes in species composition. Aboveground tissue C:N was better correlated with net nitrogen mineralization rates at the wet sites than the dry site. Thus, tissue chemistry appears to exert more control on nitrogen dynamics at the wet than the dry sites. The results suggest that plant species traits that are relevant to nutrient cycling (e.g., tissue C:N ratios, spatial patterns, productivity) reflect environmental limitations as well as species' physiological potentials. Furthermore, a dominant environmental driver such as precipitation may ameliorate or exaggerate the importance of individual species traits for nutrient cycling.