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Establishing competitive plants is essential for restoring spot ted knapweed infested grasslands. Revegetation attempts typically fail because of weed competition during the initial stages of establishment.We hypothesized that competitive interactions can be shifted from spotted knapweed to intermediate wheatgrass by increasing wheatgrass seedling density over 1,000 plants m-2. Spotted knapweed andintermediate wheatgrass were grown in addition series mixtures to assess their interference at low (0 to 1,000 plants m- 2) versus high (1,000 to 10,000 plants m-2) densities. In the spring of 1995, 7 densities (0, 100, 500, 1,000, 3,000, 6,000, and 10,000 plants m-2) of each species were seeded in a factorialarrangement (49 density combinations) in a randomized-complete-blockdesign and replicated 3 times at 2 sites in Montana. Plants were grown in pots (2,250 mm2 x 380 mm deep) for 60 days before harvesting. Regressions predicting shoot weight, root weight, total weight, leaf area, and root length were calculated using 1) low knapweed:low wheatgrass, 2) low knapweed:high wheatgrass, 3) high knapweed:low wheatgrass, and 4) high knapweed:high wheatgrass densities. Regression coefficients indicated intraspecific interference was most important in predicting intermediate wheatgrass weight at both sites. At the wet site (457 mm, annually), interspecific interference only occurred at high spotted knapweed densities. At the dry site (305 mm, annually), interspecific interference occurred at low densities. Increasing intermediate wheatgrass from low to high densities removed the effect of spotted knapweed on intermediate wheatgrass where interspecificinterference occurred. [ABSTRACT FROM AUTHOR]

Spotted knapweed (Centaurea maculosa Lam.) has reduced forage production, increased soil erosion, and lowered biodiversity on millions ofhectares of rangeland throughout the western United States. Objectives of this study were to quantify the interaction between picloram (4-amino-3,5,6-trichloropicolinic acid) and fertilizer on spotted knapweed density and grass yield. Four picloram rates (0.0, 0.14, 0.28, and 0.42 kg ha-1) and 4 fertilizer rates (N+P: 0.0+0.0, 10.5+12.2, 21.1+26.4, and 31.7+39.6 kg ha- 1) were applied to 3 spotted knapweed infested rangeland sites in a factorial combination arranged in a randomized-complete-block design during the spring of1994. Grass yield and spotted knapweed density were measured at peakstanding grass crop in 1994 and 1995. Data were analyzed as a split-plot in time using analysis of variance. Picloram and fertilizer did not interact to affect either spotted knapweed density or grass yield. All picloram treatments reduced spotted knapweed density to nearly zero. By 1995, all picloram treatments increased grass yield by an average of 1,500 kg ha-1. Fertilization did not affect spotted knapweed density, but the highest rates increased grass yield on those sites with a substantial residual grass understory. Combining fertilizer with picloram may enhance grass yield on sites with a residual of highly productive grasses. [ABSTRACT FROM AUTHOR]

Developing rangeland management strategies to minimize spotted knapweed (Centaurea maculosa Lam.) invasion will require understanding theeffects of intensity, frequency, and season of grazing on weed establishment. We studied the effects of hand-clipping 2-year-old Idaho fescue (Festuca idahoensis Elmer) plants 0, 30, 60, or 90%, 1, 2, or 3 times (14-day intervals) on spotted knapweed seedling emergence and growth in pots watered with 150 ml once weekly or 50 ml 3 times weekly. Pots were seeded with 5,000 spotted knapweed seeds m-2, replicated twice, and placed in a growth chamber in a completely randomized design. The experiment was repeated once. Plants were harvested after 50 days. Treatment effects on soil moisture, Idaho fescue andspotted knapweed shoot and root weight, and leaf area were compared using analysis of variance and regression analysis. At final harvest,Idaho fescue shoot weight and leaf area decreased with increasing defoliation level and frequency. Idaho fescue root weight was not affected by any treatment. A single Idaho fescue defoliation at 30% and 90% increased spotted knapweed weight and numbers per pot respectively,over those pots with undefoliated plants. The level of defoliation necessary to enhance spotted knapweed numbers was lower as defoliationfrequency increased. As defoliation level and frequency increased, soil water content increased resulting in a corresponding increase in spotted knapweed emergence and growth. [ABSTRACT FROM AUTHOR]

Community dynamics and dominance on cheatgrass (Bromus tectorum L.) and yellow starthistle (Centaurea solstitialis L.) infested rangelandappear to be influenced by resource acquisition rates and duration of growth. Objectives were to determine the effects of densities, proportions, and soil depth on the growth rate and duration of growth of these species. In 6 field experiments isolated individuals, monocultures (100, 1,000, 10,000 plants m{sup}-2{end}), and mixtures (same densities arranged factorially) were grown with unrestricted and restricted (0.2- and 0.5-m) soil depths. Shoot weights were determined on 12-day intervals beginning on day 24 and ending on day 72 for plants grown with restricted soil depth and day 96 (cheatgrass) and day 108 (yellow starthistle) for plants grown in unrestricted soil. Quadratic growth curves were fit for each replication for plants grown in isolation. Linear and quadratic models were developed for plants grown in monocultures and mixtures. Simple linear regression coefficients were used as growth rates and regressed over plant density. Time of injection (an indicator of the duration of growth) for plants growing in monocultures and mixtures was calculated from quadratic models. Growth rate of yellow starthistle was about 7 times faster and duration of growth 25 days longer, than those of cheatgrass when grown in isolation without soil depth restriction. As densities were increased and/or soil depth decreased, growth rate and duration of growth were loweredand the difference between species were masked. Growth rates and duration of growth of cheatgrass and yellow starthistle appear to dependon plant density, soil depth, and available soil moisture. [ABSTRACT FROM AUTHOR]

Diffuse knapweed (Centaurea diffuse Lam.), an invasive weed, has reduced forage production and biodiversity, and increased soil erosion on over a million hectares of rangeland in the western United States. This study evaluated the effects of a single grass defoliation on establishment of diffuse knapweed seeded at 2 rates into a bluebunch wheatgrass (Pseudoroegneria spicata [Pursh.] Scribn and Smith)/needle-and-thread (Stipa comata Trin. & Rupr.) community and a crested wheatgrass (Agropyron cristatum (L.) Gaertn.) community. Six defoliation levels (0, 20,40, 60, 80, 100 percent) and 2 seeding rates (3,000, 6,000 diffuse knaweed seeds) were applied to 1 m{sup}2{end} plots in a randomized-complete-block design (n=4). Diffuse knapweed was seeded in the fall of 1992, and grasses were defoliated on 28 April 1993. The number of flowering culms and weed seedlings were counted in September1993. Densities of diffuse knapweed seedlings, juveniles, and adults, as well as plant standing crop, were determined in May 1994. Seed rate had minimal effect on diffuse knapweed density. By May 1994, densities of diffuse knapweed were about 20 and 30 plants m{sup}2{end} onundefoliated bluebunch wheatgrass and crested wheatgrass plots, respectively, indicating that defoliation is not required for this noxious weed to become established. Higher levels of grass defoliation ( greater than 60 percent), especially of bluebunch wheatgrass, enhanced diffuse knapweed establishment, indicating that moderate ( less than 60 percent) defoliation would not necessarily accelerate invasion by diffuse knapweed. [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]

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]

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., (Published by Oxford University Press on behalf of the Annals of Botany Company 2021.)

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.

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(2) 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.

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.

In a previous article, Beschta et al. (Environ Manag 51(2):474-491, 2013) 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 CO₂ 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.

In recent decades, dozens of studies have involved attempts to introduce native and desirable nonnative plant species into grasslands dominated by invasive weeds. The newly introduced plants have proved capable of establishing, but because they are rarely monitored for more than four years, it is unknown if they have a high likelihood of persisting and suppressing invaders for the long-term. Beyond invaded grasslands, this lack of long-term monitoring is a general problem plaguing efforts to reintroduce a range of taxa into a range of ecosystems. We introduced species from seed and then periodically measured plant abundances for nine years at one site and 15 years at a second site. To our knowledge, our 15-year data are the longest to date from a seeding experiment in invaded, never-cultivated grassland. At one site, three seeded grasses maintained high densities for three or more years, but then all or nearly all individuals died. At the second site, one grass performed similarly, but two other grasses proliferated and at least one greatly suppressed the dominant invader (Centaurea maculosa). In one study, our point estimate suggests that the seeded grass Thinopyrum intermedium reduced C. maculosa biomass by 93% 15 years after seeding. In some cases, data from three and fewer years after seeding falsely suggested that seeded species were capable of persisting within the invaded grassland. In other cases, data from as late as nine years after seeding falsely suggested seeded populations would not become large enough to suppress the invader. These results show that seeded species sometimes persist and suppress invaders for long periods, but short-term data cannot predict if, when, or where this will occur. Because short-term data are not predictive of long-term seeded species performances, additional long-term data are needed to identify effective practices, traits, and species for revegetating invaded grasslands.