Effects of climate and snow depth on Bromus tectorum population dynamics at high elevation

Invasive plants are thought to be especially capable of range shifts or expansion in response to climate change due to high dispersal and colonization abilities. Although highly invasive throughout the Intermountain West, the presence and impact of the grass Bromus tectorum has been limited at higher elevations in the eastern Sierra Nevada, potentially due to extreme wintertime conditions. However, climate models project an upward elevational shift of climate regimes in the Sierra Nevada that could favor B. tectorum expansion. This research specifically examined the effects of experimental snow depth manipulations and interannual climate variability over 5 years on B. tectorum populations at high elevation (2,175 m). Experimentally-increased snow depth had an effect on phenology and biomass, but no effect on individual fecundity. Instead an experimentally-increased snowpack inhibited population growth in 1 year by reducing seedling emergence and early survival. A similar negative effect of increased snow was observed 2 years later. However, a strong negative effect on B. tectorum was also associated with a naturally low-snow winter, when seedling emergence was reduced by 86%. Across 5 years, winters with greater snow cover and a slower accumulation of degree-days coincided with higher B. tectorum seedling density and population growth. Thus, we observed negative effects associated with both experimentally-increased and naturally-decreased snowpacks. It is likely that the effect of snow at high elevation is nonlinear and differs from lower elevations where wintertime germination can be favorable. Additionally, we observed a doubling of population size in 1 year, which is alarming at this elevation.     

Winter cold tolerance and the geographic range separation of Bromus tectorum and Bromus rubens,two severe invasive species in North America

The invasive grasses Bromus rubens and Bromus tectorum are responsible for widespread damage to semiarid biomes of western North America. Bromus. tectorum dominates higher and more northern landscapes than its sister species B. rubens, which is a severe invader in the Mojave desert region of the American Southwest. To assess climate thresholds controlling their distinct geographic ranges, we evaluated the winter cold tolerance of B. tectorum and B. rubens. Freezing tolerance thresholds were determined using electrolyte leakage and whole‐plant mortality. The responses of the two species to winter cold and artificial freezing treatments were similar in 2007–2008 and 2009–2010. When grown at minimum temperatures of 10 °C, plants of both species had cold tolerance thresholds near ?10 °C, while plants acclimated to a daily minimum of ?10 to ?30 °C survived temperatures down to ?31 °C. In the winter of 2010–2011, a sudden severe cold event on December 9, 2010 killed all B. rubens populations, while B. tectorum was not harmed; all tested plants were 7–8 weeks old. Controlled acclimation experiments demonstrated that 8‐week‐old plants of B. rubens had a slower acclimation rate to subzero temperatures than B. tectorum and could not survive a rapid temperature drop from 1 to ?14 °C. Four‐month‐old B. rubens populations were as cold tolerant as B. tectorum. Our results show that severe and sudden freeze events in late autumn can kill young plants of B. rubens but not B. tectorum. Such events could exclude B. rubens from the relatively cold, Intermountain steppe biome of western North America where B. tectorum predominates.     

Performance of Bromus tectorum L. in relation to soil properties, water additions, and chemical amendments in calcareous soils of southeastern Utah, USA

In drylands of southeastern Utah, USA, the invasive exotic grass Bromus tectorum L. occurs in distinct spatial patterns suggesting soil control of ecosystem susceptibility to invasion. To improve our understanding of these patterns, we examined performance of B. tectorum in relation to additions of water, KCl, MgO, and CaO at seventeen 1600 m² sites distributed across a calcareous soil gradient in Canyonlands National Park. Water additions resulted in a 57% increase in B. tectorum establishment. Fall establishment was significantly correlated with silt and clay content in wet plots but not in dry plots, suggesting that texture effects on B. tectorum establishment patterns may be greater in wet years than in dry years. Applications of MgO resulted in a 49% decrease in B. tectorum establishment, although MgO had no effect on whole-plot biomass at the end of the growing season. B. tectorum–soil relations were strongest during winter (December–March) when relative growth rates were negatively related to soil acid-neutralizing potential, sand and CaCO₃ content, and a measure of bioavailable Mg; and positively related to silt and clay content, total N, measures of bioavailable Mn, P, and K, and a measure of magnetite indicating distributional patterns of eolian dust. As soils were persistently moist during this period, we attribute strong B. tectorum–soil patterns in winter to effects of low temperature on diffusion, microbial activity, and/or production of root exudates important for nutrient mobilization and uptake. In spring, there was a reversal in B. tectorum–soil relations such that loamy soils with higher B. tectorum densities were unfavorable for growth relative to sandy soils with higher warm-season water potentials. We conclude that resource limitations for B. tectorum in this study area shift seasonally, from water limitation of fall establishment, to nutrient limitation of winter growth, and back to water limitation of spring growth. Because study sites generally were arrayed along a hillslope gradient with downslope trends in soil vtexture and nutrient content, close B. tectorum–soil relations documented in this study indicate that a geomorphic framework is useful for understanding and predicting B. tectorum invasion patterns in dryland ecosystems of this region.Electronic supplementary material Electronic supplementary material is available for this article at and accessible for authorised users. Section Editor: T. KalaposThe U.S. Government’s right to retain a non-exclusive, royalty free licence in and to any copyright is acknowledged     

The consequences of multiple resource shifts on the productivity and composition of alpine tundra communities: inferences from a long-term snow and nutrient manipulation experiment

Background: Gradients in the amounts and duration of snowpack and resulting soil moisture gradients have been associated with different plant communities across alpine landscapes.

Aims: The extent to which snow additions could alter plant community structure, both alone and in combination with nitrogen (N) and phosphorus (P) additions, provided an empirical assessment of the strength of these variables on structuring the plant communities of the alpine tundra at Niwot Ridge, Colorado Front Range.

Methods: A long-term snow fence was used to study vegetation changes in responses to snowpack, both alone and in conjunction with nutrient amendments, in plots established in dry and moist meadow communities in the alpine belt. Species richness, diversity, evenness and dissimilarity were evaluated after 20 years of treatments.

Results: Snow additions, alone, reduced species richness and altered species composition in dry meadow plots, but not in moist meadow; more plant species were found in the snow-impacted areas than in nearby controls. Changes in plant community structure to N and N + P additions were influenced by snow additions. Above-ground plant productivity in plots not naturally affected by snow accumulation was not increased, and the positive responses of plant species to nutrient additions were reduced by snow addition. Plant species showed individualistic responses to changes in snow and nutrients, and indirect evidence suggested that competitive interactions mediated responses. A Permanova analysis demonstrated that community dissimilarity was affected by snow, N, and P additions, but with these responses differing by community type for snow and N. Snow influenced community patterns generated by N, and finally, the communities impacted by N + P were significantly different than those affected by the individual nutrients.

Conclusions: These results show that changes in snow cover over a 20-year interval produce measureable changes in community composition that concurrently influence and are influenced by soil nutrient availability. Dry meadow communities exhibit more sensitivity to increases in snow cover whereas moist meadow communities appear more sensitive to N enrichment. This study shows that the dynamics of multiple limiting resources influence both the productivity and composition of alpine plant communities, with, species, life form, and functional traits mediating these responses.     

Nitrogen deposition enhances Bromus tectorum invasion: biogeographic differences in growth and competitive ability between China and North America

Increased resource supply commonly facilitates invasion by exotic plants, raising concerns over atmospheric nitrogen (N) deposition; fast‐growing annual invaders may have exceptional abilities to outperform native perennials in response to N pulses. However, it remains unclear whether this advantage is due to growth differences or to shifts in competitive outcomes, and whether annual invaders are favored by N deposition in their introduced range over native range. We conducted an experiment to compare the growth and competitive ability of Bromus tectorum and its native perennial grasses either at three different N regimes or between China and North America. The soil used in this experiment was from mountain grasslands as a neutral growth medium. The total biomass of three natives from China and North America did not increase along the N deposition gradient. Nitrogen addition enhanced the growth of North American B. tectorum instead of Chinese B. tectorum. Nitrogen addition increased the competitive ability of B. tectorum, but had no effect on that of natives. North American B. tectorum was bigger and had greater competitive ability and root weight ratio than Chinese B. tectorum. In contrast, North American natives were less competitive than Chinese natives. There was a significantly positive correlation between the growth of B. tectorum grown alone and its competitive ability. These findings suggest that N deposition may enhance the B. tectorum invasion through disproportionally increasing the growth and maintaining inherent competitive advantages of North American B. tectorum, further increasing threats to introduced ranges. There were differences in the growth and competitive ability of B. tectorum and natives between China and North America, which explains why B. tectorum is a minor component at home and becomes a successful invader abroad.     

Fire, native species, and soil resource interactions influence the spatio-temporal invasion pattern of Bromus tectorum

Bromus tectorum (cheatgrass) is an invasive annual that occupies perennial grass and shrub communities throughout the western United States. Bronus tectorum exhibits an intriguing spatio‐temporal pattern of invasion in low elevation ponderosa pine Pinus ponderosa/bunchgrass communities in western Montana where it forms dense rings beneath solitary pines following fire. This pattern provides a unique opportunity to investigate several indirect effects of native vegetation that influence the invasion pattern of B. tectorum, and specifically how native species, disturbance, and soil resources interact to influence the spatio‐temporal pattern of invasion. We established four replicate field sites, each containing burned‐tree, burned‐grass, unburned‐tree, and unburned‐grass sampling locations, and initiated a series of field sampling and greenhouse experiments utilizing these locations. The objective of our first greenhouse experiment was to identify whether belowground factors contributed to the pattern of B. tectorum biomass observed in these field locations. This experiment generated a B. tectorum biomass response that was nearly identical to the invasion pattern observed in the field, suggesting further investigation of belowground factors was necessary. We measured resin‐sorbed NH₄⁺ and NO₃⁻ during one generation of B. tectorum, and measured a suite of P fractions through a sequential extraction procedure from these soils. These data revealed that a resource island of high N and P exists beneath pine trees. Through a second greenhouse experiment, we determined that N limited B. tectorum biomass in tree soil, whereas P limited biomass in bunchgrass soil. Finally, through a germination experiment we determined that pine litter strongly inhibited B. tectorum germination. These data suggest B. tectorum is regulated by P in bunchgrass soil, and by N and inhibition by pine litter beneath trees, effects that are likely alleviated by fire. These data demonstrate the combined role of direct and indirect interactions between native and invasive species in regulating biological invasions.     

Cheatgrass is favored by warming but not CO2 enrichment in a semi‐arid grassland

Elevated CO₂ and warming may alter terrestrial ecosystems by promoting invasive plants with strong community and ecosystem impacts. Invasive plant responses to elevated CO₂ and warming are difficult to predict, however, because of the many mechanisms involved, including modification of phenology, physiology, and cycling of nitrogen and water. Understanding the relative and interactive importance of these processes requires multifactor experiments under realistic field conditions. Here, we test how free‐air CO₂ enrichment (to 600 ppmv) and infrared warming (+1.5 °C day/3 °C night) influence a functionally and phenologically distinct invasive plant in semi‐arid mixed‐grass prairie. Bromus tectorum (cheatgrass), a fast‐growing Eurasian winter annual grass, increases fire frequency and reduces biological diversity across millions of hectares in western North America. Across 2 years, we found that warming more than tripled B. tectorum biomass and seed production, due to a combination of increased recruitment and increased growth. These results were observed with and without competition from native species, under wet and dry conditions (corresponding with tenfold differences in B. tectorum biomass), and despite the fact that warming reduced soil water. In contrast, elevated CO₂ had little effect on B. tectorum invasion or soil water, while reducing soil and plant nitrogen (N). We conclude that (1) warming may expand B. tectorum's phenological niche, allowing it to more successfully colonize the extensive, invasion‐resistant northern mixed‐grass prairie, and (2) in ecosystems where elevated CO₂ decreases N availability, CO₂ may have limited effects on B. tectorum and other nitrophilic invasive species.     

Effects of resource availability and propagule supply on native species recruitment in sagebrush ecosystems invaded by Bromus tectorum

Resource availability and propagule supply are major factors influencing establishment and persistence of both native and invasive species. Increased soil nitrogen (N) availability and high propagule inputs contribute to the ability of annual invasive grasses to dominate disturbed ecosystems. Nitrogen reduction through carbon (C) additions can potentially immobilize soil N and reduce the competitiveness of annual invasive grasses. Native perennial species are more tolerant of resource limiting conditions and may benefit if N reduction decreases the competitive advantage of annual invaders and if sufficient propagules are available for their establishment. Bromus tectorum, an exotic annual grass in the sagebrush steppe of western North America, is rapidly displacing native plant species and causing widespread changes in ecosystem processes. We tested whether nitrogen reduction would negatively affect B. tectorum while creating an opportunity for establishment of native perennial species. A C source, sucrose, was added to the soil, and then plots were seeded with different densities of both B. tectorum (0, 150, 300, 600, and 1,200 viable seeds m⁻²) and native species (0, 150, 300, and 600 viable seeds m⁻²). Adding sucrose had short-term (1 year) negative effects on available nitrogen and B. tectorum density, biomass and seed numbers, but did not increase establishment of native species. Increasing propagule availability increased both B. tectorum and native species establishment. Effects of B. tectorum on native species were density dependent and native establishment increased as B. tectorum propagule availability decreased. Survival of native seedlings was low indicating that recruitment is governed by the seedling stage.     

Evaluating dominance as a component of non-native species invasions

Many studies have quantified plant invasions by determining patterns of non‐native species establishment (i.e. richness and absolute cover). Until recently, dominance has been largely overlooked as a significant component of invasion. Therefore, we re‐examined a 6‐year data set of 323 0.1 ha plots within 18 vegetation types collected in the Grand Staircase‐Escalante National Monument from 1998 to 2003, including dominance (i.e. relative cover) in our analyses. We specifically focused on the non‐native species Bromus tectorum, a notable dominant annual grass in this system. We found that non‐native species establishment and dominance are both occurring in species‐rich, mesic vegetation types. Therefore, non‐native species dominance may result despite many equally abundant native species rather than a dominant few, and competitive exclusion does not seem to be a primary control on either non‐native species establishment or dominance in this study. Unlike patterns observed for non‐native species establishment, relative non‐native species cover could not be predicted by native species richness across vegetation types (R² < 0.001; P = 0.45). However, non‐native species richness was found to be positively correlated with relative non‐native species cover and relative B. tectorum cover (R² = 0.46, P < 0.01; R² = 0.17, P < 0.01). Analyses within vegetation types revealed predominantly positive relationships among these variables for the correlations that were significant. Regression tree analyses across vegetation types that included additional biotic and abiotic variables were a little better at predicting non‐native species dominance (PRE = 0.49) and B. tectorum dominance (PRE = 0.39) than at predicting establishment. Land managers will need to set priorities for control efforts on the more productive, species‐rich vegetation types that appear to be susceptible to both components of invasion.     

Established native perennial grasses out-compete an invasive annual grass regardless of soil water and nutrient availability

Competition and resource availability influence invasions into native perennial grasslands by non-native annual grasses such as Bromus tectorum. In two greenhouse experiments we examined the influence of competition, water availability, and elevated nitrogen (N) and phosphorus (P) availability on growth and reproduction of the invasive annual grass B. tectorum and two native perennial grasses (Elymus elymoides, Pascopyrum smithii). Bromus tectorum aboveground biomass and seed production were significantly reduced when grown with one or more established native perennial grasses. Conversely, average seed weight and germination were significantly lower in the B. tectorum monoculture than in competition native perennial grasses. Intraspecific competition reduced per-plant production of both established native grasses, whereas interspecific competition from B. tectorum increased production. Established native perennial grasses were highly competitive against B. tectorum, regardless of water, N, or P availability. Bromus tectorum reproductive potential (viable seed production) was not significantly influenced by any experimental manipulation, except for competition with P. smithii. In all cases, B. tectorum per-plant production of viable seeds exceeded parental replacement. Our results show that established plants of Elymus elymoides and Pascopyrum smithii compete successfully against B. tectorum over a wide range of soil resource availability.     

Soil water exploitation after fire: competition between Bromus tectorum (cheatgrass) and two native species

Summary Causes for the widespread abundance of the alien grass Bromus tectorum (cheatgrass) after fire in semiarid areas of western North America may include: (1) utilization of resources freed by the removal of fireintolerant plants; and (2) successful competition between B. tectorum and individual plants that survive fire. On a site in northwestern Nevada (USA), measurements of soil water content, plant water potential, aboveground biomass production, water use efficiency, and B. tectorum tiller density were used to determine if B. tectorum competes with either of two native species (Stipa comata and Chrysothamnus viscidiflorus) or simply uses unclaimed resources. Soil water content around native species occurring with B. tectorum was significantly lower (P<0.05) than around individuals without B. tectorum nearby. Native species had significantly more negative plant water potential when they occurred with B. tectorum. Aboveground biomass was significantly higher for native species without B. tectorum. However, the carbon isotope ratio of leaves for native species with B. tectorum was not significantly different from individuals without B. tectorum. Thus, B. tectorum competes with native species for soil water and negatively affects their wate status and productivity, but the competition for water does not affect water use efficiency of the native species. These adverse effects of B. tectorum competition on the productivity and water status of native species are also evident at 12 years after a fire. This competitive ability of B. tectorum greatly enhances its capability to exploit soil resources after fire and to enhance its status in the community.     

Effects of perennial semi-arid bunchgrass spatial patterns on performance of the invasive annual cheatgrass (Bromus tectorum L.)

Plant spatial patterns critically influence community dynamics, including plant interactions, resource distribution, and community invasibility. Research suggests that resistance of western US plant communities to further invasion by the exotic annual grass Bromus tectorum may be linked to the positions of, and spacing between, perennial plants. In particular, gaps between aggregated clusters of perennial plants may facilitate B. tectorum invasion by providing safe sites for seed germination and establishment. We tested the effects of random, regular, and aggregated bunchgrass patterns, manipulated at both community (plot) and neighborhood scales, on B. tectorum biomass and spikelet production after experimental seed addition. We found strong evidence of treatment effects on both biomass and spikelets, which varied between treatments by approximately 2.5-fold. Mean biomass and spikelet counts were lowest in plots in which bunchgrasses were aggregated at both community and neighborhood scales, likely due to the increased competition. Although not statistically distinguishable from most other treatments, B. tectorum biomass and spikelet counts were highest in plots with bunchgrass patterns that were random at the community scale and aggregated at the neighborhood scale. These plots were characterized by relatively large gaps between bunchgrass clusters, suggesting that B. tectorum may exploit gaps between aggregated perennial plants. Our results support the emerging hypothesis that community resistance to B. tectorum invasion could be increased through manipulation of perennial vegetation to reduce basal gap size and connectivity.     

Contrasting effects of resource availability and plant mortality on plant community invasion by Bromus tectorum L.

The positive effect of disturbance on plant community invasibility is one of the more consistent results in invasion ecology. It is generally attributed to a coincident increase in available resources (due to the disturbance) that allows non-resident plant species to establish (Davis MA, Grime JP Thompson K, J Ecol 88:528–534, 2000). However, most research addressing this issue has been in artificial or highly modified plant communities. Our goal in this study was to investigate the interactive effects of resource availability and plant mortality disturbance on the invasion of natural plant communities. We conducted a series of experiments that examined the response of Bromus tectorum L., a highly invasive annual grass, to experimentally created gradients of resource availability [nitrogen (N) and water] and resident plant species mortality. We found that B. tectorum biomass was co-limited by N and water. Biomass at the end of the growing season was a saturating function (i.e., increased to a maximum) of water, which determined maximum biomass, and N, which determined the rate at which maximum biomass was attained. Despite that fact that plant mortality increased N availability, it had a negative impact on invasion success. Plant mortality also decreased foliar cover, standing dead biomass, and soil cover by litter. In harsh environments, removing foliar and soil cover may increase germination and seedling stress by increasing soil temperatures and water loss. Across all treatments, B. tectorum success decreased with decreasing foliar cover and standing dead biomass. This, in combination with the strong limitation of B. tectorum biomass by water in this experiment, suggests that our plant mortality disturbance removed soil cover that may have otherwise aided B. tectorum invasion into this semi-arid plant community by reducing water stress.     

Plant phenology and life span influence soil pool dynamics: Bromus tectorum invasion of perennial C3–C4 grass communities

In water-limited ecosystems, small rainfall events can have dramatic impacts on microbial activity and soil nutrient pools. Plant community phenology and life span also affect soil resources by determining the timing and quantity of plant nutrient uptake, storage, and release. Using the replacement of C₃–C₄ perennial grasses by the invasive annual grass Bromus tectorum as a case study, we investigated the influence of phenology and life span on pulse responses and sizes of soil carbon (C) and nitrogen (N) pools. We hypothesized that available and microbial C and N would respond to small rainfall events and that B. tectorum invasion would increase soil C and N pools by reducing inter-annual plant C and N storage and alter seasonal pool dynamics by changing the timing of plant uptake and litter inputs. We tested our hypotheses by simulating small rainfall events in B. tectorum and perennial grass communities three times during the growing season. Microbial pools responded strongly to soil moisture and simulated rainfall events, but labile C and N pools were affected weakly or not at all. All pools were larger beneath B. tectorum than perennial grasses. Soil C and N pools increased after senescence in both communities. Our results suggest that transforming a perennial into a B. tectorum dominated community increases the overall size of soil C and N pools by decreasing plant C and N storage and changes seasonal pool dynamics by altering dominant plant phenology. Our results indicate strong roles for water, life span and phenology in controlling soil C and N pools and begin to elucidate the biogeochemical effects of altering plant community phenology and life span.     

Emerging climate‐driven disturbance processes: widespread mortality associated with snow‐to‐rain transitions across 10° of latitude and half the range of a climate‐threatened conifer

Climate change is causing rapid changes to forest disturbance regimes worldwide. While the consequences of climate change for existing disturbance processes, like fires, are relatively well studied, emerging drivers of disturbance such as snow loss and subsequent mortality are much less documented. As the climate warms, a transition from winter snow to rain in high latitudes will cause significant changes in environmental conditions such as soil temperatures, historically buffered by snow cover. The Pacific coast of North America is an excellent test case, as mean winter temperatures are currently at the snow–rain threshold and have been warming for approximately 100 years post‐Little Ice Age. Increased mortality in a widespread tree species in the region has been linked to warmer winters and snow loss. Here, we present the first high‐resolution range map of this climate‐sensitive species, Callitropsis nootkatensis (yellow‐cedar), and document the magnitude and location of observed mortality across Canada and the United States. Snow cover loss related mortality spans approximately 10° latitude (half the native range of the species) and 7% of the overall species range and appears linked to this snow–rain transition across its range. Mortality is commonly >70% of basal area in affected areas, and more common where mean winter temperatures is at or above the snow–rain threshold (>0 °C mean winter temperature). Approximately 50% of areas with a currently suitable climate for the species (相似文献   

Photosynthetic and phenological responses of dwarf shrubs to the depth and properties of snow

Snow is known to have a major impact on vegetation in arctic ecosystems, but little is known about how snow affects plants in boreal forests, where the snowpack is uneven due to canopy impact. The responses of two dwarf shrubs, the evergreen Vaccinium vitis‐idaea and the deciduous V. myrtillus, to snow conditions were studied in a snow manipulation experiment in southern Finland. The thermal insulation of the snowpack was expected to decrease with partial removal or compression of the snow, while addition of snow was expected to have the opposite effect. The penetration of light was manipulated by partial removal of snow or by formation of an artificial ice layer in the snowpack. CO₂ exchange measurements that were carried out at the time of maximum snow depth in late March indicated significant photosynthetic activity in the leaves of V. vitis‐idaea under snow. Net gain of CO₂ was observed in the daytime on all the manipulation plots, excluding the snow addition plots, where light intensity was very low. The subnivean photosynthesis compensated for a substantial proportion (up to 80%) of the respiratory CO₂ losses. Chlorophyll fluorescence measurements indicated reduced potential capacity of photosystem II in the leaves of V. vitis‐idaea on those plots where snow cover was thin. Neither V. vitis‐idaea nor V. myrtillus suffered from frost damage (assessed as electrolyte leakage) when thermal insulation was reduced by means of snow manipulations. No phenological responses were observed in V. vitis‐idaea, but in V. myrtillus bud burst, leaf unfolding and flowering were advanced by 1–3 days on the addition plots. The results of the present study show that dwarf shrubs respond to not only the thickness of snow but also the physical properties of snow, both of which are expected to change due to climatic warming.     

Sources of nitrate export during rain-on-snow events at forested catchments

Rain-on-snow (ROS) events are major drivers of nitrate (NO₃-N) export from seasonally snow-covered forested catchments and may cause episodic declines in stream pH. High intensity monitoring of throughfall, snow pack and stream water draining two proximal catchments (Harp 3A and Harp 6A) with very different NO₃-N export revealed that a very small percentage of ROS-induced stream discharge originates from throughfall and melting snow (new water; average = 6.4 %). However, this new water has a very high concentration of NO₃-N (throughfall/snowmelt average = 498 μg/L) compared with baseflow (average = 7.3 μg/L in Harp 6A; average = 41 μg/L in Harp 3A) and as a result, throughfall and snowmelt contribute the majority of NO₃-N export (average = 62 %) during ROS events. In contrast, concentrations of sulphate, dissolved organic carbon and calcium in rain and snowpack are similar to baseflow and therefore ROS-induced declines in pH (often to below pH 6.0) are attributed entirely to increases in NO₃-N concentration. Differences in absolute magnitude of ROS NO₃-N export between catchments are explained through differences in baseflow NO₃-N concentrations. The frequency and magnitude of ROS events in this region are affected by both NO₃-N deposition and winter temperature, and thus the impact of these events in the future depends on changes in both atmospheric deposition and winter climate.     

Controls over invasion of Bromus tectorum: The importance of climate,soil, disturbance and seed availability

Question: Predicting the future abundance and distribution of invasive plants requires knowing how they respond to environmental conditions. In arid and semi‐arid ecosystems where water is a limiting resource, environmental conditions and disturbance patterns influence invasions by altering acquisition and utilization of water over space and time. We ask: 1. How do variations in climatic and soil properties influence temporal soil water dynamics? 2. How does this variation affect the establishment of Bromus tectorum (cheatgrass), a cool‐season annual grass that has successfully colonized much of the U.S. Great Basin? Location: Short‐grass Steppe in northeastern Colorado, USA; Arid Lands Ecology reserve in southeastern Washington, USA; and the Patagonian steppe of the Chubut province in Argentina. Methods: We utilized a soil water model to simulate seasonal soil water dynamics in multiple combinations of climatic and soil properties. In addition, we utilized a gap dynamics model to simulate the impact of disturbance regime and seed availability on competition between B. tectorum and native plants. Results: Our results suggest that climate is very important, but that soil properties do not significantly influence the probability of observing conditions suitable for B. tectorum establishment. Results of the plant competition model indicate that frequent disturbance causes more Bromus tectorum in invaded areas and higher seed availability causes faster invasion. Conclusions: These results imply a general framework for understanding Bromus tectorum invasion in which climatic conditions dictate which areas are susceptible to invasion, disturbance regime dictates the severity of invasion and seed availability dictates the speed of invasion.     

The effects of snow-N deposition and snowmelt dynamics on soil-N cycling in marginal terraced grasslands in the French Alps

Atmospheric nitrogen (N) deposition increasingly impacts remote ecosystems. At high altitudes, snow is a key carrier of water and nutrients from the atmosphere to the soil. Medium-sized subalpine grassland terraces are characteristic of agricultural landscapes in the French Alps and influence spatial and temporal snow pack variables. At the Lautaret Pass, we investigated snow and soil characteristics along mesotopographic gradients across the terraces before and during snowmelt. Total N concentrations in the snowpack did not vary spatially and were dominated by organic N forms either brought by dry deposition trapped by the snow, or due to snow-microbial immobilization and turnover. As expected, snowpack depth, total N deposited with snow and snowmelt followed the terrace toposequence; more snow-N accumulated towards the bank over longer periods. However, direct effects of snow-N on soil-N cycling seem unlikely since the amount of nitrogen released into the soil from the snowpack was very small relative to soil-N pools and N mineralization rates. Nevertheless, some snow-N reached the soil at thaw where it underwent biotic and abiotic processes. In situ soil-N mineralization rates did not vary along the terrace toposequence but soil-N cycling was indirectly affected by the snowpack. Indeed, N mineralization responded to the snowmelt dynamic via induced temporal changes in soil characteristics (i.e. moisture and T°) which cascaded down to affect N-related microbial activities and soil pH. Soil-NH₄ and DON accumulated towards the bank during snowmelt while soil-NO₃ followed a pulse-release pattern. At the end of the snowmelt season, organic substrate limitation might be accountable for the decrease in N mineralization in general, and in NH₄ ⁺ production in particular. Possibly, during snowmelt, other biotic or abiotic processes (nitrification, denitrification, plant uptake, leaching) were involved in the transformation and transfer of snow and soil-N pools. Finally, subalpine soils at the Lautaret Pass during snowmelt experienced strong biotic and abiotic changes and switched between a source and a sink of N.     

Natural variations in snow cover do not affect the annual soil CO2 efflux from a mid‐elevation temperate forest

Climate change might alter annual snowfall patterns and modify the duration and magnitude of snow cover in temperate regions with resultant impacts on soil microclimate and soil CO₂ efflux (F_soil). We used a 5‐year time series of F_soil measurements from a mid‐elevation forest to assess the effects of naturally changing snow cover. Snow cover varied considerably in duration (105–154 days) and depth (mean snow depth 19–59 cm). Periodically shallow snow cover (<10 cm) caused soil freezing or increased variation in soil temperature. This was mostly not reflected in F_soil which tended to decrease gradually throughout winter. Progressively decreasing C substrate availability (identified by substrate induced respiration) likely over‐rid the effects of slowly changing soil temperatures and determined the overall course of F_soil. Cumulative CO₂ efflux from beneath snow cover varied between 0.46 and 0.95 t C ha^?1 yr^?1 and amounted to between 6 and 12% of the annual efflux. When compared over a fixed interval (the longest period of snow cover during the 5 years), the cumulative CO₂ efflux ranged between 0.77 and 1.18 t C ha^?1 or between 11 and 15% of the annual soil CO₂ efflux. The relative contribution (15%) was highest during the year with the shortest winter. Variations in snow cover were not reflected in the annual CO₂ efflux (7.44–8.41 t C ha^?1) which did not differ significantly between years and did not correlate with any snow parameter. Regional climate at our site was characterized by relatively high amounts of precipitation. Therefore, snow did not play a role in terms of water supply during the warm season and primarily affected cold season processes. The role of changing snow cover therefore seems rather marginal when compared to potential climate change effects on F_soil during the warm season.