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The invasion of annual grasses into historically perennial-dominated systems is a global biodiversity threat. While significant, it is not known how evolutionary changes in functional traits during the invasion process may contribute to invader success. We examined how functional traits of a widely distributed invasive annual grass medusahead (Taeniatherum caput-medusae (L.) Nevski) differed across native-range and invaded-range populations and compared these trait values to those of a common native perennial, squirreltail (Elymus elymoides (Raf.) Swezey). In greenhouse and growth chamber studies we screened a range of plant, seed and seedling functional traits and determined how variation in these traits contributed to variation in plant biomass. Our results show medusahead maintains a higher relative growth rate (RGR) than squirreltail, but the mechanism differed depending on origin with native-range medusahead achieving a higher RGR through a higher specific leaf area and the invaded-range medusahead achieving a higher RGR though a higher net assimilation rate. Contrary to previous literature, we did not find that medusahead evolved traits allowing larger biomass in its invaded range. Instead, native-range medusahead populations grew bigger than invaded-range populations, primarily due to differences in seed size. While counter to expectations, smaller seed mass may provide invaded-range medusahead populations an advantage in terms of dispersal and higher seed bank density compared to native-range populations. Thus, life history differences, altered disturbance regimes, and a shift towards production of smaller but a greater number of seeds may act, in combination, to facilitate invasion of annual grasses into perennial dominated systems. [ABSTRACT FROM AUTHOR]

Restoration of dryland ecosystems is often limited by low seedling establishment and survival. Defoliation caused by insects and small mammals could be an overlooked cause of seedling mortality. In the sagebrush steppe, we examined the effect of seedling defoliation on the survival of perennial grasses commonly used as restoration materials. Under field conditions, seedlings of three perennial bunchgrass species (non-native Agropyron cristatum , and native grasses Poa secunda and Pseudoroegneria spicata) were defoliated at two intensities (30 % and 70 % leaf length removal) and frequencies (one or two clippings) and compared to a non-defoliated control. Following emergence the first year, clippings occurred at the two-leaf stage; a second clipping occurred 1 month later for repeated defoliation treatments. We monitored seedling survival and tillering for 2 years. We expected higher defoliation intensity and frequency to reduce survival for all species, but only a few treatments reduced Po. secunda survival. Conversely, larger-statured Triticeae (wheatgrasses) benefited from some defoliation treatments. In both years, A. cristatum survival increased with repeated defoliation at both intensities. Defoliation did not affect Ps. spicata survival in the first year, but a single defoliation in the second year resulted in increased survival. In both A. cristatum and Ps. spicata , higher-intensity defoliation reduced the boost to survival resulting from defoliation frequency. Seedlings with more tillers had greater survival probabilities, but tiller number was unaffected by defoliation. Further research may elucidate mechanisms seedlings use to compensate for or benefit from defoliation. In the meantime, managers should aim to select defoliation-tolerant species if they anticipate herbivory will be problematic for restoration sites. [ABSTRACT FROM AUTHOR]

Plant reproduction is highly sensitive to stress from severe weather. While facilitation has been shown to buffer negative impacts along stress gradients, less is known about facilitating plant reproduction in drought periods. Because intensity and frequency of drought are predicted to increase, plant reproductive facilitation has important implications for a species ability to adapt to changes in climate. Our primary study objective is to test if nurse shrubs act as reproductive micro-refugia across soil types, by improving reproductive potential of perennial bunchgrass neighbors subjected to severe drought. To investigate this objective, we designed a fully factored study testing direct interaction between shrub and bunchgrasses in eastern Oregon sagebrush steppe, at two sites with different soil types. The study consisted of six simple effect treatments combining three moisture regimes (moist, ambient, and drought) with two shrub conditions (shrub intact or shrub removed). Our results indicate when facilitation of reproductive potential occurs, it occurs strongly and particularly in drought, consistent with the stress gradient hypothesis (SGH), where several species produced at least 54% more inflorescences in the presence of shrub neighbors. In addition, we found facilitation to be consistent with the SGH at the species level likely reflecting differences in plant strategy and perception of strain, but to follow alternative SGH models more closely at the site level where facilitation declined on the drier soil. Ultimately, our findings highlight the importance of facilitation in improving plant reproductive potential in drought, and support the role of nurse shrubs as micro-refugia in a changing climate. [ABSTRACT FROM AUTHOR]

Seed‐based restoration is one of the most difficult challenges for dryland restoration. Identifying environmental conditions that drive variation in seed and seedling mortality across similar restoration efforts could increase understanding of when and where restoration outcomes are likely to be favourable and identify new tools and strategies to improve outcomes.We asked how variation in a suite of environmental predictors influenced germination, emergence, seedling establishment and juvenile survival of four commonly sown perennial grass species across 33 seeding experiments distributed over an ~160,000‐km2 area of the Great Basin, a cold desert system in the western United States.Across experiments, we observed wide variation in the rates of four demographic transitions and wide variation in environmental conditions experienced by plants at each stage. For all species, higher precipitation during the first 30 days following seeding was associated with an increase in germination. Conversely, higher soil temperature over this same time period was associated with a significant decrease in germination and emergence, and soil temperature was associated with a substantial portion of the variation in germination and emergence probabilities observed across our seeding experiments.Within the range of precipitation variation observed, we were unable to detect a significant relationship between seedling establishment the first growing season and cumulative precipitation the first year, precipitation during the first spring growing season or annual climatic water deficit (CWD) the first year. Higher CWD the second growing season reduced seedling survival over that time period.Synthesis and applications. Our results show higher soil temperature negatively impacts grass seedling establishment. Restoration efforts should be prioritized when seasonal or subseasonal temperature forecasts indicate likely periods of below average temperatures. These results also indicate that climate warming will make restoration even more difficult, with our model estimates suggesting the 2°C increase in temperature expected in the Great Basin over the coming decades will decrease germination and emergence by about 30%. Lastly, while our field‐based approach provided insight into short‐term drivers of mortality, it did not provide insight into drivers of longer term survival, suggesting a need for further work on predicting long‐term restoration outcomes. [ABSTRACT FROM AUTHOR]

Background: Annual grass invasion and dominance creates a self-sustaining cycle that promotes wildfires and reduces forage abundance. Restoring native plant species to annual grass-invaded ecosystems is fundamental to fostering self-sustaining native plant communities. Aims: We sought to clarify varying restoration strategies on invasive annual and native perennial grass assembly, including strategically modified seeding times, seeding rates, and added water. Methods: We tested the effects of seeding perennial grasses in autumn, spring, or seeding half in autumn and the remaining half in spring, adding water, and varying annual and perennial grass seeding rates on annual and perennial grass life history. Results: While varying perennial grass seeding times did not affect perennial grass germination rates, annual grass germination rates were highest when perennial grasses were seeded in autumn. Seeding perennial grasses in spring produced the highest adult perennial grass density in the first-growing season, but adult perennial grass density in the second-growing season was greatest when seeding occurred in autumn. Second-growing season perennial grass density was highest where annual grass seeding rates were lowest and perennial grass seeding rates were highest. Adding water in the first-growing season produced almost two-times more second-year adult perennial grasses compared to where water was not added. Conclusions: High water availability during the seeding year appears to be the most important factor for retaining perennial grasses in annual grass-invaded ecosystems. [ABSTRACT FROM AUTHOR]

Hydrothermal (HT) models can be used to characterize seed germination response to fieldvariable conditions of temperature (T) and water potential (Y). Hydrothermal response data are relatively difficult to generate, which limits their utility for large-scale comparisons of inter- and intraspecies germination response. Previous studies have hypothesized that HT germination response can be estimated using a simple model for thermal-time accumulation above a fixed threshold of environmental Y (wet-thermal [WT] model). The purpose of this study was to test this hypothesis by explicitly comparing HT and WT germination rates and estimated cumulative germination response of 13 rangeland grass seedlots under simulated conditions of field-variable T and Y. We used a 44-yr weather record to parameterize a seedbed-microclimate model for estimation of hourly T and Y at seeding depth for a sandy loam soil type at the Orchard field test site in southwestern Ada County, Idaho. Hydrothermal and WT germination responses for each hour of the simulation were estimated for 13 range grass seedlots. We found that overestimation of germination rate by thermal accumulation at low Y contributed relatively little to WT model error rates, and that >95% of the variability in predicted germination response could be explained by WT germination response above a Y threshold of -0.3 to -0.5 MPa. Given the pulse-like nature of favorable germination conditions in the field, this modeling approach may have immediate and wide potential application, as there are a relatively large number of thermal-germination datasets currently available for rangeland plant species. [ABSTRACT FROM AUTHOR]

Our ability to restore rangelands is limited, and it is unknown if seedling herbivory on its own, or in interaction with other stressors, is a major contributor to restoration failure. To address this, we conducted two experiments: a No Defoliation (ND) experiment ( n = 48), in which seedlings from three perennial grasses (crested wheatgrass [ Agropyron cristatum {(L.} Gaertn.], bluebunch wheatgrass [ Psuedoroegnaria spicata {Pursh} Á. Love], Sandberg bluegrass [ Poa secunda J Presl]) were subjected to wet and dry water regimes for 4 mo, and a concurrent Defoliation (D) experiment ( n = 95), in which seedlings were factorially assigned to two defoliation treatments—frequency (LOW, HIGH) and intensity (30% vegetation removal, 70% vegetation removal). Indicators of seedling performance were aboveground and belowground biomass (AGB and BGB), root:shoot ratio, tillering, and mortality. The effect size statistic, Hedge’s g, allowed for comparisons between performance measures. Water stress induced reductions in most performance measures: BGB ( g = ND: –1.3; D: –1.6), root:shoot ratio ( g = ND: n.s.; D: –0.2), and tillering ( g = ND: –1.7; D: –1.2), though not significantly for all species. For ABG, water stress interacted with defoliation, reducing performance less at an intensity of 70% ( g = –2.0) as opposed to 30% ( g = –3.0), but not always significantly in the former. Water stress also caused less reduction in AGB when no defoliation occurred (ND: –0.8; g = D: –2.5). Intensity and frequency of defoliation interacted; seedlings were generally resistant to reductions in performance except at high frequency, 70% defoliation. Agropyron cristatum and P. spicata displayed similar sensitivity to treatments, mostly in terms of changes in AGB and BGB, while P. secunda also experienced increased mortality and reduced tillering. If these differences in sensitivity result in differential survival, herbivory could impact species postrestoration population demographics. [ABSTRACT FROM AUTHOR]

Hydrothermal models used to predict germination response in the field are usually parameterized with data from laboratory experiments that examine the full range of germination response to temperature (T) and water potential (ψ). Inclusion of low-ψ and high- and low-T treatments, however, may not be necessary if their contribution to germination rate in the field is negligible. In this study, we simulated cumulative germination response of 13 range grass seedlots to a typical seasonal planting scenario used for rangeland restoration in the Great Basin region of the western United States. Given realistic estimates of actual time spent within the full hydrothermal range of these seedlots, >95% of germination response in the field could be predicted by models that only considered germination at ψ values less negative than -0.5 MPa and T values <25°C. Limiting laboratory treatments to a restricted hydrothermal range could greatly increase the efficiency of parameterizing hydrothermal models for predicting field response. Laboratory expenditures in this example could be reduced by 80 to 90% with only a relatively small reduction in the potential accuracy of field predictions. [ABSTRACT FROM AUTHOR]

Developing land-use practices that lead to sustainable net primary productivity in rangelands are important, but understanding their consequences to population and community processes is not often accounted for in basic ecosystem studies. Grazed and ungrazed upland ecosystems generally do not differ in net ecosystem CO2 exchange (NEE), but the underlying mechanisms and the concurrent effects of defoliation to vegetative and reproductive biomass allocation are unclear. To address this, we measured evapotranspiration (ET), NEE, and its constituent fluxes of ecosystem respiration (Reco) and gross ecosystem photosynthesis (GEP) with live canopy leaf area index (LAIlive; m² live leaf area/m² ground area) and aboveground leaf, culm, and reproductive biomass in plots of clipped and unclipped squirreltail (Elymus elymoides) and bluebunch wheatgrass (Pseudoroegneria spicata) growing in intact sagebrush steppe. Clipping reduced LAIlive by 75%, but subsequent re-growth rates in clipped plots was similar to LAIlive accumulation in unclipped plots. Concurrently, ET and NEE was similar between clipped and unclipped plots, with NEE primarily determined by GEP. GEP was initially lower in clipped plots, but then converged with unclipped GEP even as LAIlive continued to increase in both treatments. GEP convergence was driven by higher whole-plant photosynthesis (GEPlive = GEP/LAIlive) in clipped plots. Ecosystem water use efficiency (GEP/ET) was reduced by 16% with clipping, due to low GEP/ET 2 weeks following defoliation, but GEP/ET converged before GEP levels did. Proportional reproductive biomass was higher in E. elymoides (21.4% total biomass) than in P. spicata (0.5% total biomass) due to lower allocation to specific leaf and culm mass. Clipping reduced reproductive effort in E. elymoides, in terms of total reproductive biomass (-56%), seed mass per unit leaf area (-64%), and seed mass per flowering head (-77%). We concluded defoliation increased canopy-level light penetration, facilitating rapid Recovery of ecosystem fluxes, but that allocation to vegetative regrowth supporting this led to lower reproductive effort in these range grasses. Insights from studies such as this will be useful in formulating systems-based land management strategies aimed at maintaining annual productivity and long-term population and community goals in semiarid rangeland ecosystems. [ABSTRACT FROM AUTHOR]

The annual grass Bromus tectorum has invaded millions of hectares in western North America and has transformed former perennial grass and shrub-dominated communities into annual grasslands. Fire plays a key role in the maintenance of B. tectorum on the landscape but the type of disturbance responsible for initial invasion is less well understood. We conducted an experiment in a perennial shrub/grass/forb community in eastern Idaho, USA to examine the roles of plant community and soil disturbance on B. tectorum emergence and establishment prior to state-changing fires. Our experiment consisted of a plant community disturbance treatment where we (1) removed the shrub component, (2) removed the grass/forb component, or (3) removed all shrubs, grasses, and forbs. We followed this treatment with seeding of B. tectorum onto the soil surface that was (1) intact, or (2) disturbed. Each experimental plot had an associated control with no plant community disturbance but was seeded in the same manner. The experiment was replicated 20 times in two sites (high and low aboveground biomass). We measured emergence by counting seedlings in late spring and establishment by counting, removing, and weighing B. tectorum individuals in mid-summer. We also examined the influence of plant community disturbance on the soil environment by measuring extractable NH4+ and NO3- four times each summer. Soil disturbance greatly influenced the number of B. tectorum individuals that emerged each spring. Plant community disturbance, specifically disturbance of the grass/forb component, increased N availability in the late growing season and biomass of B. tectorum the following summer. We conclude that soil disturbance and plant community disturbance interact to promote the initial invasion of B. tectorum in Intermountain West valley ecosystems. [ABSTRACT FROM AUTHOR]

The invasion by winter-annual grasses (AGs) such as Bromus tectorum into sagebrush steppe throughout the western USA is a classic example of a biological invasion with multiple, interacting climate, soil and biotic factors driving the invasion, although few studies have examined all components together. Across a 6000-km area of the northern Great Basin, we conducted a field assessment of 100 climate, soil, and biotic (functional group abundances, diversity) factors at each of 90 sites that spanned an invasion gradient ranging from 0 to 100 % AG cover. We first determined which biotic and abiotic factors had the strongest correlative relationships with AGs and each resident functional group. We then used regression and structural equation modeling to explore how multiple ecological factors interact to influence AG abundance. Among biotic interactions, we observed negative relationships between AGs and biodiversity, perennial grass cover, resident species richness, biological soil crust cover and shrub density, whereas perennial and annual forb cover, tree cover and soil microbial biomass had no direct linkage to AG. Among abiotic factors, AG cover was strongly related to climate (increasing cover with increasing temperature and aridity), but had weak relationships with soil factors. Our structural equation model showed negative effects of perennial grasses and biodiversity on AG cover while integrating the negative effects of warmer climate and positive influence of belowground processes on resident functional groups. Our findings illustrate the relative importance of biotic interactions and climate on invasive abundance, while soil properties appear to have stronger relationships with resident biota than with invasives. [ABSTRACT FROM AUTHOR]

Seedling recruitment is a critical driver of population dynamics and community assembly, yet we know little about functional traits that define different recruitment strategies. For the first time, we examined whether trait relatedness across germination and seedling stages allows the identification of general recruitment strategies which share core functional attributes and also correspond to recruitment outcomes in applied settings. We measured six seed and eight seedling traits (lab- and field-collected, respectively) for 47 varieties of dryland grasses and used principal component analysis (PCA) and cluster analysis to identify major dimensions of trait variation and to isolate trait-based recruitment groups, respectively. PCA highlighted some links between seed and seedling traits, suggesting that relative growth rate and root elongation rate are simultaneously but independently associated with seed mass and initial root mass (first axis), and with leaf dry matter content, specific leaf area, coleoptile tissue density and germination rate (second axis). Third and fourth axes captured separate tradeoffs between hydrothermal time and base water potential for germination, and between specific root length and root mass ratio, respectively. Cluster analysis separated six recruitment types along dimensions of germination and growth rates, but classifications did not correspond to patterns of germination, emergence or recruitment in the field under either of two watering treatments. Thus, while we have begun to identify major threads of functional variation across seed and seedling stages, our understanding of how this variation influences demographic processes-particularly germination and emergence-remains a key gap in functional ecology. [ABSTRACT FROM AUTHOR]

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Seeding native plants is a key management practice to counter land degradation across the globe, yet the majority of seeding efforts fail, limiting our ability to accelerate ecosystem recovery., Recruitment requires transitions through several seed and seedling stages, some of which may have overriding influences on restoration outcomes. We lack, however, a general framework to understand and predict differences in these critical demographic processes across species. Functional traits influence fitness, and consequently, trait variation could provide the basis for a framework to explain and predict variation in life stage transition probabilities., We used seed and seedling traits, and field probabilities of germination, emergence, seedling establishment, and survival for 47 varieties of drylands grasses under two watering treatments to identify critical life stage transitions and quantify the effect of traits on cumulative survival through the first growing season., Variation in germination and emergence probabilities explained over 90% of the variation in cumulative survival regardless of seedling survival probabilities or watering treatment, with emergence probability being the strongest predictor of cumulative survival., Coleoptile tissue density and seed mass had significant effects on emergence and germination, respectively, explaining 10-23% of the variation in transition probabilities., Synthesis and applications. While the majority of functional trait work has centred on linking leaf and root traits to resource acquisition and utilization, our study demonstrates that traits associated with germination and emergence may have prevailing influences on restoration outcomes. A portion of these traits have been examined, but there is substantial opportunity to identify other key traits driving these demographic processes. These advancements will underpin our ability to develop trait-based frameworks for overcoming recruitment barriers and facilitating recovery of degraded systems across the globe. [ABSTRACT FROM AUTHOR]

Climate change effects on plants are expected to be primarily mediated through early life stage transitions. Snowfall variability, in particular, may have profound impacts on seedling recruitment, structuring plant populations and communities, especially in mid-latitude systems. These water-limited and frequently invaded environments experience tremendous variation in snowfall, and species in these systems must contend with harsh winter conditions and frequent disturbance. In this study, we examined the mechanisms driving the effects of snowpack depth and soil disturbance on the germination, emergence, and establishment of the native Pseudoroegnaria spicata and the invasive Bromus tectorum, two grass species that are widely distributed across the cold deserts of North America. The absence of snow in winter exposed seeds to an increased frequency and intensity of freeze-thaw cycles and greater fungal pathogen infection. A shallower snowpack promoted the formation of a frozen surface crust, reducing the emergence of both species (more so for P. spicata). Conversely, a deeper snowpack recharged the soil and improved seedling establishment of both species by creating higher and more stable levels of soil moisture availability following spring thaw. Across several snow treatments, experimental disturbance served to decrease the cumulative survival of both species. Furthermore, we observed that, regardless of snowpack treatment, most seed mortality (70-80 %) occurred between seed germination and seedling emergence (November-March), suggesting that other wintertime factors or just winter conditions in general limited survival. Our results suggest that snowpack variation and legacy effects of the snowpack influence emergence and establishment but might not facilitate invasion of cold deserts. [ABSTRACT FROM AUTHOR]

Plant invasion and restoration outcomes are largely driven by the timing and magnitude of seed dispersal, and by the performance of dispersed species in an environment. Because seed dispersal controls recruitment of newly arriving species and facilitates safe site occupation, assembly will differ depending on seed dispersal processes and variable environmental conditions. The objective of this study was to identify how annual and perennial grasses assembled when dispersal times, propagule pressure, and water availability were modified. To assess these effects, we conducted a field experiment in an annual grass invaded shrub-steppe ecosystem in eastern Oregon. We tested the effects of seeding annual and perennial grasses in autumn or delaying annual grass seeding until spring, adding water, and varying annual and perennial grass seeding rate by 150, 1,500, 2,500, or 3,500 seeds m on perennial and annual grass seedling emergence through time and final density and biomass. Providing perennial grasses a priority effect by delaying annual grass seeding until spring initially facilitated perennial grass establishment, but this effect did not persist into the second growing season. We found that if annual grass propagule pressure exceeded 150 seeds m, perennial grass recruitment was limited. In addition, higher water availability increased perennial grass establishment, but was dependent upon annual grass propagule pressure. These findings suggest that seeding perennial grasses into annual grass dominated systems is more dependent upon the existing propagule pressure of annual grasses than the priority effects of perennial grasses, the propagule pressure of perennial grasses, or water availability. [ABSTRACT FROM AUTHOR]

In a previous article, Beschta et al. (Environ Manag 51(2):474-491, ) argue that grazing by large ungulates (both native and domestic) should be eliminated or greatly reduced on western public lands to reduce potential climate change impacts. The authors did not present a balanced synthesis of the scientific literature, and their publication is more of an opinion article. Their conclusions do not reflect the complexities associated with herbivore grazing. Because grazing is a complex ecological process, synthesis of the scientific literature can be a challenge. Legacy effects of uncontrolled grazing during the homestead era further complicate analysis of current grazing impacts. Interactions of climate change and grazing will depend on the specific situation. For example, increasing atmospheric CO and temperatures may increase accumulation of fine fuels (primarily grasses) and thus increase wildfire risk. Prescribed grazing by livestock is one of the few management tools available for reducing fine fuel accumulation. While there are certainly points on the landscape where herbivore impacts can be identified, there are also vast grazed areas where impacts are minimal. Broad scale reduction of domestic and wild herbivores to help native plant communities cope with climate change will be unnecessary because over the past 20-50 years land managers have actively sought to bring populations of native and domestic herbivores in balance with the potential of vegetation and soils. To cope with a changing climate, land managers will need access to all available vegetation management tools, including grazing. [ABSTRACT FROM AUTHOR]

Changes in the quantity and quality of plant litter occur in many ecosystems as they are invaded by exotic species, which impact soil nutrient cycling and plant community composition. Such changes in sagebrush-steppe communities are occurring with invasion of annual grasses (AG) into a perennial grass (PG) dominated system. We conducted a 5-year litter manipulation study located in the northern Great Basin, USA. Springtime litter was partially or completely removed in three communities with differing levels of invasion (invaded, mixed, and native) to determine how litter removal and litter biomass affected plant-available soil N and plant community composition. Litter biomass (prior to the removal treatment) was negatively correlated with plant-available N in the invaded community, but was positively correlated in the native community. Plant-available N had greater intra- and inter-annual fluctuations in the invaded compared to the mixed or native communities, but was not generally affected by removal treatments. Litter removal had negative effects on AG cover during a warm/dry year and negative effects on PG cover during a cool/wet year in the mixed community. Overall, the effectiveness of springtime litter manipulations on plant-available N were limited and weather dependent, and only removal treatments >75 % had effects on the plant community. Our study demonstrates how communities invaded by AGs have significantly increased temporal variability in nutrient cycling, which may decrease ecosystem stability. Further, we found that the ecological impacts from litter manipulation on sagebrush communities were dependent on the extent of AG invasion, the timing of removal, and seasonal precipitation. [ABSTRACT FROM AUTHOR]

Functional differences between native and exotic species potentially constitute one factor responsible for plant invasion. Differences in trait values between native and exotic invasive species, however, should not be considered fixed and may depend on the context of the comparison. Furthermore, the magnitude of difference between native and exotic species necessary to trigger invasion is unknown. We propose a criterion that differences in trait values between a native and exotic invasive species must be greater than differences between co-occurring natives for this difference to be ecologically meaningful and a contributing factor to plant invasion. We used a meta-analysis to quantify the difference between native and exotic invasive species for various traits examined in previous studies and compared this value to differences among native species reported in the same studies. The effect size between native and exotic invasive species was similar to the effect size between co-occurring natives except for studies conducted in the field; in most instances, our criterion was not met although overall differences between native and exotic invasive species were slightly larger than differences between natives. Consequently, trait differences may be important in certain contexts, but other mechanisms of invasion are likely more important in most cases. We suggest that using trait values as predictors of invasion will be challenging. [ABSTRACT FROM AUTHOR]

Drylands support over 2 billion people and are major providers of critical ecosystem goods and services across the globe. Drylands, however, are one of the most susceptible biomes to degradation. International programmes widely recognize dryland restoration as key to combating global dryland degradation and ensuring future global sustainability. While the need to restore drylands is widely recognized and large amounts of resources are allocated to these activities, rates of restoration success remain overwhelmingly low., Advances in understanding the ecology of dryland systems have not yielded proportional advances in our ability to restore these systems. To accelerate progress in dryland restoration, we argue for moving the field of restoration ecology beyond conceptual frameworks of ecosystem dynamics and towards quantitative, predictive systems models that capture the probabilistic nature of ecosystem response to management., To do this, we first provide an overview of conceptual dryland restoration frameworks. We then describe how quantitative systems framework can advance and improve conceptual restoration frameworks, resulting in a greater ability to forecast restoration outcomes and evaluate economic efficiency and decision-making. Lastly, using a case study from the western United States, we show how a systems approach can be integrated with and used to advance current conceptual frameworks of dryland restoration., Synthesis and applications. Systems models for restoration do not replace conceptual models but complement and extend these modelling approaches by enhancing our ability to solve restoration problems and forecast outcomes under changing conditions. Such forecasting of future outcomes is necessary to monetize restoration benefits and cost and to maximize economic benefit of limited restoration dollars. [ABSTRACT FROM AUTHOR]

Cultivation legacies affect native vegetation in old fields of the Great Basin, USA for nearly a century after these fields are abandoned. We hypothesized that cultivation lowered soil nutrients and that this legacy would differentially impact plant performance of four representative Great Basin species. To test these hypotheses, we compared soil nutrients (C, N, P, K, Mg and Ca) between two formerly cultivated and adjacent noncultivated sites in two soil series. We then compared the plant growth and foliar nutrient content of an exotic grass ( Bromus tectorum L.), two native grasses ( Elymus elymoides [Raf.] Swezey and Achnatherum hymenoides [Roem. and Schult.] Barkworth), and a native forb ( Sphaeralcea grossulariifolia [Hook. and Arn.] Rydb) grown in these soils in the greenhouse and in the field. Only one sampling site had reduced soil nutrients associated with cultivation legacies, where most of the negative effects on plant performance were found. E. elymoides appeared to be less affected by cultivation legacies than did A. hymenoides, which had a reduced survivorship and 20 % less above-ground biomass in cultivated soils. No species, including B. tectorum, were favored by cultivation. Our findings suggest that cultivation legacies can affect plant performance of different species in different ways and that altered soil nutrients may interact with other abiotic and biotic cultivation legacies in complex ways. [ABSTRACT FROM AUTHOR]

Plant competition is a primary ecological process limiting grassland restoration success. Appropriate restoration techniques require an understanding of the degree to which intra and interspecific competition control invasive and native plant growth. The objective of this study was to determine how the intensity of intra and interspecific competition changes during early stages of plant growth. Two invasive ( Bromus tectorum and Taeniatherum caput- medusae) and two native ( Pseudoroegneria spicata and Poa secunda) species were grown in a diallel competition experiment, either alone or in 1:1 binary combinations and exposed to two levels of N (no N or 400 mg N kg soil added) in a greenhouse. Total biomass for each species was quantified over four harvests and competitive effects were calculated. Our results show that the relative magnitude of intra and interspecific competition changes through time. Intraspecific competition was intense for native species at the initial harvests and therefore important in contributing to the outcome of final size of native species seedlings. Interestingly, bluebunch wheatgrass imposed interspecific competition on annual grasses at the first two harvests and appeared to be a better competitor than Sandberg's bluegrass. We found that fast growing invasive species became more competitive compared to slow growing native species with increasing N and appear to establish a positive feedback mechanism between size and resource uptake. Opportunities to improve restoration success exist from determining the optimum combination of density, species proportion, and their spatial arrangement in various ecosystems and environments. [ABSTRACT FROM AUTHOR]

Invasive plants are degrading wildlands around the globe by displacing native species, reducing biodiversity, and altering ecological functions. The current approach of applying herbicides to invasive plants in wildlands has not been effective at curtailing their expansion and, in certain circumstances, may do more harm than good. Preventing the spread of invasive species has been identified as an important strategy to protect wildlands. However, few prevention strategies have actually been tested. We hypothesized that establishing competitive vegetation next to infestations would increase the biotic resistance of the plant community to invasion and decrease the invasive species propagule pressure beyond the competitive vegetation. To evaluate this, we established twelve competitive vegetation barriers in front of invasive annual grass, Taeniatherum caput- medusae (L.) Nevski, infestations. The non-native perennial grass Agropyron desertorum (Fisch. ex Link) Schult. was seeded into plant communities adjacent to the infestations to create the competitive vegetation barriers. Soil nutrient concentrations and the spread of T. caput- medusae were compared between where A. desertorum was seeded and not seeded (control treatment) 3 years after treatment. Less T. caput- medusae and lower soil ammonium and potassium concentrations in the competitive vegetation barrier than control treatment ( P ≤ 0.01) suggest that establishing competitive vegetation increased the biotic resistance of the plant communities to invasion. Taeniatherum caput- medusae cover and density in the plant communities protected by the competitive vegetation barrier (locales across the barriers from the infestations) were ~42- and 47-fold less, respectively, than unprotected plant communities ( P < 0.01). This suggests that invasive plant propagule pressure was decreased in the plant communities protected by competitive vegetation barriers. The establishment of competitive vegetation around infestations may be an effective strategy to prevent or at least reduce the spread of invasive plant species. [ABSTRACT FROM AUTHOR]

The article presents a study which examines the grassland natives exhibiting long-term coexistence with an invader, and determines whether herbicide use can affect invasive species. One-time herbicide use made two native forbs exceedingly rare for the entire study period. It was found that the dominant invader became more abundant in response to the decreases in native-forb abundances. However, some applications are not warranted because they target small invader patches or larger areas with no remaining natives.

Seeding of non-native species following wildfires to stabilize soils and prevent erosion has become a controversial practice because it risks inhibiting tree regeneration, introducing noxious weeds, and permanently replacing native species. This paper reports the fate of non-native seeded species during 31 yean; of postfire forest,succession in northcentral Washington state, USA. In that region, catastrophic wildfires in grand fir (Abies grandis)/pine grass (Calamagrostis rubescens) associations characteristically result in a flush of lodgepole pine (Pinus contorta) regeneration that creates a dense tree monoculture. In the study, seven grass cultivars were broadcast seeded to prevent erosion, limit tree regeneration, and increase forage production for wildlife and livestock. Tree regeneration, native and non-native species cover, and above- ground biomass were evaluated in 1971, 1975, 1980, 1989, and 2001 . The seeded species quickly established dominant cover with levels of biomass production two to three times the level of native species. Density of tree regeneration was inversely correlated with perennial grass cover during the first 10 years. Between 1980 and 1 989, the seeded grasses disappeared, long before tree canopyclosure. Within 15 to 20 years native species had regained dominance, and after 30 years the last remnants of the non-native cultivars were replaced in the seeded areas by a diverse mixture of native graminoids, forbs, shrubs, and trees. In contrast, a monoculture of lodgepole pine dominated the unseeded areas. This study showed that non-native grasses seeded after wildfires do not always persist and can serve as a transition to restoring a more diverse seral community. [ABSTRACT FROM AUTHOR]

Revegetation of degraded shrub-steppe often fails due to intense competition from weeds, highly variable environmental conditions, and limited soil moisture. The objective of this study was to test whether a commercially available seed coating and a water-retaining acrylamide copolymer hydrogel would increase seedling emergence and establishment of crested wheatgrass (Agropyron cristatum) under three watering frequencies and two soil textures. Pots were filled with one of four soil treatments: field soil, two parts field soil mixed with one part sand, field soil plus hydrogel, or field soil plus sand plus hydrogel. We seeded the pots with coated or uncoated seed and placed them in a greenhouse for 66 days. The pots were assigned one of three watering treatments: 4.5 oz (150 ml) applied one time per week, 1 .5 oz (50 ml) applied three times per week, or 0.9 oz (30 ml) applied five times per week. We recorded seedling emergence at three and six weeks. At the end of 66 days, the numbers of seedlings that survived were counted and aboveground biomass was collected, dried, and weighed. We found that uncoated seed had 1.6 times greater seedling density than coated seed. Incorporation of the acrylamide copolymer hydrogel into the potting medium conferred some benefit to emergence, biomass, and survivorship of crested wheatgrass seedlings. However, it was watering frequency that produced the most consistent influence on seedling emergence, survival, and biomass. Watering three or five times per week increased emergence more than watering one time per week, but watering one time per week generally led to greater survivorship and biomass. This suggests that the use of water-retaining hydrogels may help to overcome soil moisture limitations and improve seedling establishment during revegetation of degraded shrub-steppe. [ABSTRACT FROM AUTHOR]

Establishing native forbs is crucial for invasive plant management and restoring a desirable plant community. Our objectives were to determine (1) if increasing forb seed density results in increased forb establishment; (2) if a species-rich mixture of forbs has greater establishment and survivorship than a single species; and (3) if mixtures of forbs are more competitive with Spotted knapweed ( Centaurea maculosa) than a forb monoculture. To test our first two objectives, we seeded monocultures of Purple coneflower ( Echinacea angustifolia), Arrowleaf balsamroot ( Balsamorhiza sagittata), Annual sunflower ( Helianthus annuus), Dotted gayfeather ( Liatris punctata), Western white yarrow ( Achillea millefolium), Sticky geranium ( Geranium viscosissimum), as well as a mixture of all forbs. Pots were seeded at 800 or 2,000 seeds/m2 and watered twice or thrice weekly. The highest seed density produced the highest plant density, which averaged 4.35 plants/pot. The density of the mixture was similar to the mean density seen for individual species, and it doubled in response to the highest seed density. To test our third objective, Spotted knapweed and Purple coneflower were arranged in an addition series matrix with a maximum total density of 4,000 seeds/pot. We found that the forb mixture was seven times more competitive with Spotted knapweed than Purple coneflower alone. Using a mixture of forbs rather than a single species enhances forb establishment in various and unpredictable environments because the mixture possesses a variety of traits that may match year–year and site–site conditions. Once established, the mixture may have a greater chance of persisting than a monoculture. [ABSTRACT FROM AUTHOR]

Environmental resource gradients affect plant establishment during invasive weed management and ecological restoration. We investigated how hydrology and nitrogen (N) availability influence establishment of native and invasive species along a dry-wet gradient. We hypothesized that 1) emergence and growth would be higher in subirrigaated environments than in saturated or dry environments; 2) annuals and invasive exotics would respond more to N availability than late-successional perennials; and 3) N effects would be greatest in subirrigated environments. In the greenhouse, seedling emergence was uniform across dry to subirrigated pots for upland species and across subirrigated to saturated, but not flooded, pots for riparian species. Seedling biomass was greatest in subirrigated environments. Seedling emergence in the field was at or near zero for most species in subirrigated sites. In the greenhouse, biomass increased with N availability, and responses to N were greatest in subirrigated pots. In both experiments, the upland invasive, spotted knapweed (Centaurea maculosa), responded more to N availability than bluebunch wheatgrass (Agropyron spicatum). However, such differences were not found for the riparian species, possibly due to low emergence. Possible explanations for low emergence in subirrigated field sites include predation by small rodents, which were more active in the transition than upland or wetland areas. Establishment of spotted knapweed and Canada thistle (Cirsium arvense) in riparian sites was limited by poor germination and seedling survival. Once established, growth was good in subirrigated environments, indicating that weeds may invade riparian sites in spite of poor initial establishment, which suggests caution when contemplating ecological restoration practices that increase N availability. [ABSTRACT FROM AUTHOR]

Ecologists are searching for models, frameworks, and principles that provide a bridge between theory and the practice of restoration. Successional management has been proposed as a useful model for managing and restoring invasive-plant-dominated rangeland because it provides a framework in which ecological processes can be manipulated by managers to achieve a desired plant community. Successional management identifies three general causes of succession (site availability [disturbance], species availability [colonization], and species performance) and suggests that managers address the ecological process influencing each general cause in a coordinated fashion to direct plant community dynamics. We tested successional management using various techniques to restore invasive-weed-dominated rangeland. Our hypothesis was that successively modifying the factors influencing the causes of succession in an integrated fashion would favor the establishment and abundance of native grasses over singularly applied treatments. Thus, we anticipated that the majority of responses to multiple treatments would be explained by higher order interactions, especially in the final year of the study (2004). To test this hypothesis; we used a model system within a Festuca campestris/Pseudoroegneria spicata habitat among pothole wetlands dominated by Centaurea maculosa and Potentilla recta, two invasive species. We used three herbicide treatments (none, 2,4-D, and picloram) to influence species performance; two seeding methods (imprinting, i.e., creating a small depression and broadcasting, and no-till drilling) to influence disturbance; three seeding rates (977, 1322, and 1557 seeds/m2) to influence colonization; and two cover crop treatments (with and without Triticum aestivum) to influence soil N and favor native grasses. Treatments were factorially arranged and replicated four times in a randomized complete block design in 2001 and sampled in 2002 and 2004. As predicted, plant response to treatments was dominated by two- and three-way interactions in 2004. The highest seeding rate (colonization) combined with no-till drilling (disturbance) produced the highest native grass density in 2002. These effects persisted into 2004 for P. spicata. but not for F. campestris or F. idahoensis. Combining picloram with no-till drill seeding also produced a high density of P. spicata. Drill seeding at 977 seeds/m2 favored F. idahoensis density, while no-till drilling at 1322 seeds/m2 favored its biomass in 2004. F. idahoensis established well after drill seeding with a cover crop and applying 2,4-D. Herbicides reduced native forb density and/or biomass, with early season forbs being more sensitive topicloram and summer forbs being more sensitive to 2,4-D. Herbicides increased exotic grasses’ density and biomass but had no effect on native grasses. In most cases, integrating treatments that addressed multiple causes of succession favored a desired plant community. Thus, we accomplished our goal of using successional management to direct plant communities toward native desired species, but the treatments used did not improve species richness. Since naturally occurring native forbs did not respond favorably to any treatment combination, ecological restoration using successional management may best be thought of as an iterative procedure where various components and processes of the system are methodically repaired or replaced over time. [ABSTRACT FROM AUTHOR]

A commonly cited mechanism for invasion resistance is more complete resource use by diverse plant assemblages with maximum niche complementarity. We investigated the invasion resistance of several plant functional groups against the nonindigenous forb Spotted knapweed ( Centaurea maculosa). The study consisted of a factorial combination of seven functional group removals (groups singularly or in combination) and two C. maculosa treatments (addition vs. no addition) applied in a randomized complete block design replicated four times at each of two sites. We quantified aboveground plant material nutrient concentration and uptake (concentration × biomass) by indigenous functional groups: grasses, shallow-rooted forbs, deep-rooted forbs, spikemoss, and the nonindigenous invader C. maculosa. In 2001, C. maculosa density depended upon which functional groups were removed. The highest C. maculosa densities occurred where all vegetation or all forbs were removed. Centaurea maculosa densities were the lowest in plots where nothing, shallow-rooted forbs, deep-rooted forbs, grasses, or spikemoss were removed. Functional group biomass was also collected and analyzed for nitrogen, phosphorus, potassium, and sulphur. Based on covariate analyses, postremoval indigenous plot biomass did not relate to invasion by C. maculosa. Analysis of variance indicated that C. maculosa tissue nutrient percentage and net nutrient uptake were most similar to indigenous forb functional groups. Our study suggests that establishing and maintaining a diversity of plant functional groups within the plant community enhances resistance to invasion. Indigenous plants of functionally similar groups as an invader may be particularly important in invasion resistance. [ABSTRACT FROM AUTHOR]

Abstract Centaurea maculosa (Lam.) (spotted knapweed) reduces wildlife and livestock habitat biodiversity and increases erosion. Nutrient availability to plants may be used to accelerate succession away from spotted knapweed. Early-successional plant communities often have high nutrient availability, whereas late-successional communities are often found on lower nutrient soils. We hypothesized that removal of nutrients would change the competitive advantage from spotted knapweed to Pseudoroegneria spicatum (bluebunch wheatgrass) (late seral). In two addition series matrices, background densities of Secale cereale (annual rye) and Elymus elimoides (bottlebrush squirreltail) (3,000 seeds/m2) were used to remove nutrients from the soil. In another set of addition series matrices, nitrogen (33 kg/ha) or phosphorus (33 kg/ha) were added to the soil. Nutrient analysis of soil and vegetation indicated that annual rye and bottlebrush squirreltail reduced nutrient availability in soils. In another matrix, neither a background density nor nutrients were added. Data were fit into Watkinson's curvilinear model to determine the competitive relationship between bluebunch wheatgrass and spotted knapweed. This allowed comparison of the equivalence ratios (C) generated from each addition series. The C parameters are the per-plant equivalent of bluebunch wheatgrass or spotted knapweed and can be interpreted as the ratio of intra-to-interspecific competition. The C parameters are also the equivalence ratio of the number of spotted knapweed it takes to have equivalent effect on bluebunch wheatgrass or the number of bluebunch wheatgrass having the equivalent effect on spotted knapweed. Without nutrient manipulation, spotted knapweed was more competitive than bluebunch wheatgrass. The C for bluebunch wheatgrass was 0.17, indicating that 0.17 knapweed plants were competitively equivalent to one wheatgrass. Annual rye changed the competitive balance in favor of bluebunch... [ABSTRACT FROM AUTHOR]

Herbicidal control of spotted knapweed, Centaurea maculosa Lamarck, is rarely cost-effective, and sustainable control may require an integrated approach. Cyphocleonus achates (Fahraeus) is a flightless root-feeding weevil of Eurasian origin that has been introduced into North America for biological control of spotted knapweed. We hypothesizedthat reducing the density of spotted knapweed using reduced rates ofpicloram would improve the establishment of C. achates. At two sitesin western Montana, three adult weevil densities (none, three, and six weevils m-2) were released in the fall (1995 at site 1 and 1996 atsite 2) in 2-m2 plots encircled with enclosures to prevent weevil escape. Six picloram rates (0, 0.03, 0.06, 0.09, 0.12 or 0.15 kg ha-1) were applied the following spring before weevil emergence in a randomized complete-block design with four replications (18 treatments per replication). Spotted knapweed density and spotted knapweed and grass cover were sampled in July each year following treatment.Weevil numbers were counted annually in August. By 1998, picloram rates of >0.09 kg ha-1 had reduced spotted knapweed density from -500 to -175 plants m-2. Cover was reduced from ~60 to 10% at these rates. There was no detectable impact of weevils on spotted knapweed. Weevil numbers in plots treated with picloram >0.09 kg ha-1 were about half (0.5 weevils per plot) the numbers found in the other picloram treatments (1.25 weevils per plot). Weevil numbers in plots treated with 0.03, 0.06, and 0.09 kg ha-1 were similar to the unsprayed control. Weevil numbers were highestwhen spotted knapweed cover was between 30 and 70%. Results show than reduced rates of picloram do not limit weevil establishment. [ABSTRACT FROM AUTHOR]