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Grass-dominated, dry rangelands account for over 30% of Earth's terrestrial surface (9,10) and provide most of the forage for the world's domestic livestock. Among the most important of these include [...], Global warming is predicted to induce desiccation in many world regions through increases in evaporative demand (1-3). Rising C[O.sub.2] may counter that trend by improving plant water-use efficiency (4,5). However, it is not clear how important this C[O.sub.2]-enhanced water use efficiency might be in offsetting warming-induced desiccation because higher C[O.sub.2] also leads to higher plant biomass, and therefore greater transpirational surface (2,6,7). Furthermore, although warming is predicted to favour warm-season, [C.sub.4] grasses, rising C[O.sub.2] should favour [C.sub.3], or cool-season plants (8). Here we show in a semi-arid grassland that elevated C[O.sub.2] can completely reverse the desiccating effects of moderate warming. Although enrichment of air to 600 p.p.m.v. C[O.sub.2] increased soil water content (SWC), 1.5/3.0 °C day/night warming resulted in desiccation, such that combined C[O.sub.2] enrichment and warming had no effect on SWC relative to control plots. As predicted, elevated C[O.sub.2] favoured [C.sub.3] grasses and enhanced stand productivity, whereas warming favoured [C.sub.4] grasses. Combined warming and C[O.sub.2] enrichment stimulated above-ground growth of [C.sub.4] grasses in 2 of 3 years when soil moisture most limited plant productivity. The results indicate that in a warmer, C[O.sub.2]-enriched world, both SWC and productivity in semi-arid grasslands may be higher than previously expected.

Understanding local adaptation to climate is critical for managing ecosystems in the face of climate change. While there have been many provenance studies in trees, less is known about local adaptation in herbaceous species, including the perennial grasses that dominate arid and semiarid rangeland ecosystems. We used a common garden study to quantify variation in growth and drought resistance traits in 99 populations of Elymus elymoides from a broad geographic and climatic range in the western United States. Ecotypes from drier sites produced less biomass and smaller seeds, and had traits associated with greater drought resistance: small leaves with low osmotic potential and high integrated water use efficiency (δ13C). Seasonality also influenced plant traits. Plants from regions with relatively warm, wet summers had large seeds, large leaves, and low δ13C. Irrespective of climate, we also observed trade‐offs between biomass production and drought resistance traits. Together, these results suggest that much of the phenotypic variation among E. elymoides ecotypes represents local adaptation to differences in the amount and timing of water availability. In addition, ecotypes that grow rapidly may be less able to persist under dry conditions. Land managers may be able to use this variation to improve restoration success by seeding ecotypes with multiple drought resistance traits in areas with lower precipitation. The future success of this common rangeland species will likely depend on the use of tools such as seed transfer zones to match local variation in growth and drought resistance to predicted climatic conditions. [ABSTRACT FROM AUTHOR]

Global changes that alter soil water availability may have profound effects on semiarid ecosystems. Although both elevated CO2 (eCO2) and warming can alter water availability, often in opposite ways, few studies have measured their combined influence on the amount, timing, and temporal variability of soil water. Here, we ask how free air CO2 enrichment (to 600 ppmv) and infrared warming (+ 1.5 °C day, + 3 °C night) effects on soil water vary within years and across wet-dry periods in North American mixed-grass prairie. We found that eCO2 and warming interacted to influence soil water and that those interactions varied by season. In the spring, negative effects of warming on soil water largely offset positive effects of eCO2. As the growing season progressed, however, warming reduced soil water primarily (summer) or only (autumn) in plots treated with eCO2. These interactions constrained the combined effect of eCO2 and warming on soil water, which ranged from neutral in spring to positive in autumn. Within seasons, eCO2 increased soil water under drier conditions, and warming decreased soil water under wetter conditions. By increasing soil water under dry conditions, eCO2 also reduced temporal variability in soil water. These temporal patterns explain previously observed plant responses, including reduced leaf area with warming in summer, and delayed senescence with eCO2 plus warming in autumn. They also suggest that eCO2 and warming may favor plant species that grow in autumn, including winter annuals and C3 graminoids, and species able to remain active under the dry conditions moderated by eCO2. [ABSTRACT FROM AUTHOR]

Plant roots mediate the impacts of environmental change on ecosystems, yet knowledge of root responses to environmental change is limited because few experiments evaluate multiple environmental factors and their interactions. Inferences about root functions are also limited because root length dynamics are rarely measured.Using a 5‐year experiment in a mixed‐grass prairie, we report the responses of root biomass, length and life span to elevated carbon dioxide (CO2), warming, elevated CO2 and warming combined, and irrigation. Root biomass was quantified using soil cores and root length dynamics were assessed using minirhizotrons. By comparing root dynamics with published results for soil resources and above‐ground productivity, we provide mechanistic insights into how climate change might impact grassland ecosystems.In the upper soil layer, 0–15 cm depth, both irrigation and elevated CO2 alone increased total root length by twofold, but irrigation decreased root biomass and elevated CO2 had only small positive effects on root biomass. The large positive effects of irrigation and elevated CO2 alone on total root length were due to increases in both root length production and root life span. The increased total root length and life span under irrigation and elevated CO2 coincided with apparent shifts from water limitation of plant growth to nitrogen limitation. Warming alone had minimal effects on root biomass, length and life span in this shallow soil layer. Warming and elevated CO2 combined increased root biomass and total root length by c. 25%, but total root length in this treatment was lower than expected if the effects of CO2 and warming alone were additive. Treatment effects on total root length and root life span varied with soil depth and root diameter.Synthesis. Sub‐additive effects of CO2 and warming suggest studies of elevated CO2 alone might overestimate the future capacity of grassland root systems to acquire resources. In this mixed‐grass prairie, elevated CO2 with warming stimulated total root length and root life span in deeper soils, likely enhancing plant access to more stable pools of growth‐limiting resources, including water and phosphorus. Thus, these root responses help explain previous observations of higher, and more stable, above‐ground productivity in these projected climate conditions. Sub‐additive effects of CO2 and warming suggest studies of elevated CO2 alone might overestimate the future capacity of grassland root systems to acquire resources. In this mixed‐grass prairie, elevated CO2 with warming stimulated total root length and root life span in deeper soils, likely enhancing plant access to more stable pools of growth‐limiting resources, including water and phosphorus. Thus, these root responses help explain previous observations of higher, and more stable, above‐ground productivity in these projected climate conditions [ABSTRACT FROM AUTHOR]

Abstract: Increasing atmospheric [CO2] and temperature are expected to affect the productivity, species composition, biogeochemistry, and therefore the quantity and quality of forage available to herbivores in rangeland ecosystems. Both elevated CO2 (eCO2) and warming affect plant tissue chemistry through multiple direct and indirect pathways, such that the cumulative outcomes of these effects are difficult to predict. Here, we report on a 7‐yr study examining effects of CO2 enrichment (to 600 ppm) and infrared warming (+1.5°C day/3°C night) under realistic field conditions on forage quality and quantity in a semiarid, mixedgrass prairie. For the three dominant forage grasses, warming effects on in vitro dry matter digestibility (IVDMD) and tissue [N] were detected only in certain years, varied from negative to positive, and were relatively minor. In contrast, eCO2 substantially reduced IVDMD (two most abundant grasses) and [N] (all three dominant grass species) in most years, except the two wettest years. Furthermore, eCO2 reduced IVDMD and [N] independent of warming effects. Reduced IVDMD with eCO2 was related both to reduced [N] and increased acid detergent fiber (ADF) content of grass tissues. For the six most abundant forage species (representing 96% of total forage production), combined warming and eCO2 increased forage production by 38% and reduced forage [N] by 13% relative to ambient climate. Although the absolute magnitude of the decline in IVDMD and [N] due to combined warming and eCO2 may seem small (e.g., from 63.3 to 61.1% IVDMD and 1.25 to 1.04% [N] for Pascopyrum smithii), such shifts could have substantial consequences for the rate at which ruminants gain weight during the primary growing season in the largest remaining rangeland ecosystem in North America. With forage production increases, declining forage quality could potentially be mitigated by adaptively increasing stocking rates, and through management such as prescribed burning, fertilization at low rates, and legume interseeding to enhance forage quality. [ABSTRACT FROM AUTHOR]

Nutrient-rich organic waste derived from sewage treatment facilities or livestock manure is often applied to rangelands of western North America to increase soil fertility, forage production, forage quality, and soil carbon storage. This practice can have a number of undesirable side effects, however, including plant invasion. While characteristics of both rangeland ecosystems and invasive plants suggest that organic waste application might often promote invasion, results to date are mixed, perhaps due in part to the paucity of long-term studies. Here, we describe the long-term (22 yr) effects of three types of organic waste-composted biosolids, composted cattle manure, and fresh cattle manure-on plant productivity and invasion in native mixed-grass prairie. Although composted manure and biosolids increased plant productivity and forage quality in the second year of the study, these effects did not persist. In contrast, Bromus tectorum (cheatgrass), which invaded the study site over the course of the experiment, was strongly facilitated by composted manure addition, with particularly large effects observed in years 17 and 22. These results show that nutrient-rich organic waste can favor invasive species even in the relatively invasion- resistant grasslands of the western Great Plains. They also demonstrate that effects of organic waste on invasion can become apparent quite gradually and persist for decades following the initial organic waste application. Together, results from experimental additions of organic and inorganic nutrients to native rangelands suggest that the risk of promoting invasive species is significant, and should be considered in programs that apply organic waste to rangelands of western North America. [ABSTRACT FROM AUTHOR]

Climate controls vegetation distribution across the globe, and some vegetation types are more vulnerable to climate change, whereas others are more resistant. Because resistance and resilience can influence ecosystem stability and determine how communities and ecosystems respond to climate change, we need to evaluate the potential for resistance as we predict future ecosystem function. In a mixed-grass prairie in the northern Great Plains, we used a large field experiment to test the effects of elevated CO2, warming, and summer irrigation on plant community structure and productivity, linking changes in both to stability in plant community composition and biomass production. We show that the independent effects of CO2 and warming on community composition and productivity depend on interannual variation in precipitation and that the effects of elevated CO2 are not limited to water saving because they differ from those of irrigation. We also show that production in this mixed-grass prairie ecosystem is not only relatively resistant to interannual variation in precipitation, but also rendered more stable under elevated CO2 conditions. This increase in production stability is the result of altered community dominance patterns: Community evenness increases as dominant species decrease in biomass under elevated CO2. In many grasslands that serve as rangelands, the economic value of the ecosystem is largely dependent on plant community composition and the relative abundance of key forage species. Thus, our results have implications for how we manage native grasslands in the face of changing climate. [ABSTRACT FROM AUTHOR]

As global changes reorganize plant communities, invasive plants may benefit. We hypothesized that elevated CO2 and warming would strongly influence invasive species success in a semi-arid grassland, as a result of both direct and water-mediated indirect effects., To test this hypothesis, we transplanted the invasive forb Linaria dalmatica into mixed-grass prairie treated with free-air CO2 enrichment and infrared warming, and followed survival, growth, and reproduction over 4 yr. We also measured leaf gas exchange and carbon isotopic composition in L. dalmatica and the dominant native C3 grass Pascopyrum smithii., CO2 enrichment increased L. dalmatica biomass 13-fold, seed production 32-fold, and clonal expansion seven-fold, while warming had little effect on L. dalmatica biomass or reproduction. Elevated CO2 decreased stomatal conductance in P. smithii, contributing to higher soil water, but not in L. dalmatica. Elevated CO2 also strongly increased L. dalmatica photosynthesis (87% versus 23% in P. smithii), as a result of both enhanced carbon supply and increased soil water., More broadly, rapid growth and less conservative water use may allow invasive species to take advantage of both carbon fertilization and water savings under elevated CO2. Water-limited ecosystems may therefore be particularly vulnerable to invasion as CO2 increases. [ABSTRACT FROM AUTHOR]

In semiarid western North American riparian ecosystems, increased drought and lower streamflows under climate change may reduce plant growth and recruitment, and favor drought-tolerant exotic species over mesic native species. We tested whether elevated atmospheric CO2 might ameliorate these effects by improving plant water-use efficiency., We examined the effects of CO2 and water availability on seedlings of two native ( Populus deltoides spp. monilifera, Salix exigua) and three exotic ( Elaeagnus angustifolia, Tamarix spp., Ulmus pumila) western North American riparian species in a CO2-controlled glasshouse, using 1-m-deep pots with different water-table decline rates., Low water availability reduced seedling biomass by 70-97%, and hindered the native species more than the exotics. Elevated CO2 increased biomass by 15%, with similar effects on natives and exotics. Elevated CO2 increased intrinsic water-use efficiency (Δ13Cleaf), but did not increase biomass more in drier treatments than wetter treatments., The moderate positive effects of elevated CO2 on riparian seedlings are unlikely to counteract the large negative effects of increased aridity projected under climate change. Our results suggest that increased aridity will reduce riparian seedling growth despite elevated CO2, and will reduce growth more for native Salix and Populus than for drought-tolerant exotic species. [ABSTRACT FROM AUTHOR]

1. It has been suggested that much of the elevated CO2 effect on plant productivity and N cycling in semi-arid grasslands is related to a CO2-induced increase in soil moisture, but the relative importance of moisture-mediated and direct effects of CO2 remain unclear. 2. We grew five grassland species common to the semi-arid grasslands of northern Colorado, USA, as monocultures and as mixtures of all five species in pots. We examined the effects of atmospheric CO2 concentration (ambient vs. 780 p.p.m.) and soil moisture (15 vs. 20% m/m) on plant biomass and plant N uptake. Our objective was to separate CO2 effects not related to water from water-mediated CO2 effects by frequently watering the pots, thereby eliminating most of the elevated CO2 effects on soil moisture, and including a water treatment similar in magnitude to the water-savings effect of CO2. 3. Biomass of the C3 grasses Hesperostipa comata and Pascopyrum smithii increased under elevated CO2, biomass of the C4 grass Bouteloua gracilis increased with increased soil moisture, while biomass of the forbs Artemisia frigida and Linaria dalmatica had no or mixed responses. Increased plant N uptake contributed to the increase in plant biomass with increased soil moisture while the increase in plant biomass with CO2 enrichment was mostly a result of increased N use efficiency (NUE). Species-specific responses to elevated CO2 and increased soil moisture differed between monocultures and mixtures. Both under elevated CO2 and with increased soil moisture, certain species gained N in mixtures at the expense of species that lost N, but elevated CO2 led to a different set of winners and losers than did increased water. 4. Elevated CO2 can directly increase plant productivity of semi-arid grasslands through increased NUE, while a CO2-induced increase in soil moisture stimulating net N mineralization could further enhance plant productivity through increased N uptake. Our results further indicate that the largest positive and negative effects of elevated CO2 and increased soil moisture on plant productivity occur with interspecific competition. Responses of this grassland community to elevated CO2 and water may be both contingent upon and accentuated by competition. [ABSTRACT FROM AUTHOR]

Elevated CO2 affects plant productivity, but also water availability and plant species composition in semi-arid grasslands, thereby potentially causing complex effects on CH4 consumption and N2O emission. We studied the effects of atmospheric CO2 concentration (400 vs 780 μL L−1), water content (15 vs 20% gravimetric soil moisture), and composition of semi-arid grassland species (perennial grasses Bouteloua gracilis, Hesperostipa comata, and Pascopyrum smithii; sub-shrub Artemisia frigida; invasive forb Linaria dalmatica grown in monoculture and all five species together) on CH4 consumption and N2O emission in a full factorial greenhouse experiment. We used a unique method where we measured microbial effects on CH4 consumption and N2O emission in isolation from effects of gas diffusivity. Microbially mediated CH4 consumption was significantly higher under elevated CO2 (by 20%), but was not affected by soil water content or plant species composition. Microbially mediated N2O emission was not significantly affected by elevated CO2, but was significantly higher with high water content (by 67%) and differed significantly among species. Treatment effects on CH4 consumption and N2O emission often could not be explained simply by differences in soil moisture, suggesting that treatment-induced changes in other soil and microbial properties played a role in causing these effects. [ABSTRACT FROM AUTHOR]

Increasing atmospheric CO2 has potentially significant impacts on the dynamics of water use and conservation in semi-arid rangelands. In this study we used large (15.5 m2) open top chambers to investigate effects of twice ambient CO2 concentration (720 μL L-1) on plant and soil water relations of semi-arid shortgrass steppe (SGS) of northeastern Colorado from 1997 to 2001. Seasonal average soil moisture throughout the soil profile (0–15, 15–45, 45–75, 75–105 cm) was increased under elevated CO2 compared to ambient CO2 for much of the study period. When averaged across years, the greatest relative increase (elevated vs. ambient) in soil moisture occurred in the 75–105 cm depth increment (16.4%). Averaged over the study period, leaf water potential (Ψleaf) was enhanced 24–30% under elevated CO2 in the major warm- and cool-season grass species of the SGS (Bouteloua gracilis, C4, 28.5%; Pascopyrum smithii, C3, 24.7%; Stipa comata, C3, 30.4%), and the degree of responsiveness in Ψleaf to elevated CO2 did not differ between C3 and C4 plant functional types, but did differ between C3 species. Water-use efficiency (WUE; g aboveground biomass harvested/ kg water consumed) was 43% higher in elevated (6.10) than ambient (4.27) CO2 plots over the study period. Results suggest that a future, elevated CO2 environment may result not only in increased plant productivity due to improved WUE, but also lead to increased water drainage and deep soil moisture storage in this semi-arid grassland ecosystem. This, along with the ability of the major grass species to maintain a favorable water status under elevated CO2, should result in the SGS being less susceptible to prolonged periods of drought. However, increases in deep soil water may eventually favor deeper-rooted over shallow-rooted species. [ABSTRACT FROM AUTHOR]

The article presents a study which aims to explore the CO2 response of the shortgrass steppe used for livestock grazing in the North American Great Plains. The study uses three with ambient air and three with air CO2 chambers as well as three unchambered controls to evaluate responses to rising atmospheric CO2. It also suggests the enhancement of the rising atmospheric CO2 production of lower quality forage along with a species composition shift.

Increasing atmospheric [CO 2 ] and temperature are expected to affect the productivity, species composition, biogeochemistry, and therefore the quantity and quality of forage available to herbivores in rangeland ecosystems. Both elevated CO 2 (eCO 2 ) and warming affect plant tissue chemistry through multiple direct and indirect pathways, such that the cumulative outcomes of these effects are difficult to predict. Here, we report on a 7-yr study examining effects of CO 2 enrichment (to 600 ppm) and infrared warming (+1.5°C day/3°C night) under realistic field conditions on forage quality and quantity in a semiarid, mixedgrass prairie. For the three dominant forage grasses, warming effects on in vitro dry matter digestibility (IVDMD) and tissue [N] were detected only in certain years, varied from negative to positive, and were relatively minor. In contrast, eCO 2 substantially reduced IVDMD (two most abundant grasses) and [N] (all three dominant grass species) in most years, except the two wettest years. Furthermore, eCO 2 reduced IVDMD and [N] independent of warming effects. Reduced IVDMD with eCO 2 was related both to reduced [N] and increased acid detergent fiber (ADF) content of grass tissues. For the six most abundant forage species (representing 96% of total forage production), combined warming and eCO 2 increased forage production by 38% and reduced forage [N] by 13% relative to ambient climate. Although the absolute magnitude of the decline in IVDMD and [N] due to combined warming and eCO 2 may seem small (e.g., from 63.3 to 61.1% IVDMD and 1.25 to 1.04% [N] for Pascopyrum smithii), such shifts could have substantial consequences for the rate at which ruminants gain weight during the primary growing season in the largest remaining rangeland ecosystem in North America. With forage production increases, declining forage quality could potentially be mitigated by adaptively increasing stocking rates, and through management such as prescribed burning, fertilization at low rates, and legume interseeding to enhance forage quality., (© 2018 by the Ecological Society of America.)