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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]

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]

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]

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]

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]