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Questions: (1) What combinations of overlapping water and heat stress and herbivory disturbance gradients are associated with shifts in interactions between Artemisia tridentata subsp. wyomingensis (Artemisia) and herbaceous beneficiary species? (2) Do interactions between Artemisia and beneficiaries shift from competition to facilitation with increasing stress-disturbance where facilitation and competition are most frequent and strongest at the highest and lowest levels, respectively? (3) Do such relationships differ for native and non-native beneficiaries? (4) What are the implications of any observed shifts in interactions between community compositional stability in space and susceptibility to invasion? Location: North American Artemisia communities. Methods: We tested the stress-gradient hypothesis (SGH) in an observational study consisting of 75 sites located along overlapping water and heat stress and disturbance gradients. We used spatial patterns of association among Artemisia and six native and two non-native beneficiary species; including the invasive annual grass Bromus tectorum, representing a diverse array of life history strategies, to infer whether the net outcome of interactions was facilitation or competition. We assessed implications for community stability by examining shifts in community composition in space and resistance to invasion. Results/Conclusions: Cattle herbivory, a novel disturbance and selective force, was a significant component of two overlapping stress gradients most strongly associated with observed shifts in interactions. Facilitation and competition were strongest and most frequent at the highest and lowest stress levels along both gradients, respectively. Contrasting ecological optima among native and non-native beneficiaries led to strikingly different patterns of interactions. The four native bunchgrasses with the strongest competitive response abilities exhibited the strongest facilitation at their upper limits of stress tolerance, while the two non-natives exhibited the strongest competition at the highest stress levels, which coincided with their maximum abundance. Artemma facilitation enhanced stability at intermediate stress levels by providing a refuge for native bunchgrasses, which in turn reduced the magnitude of B. tectorum invasion. However, facilitation was a destabilizing force at the highest stress levels when native bunchgrasses became obligate beneficiaries dependent on facilitation for their persistence. B. tectorum dominated these communities, and the next fire may convert them to annual grasslands.

Plant–plant interactions are important drivers of ecosystem structure and function, yet predicting interaction outcomes across environmental gradients remains challenging. Understanding how interactions are affected by ontogenetic shifts in plant characteristics can provide insight into the drivers of interactions and improve our ability to anticipate ecosystem responses to environmental change. We developed a conceptual framework of nurse shrub facilitation of tree establishment. We used a combination of field experiments and environmental measurements to test the framework with a shrub (Artemisia tridentata) and a tree (Pinus monophylla), two foundation species in a semiarid environment. Shrub microsites allowed trees to overcome an early population bottleneck and successfully establish in areas without tree cover. Shrubs facilitated trees at multiple ontogenetic stages, but the net outcome of the interaction shifted from strongly positive to neutral after the transition of P. monophylla from juvenile to adult foliage. Microhabitat conditions varied across a broad elevational gradient, but interaction outcomes were not strongly related to elevation. Favorable microsites provided by A. tridentata cover are crucial for P. monophylla recovery after stand-replacing disturbance. Models of vegetation response to rapid global environmental change should incorporate the critically important role of nurse shrub interactions for ameliorating population bottlenecks in tree establishment.

Fire refugia and patchiness are important to the persistence of fire-sensitive species and may facilitate biodiversity conservation in fire-dependent landscapes. Playing the role of ecosystem engineers, large herbivores alter vegetation structure and can reduce wildfire risk. However, herbivore effects on the spatial variability of fire and the persistence of fire-sensitive species are not clear. To examine the hypothesis that large herbivores support the persistence of firesensitive species through the creation of fire refugia in fire-prone landscapes, we examined the response of a fire-sensitive plant, Wyoming big sagebrush (Artemisia tridentata ssp. wyomingensis [Beetle & Young]) to fire and grazing in the fire-dependent mixed-grass prairie of the northern Great Plains. We carried out a controlled burn in 2010 within pre-established exclosures that allowed differential access to wild and domestic herbivores and no record of fire in the previous 75 years due to fire suppression efforts. The experiment was set up with a split-plot design to also examine potential changes in plots that were not burned. Canopy cover of big sagebrush was recorded before the burn in 2010 and again in 2011 with percent area burned recorded within 1-month post-fire in the burned plots. Percentage area burned was the greatest in ungulate exclosures (92% ± 2%) and the least in open areas (55% ± 21%), suggesting that large herbivores influenced fire behavior (e.g., reducing fire intensity and rate of spread) and are likely to increase fire patchiness through their alterations to the fuel bed. Regression analysis indicated that the proportion of sagebrush cover lost was significantly correlated with the proportion of area burned (𝑅² = 0.76, 𝑝 = 0.05). No differences in the non-burn plots were observed among grazing treatments or among years. Altogether, this illustrates the potential importance of large herbivores in creating biotic-driven fire refugia for fire-sensitive species to survive within the flammable fuel matrix of fire-dependent grassland ecosystems such as the mixed-grass prairie. Our findings also attest to the resiliency of the northern Great Plains to fire and herbivory and underscore the value of managing grasslands for heterogeneity with spatial and temporal variations in these historic disturbances.

Sagebrush is one of the most imperiled ecosystems in western North America, having lost about half of its original 62 million hectare extent. Annual grass invasions are known to be increasing wildfire occurrence and burned area, but the lasting effects (greater than five years post‐fire) that the resulting reburns have on these plant communities are unclear. We created a fire history atlas from 31 yr (1984–2014) of Landsat‐derived fire data to sample along a fire frequency gradient (zero to three fires) in an area of northern Nevada that has experienced frequent fire in this time period. Thirty‐two percent of our study area (13,000 km2) burned in large fires (over 404 ha) at least once, 7% burned twice, and 2% burned three or more times. We collected plant abundance data at 28 plots (N = 7 per fire frequency), with an average time since fire of 17 yr. We examined fire's effect on plant diversity using species accumulation curves, alpha diversity (Shannon's dominance, Pielou's evenness, and number of species), and beta diversity (Whittaker, Simpson, and Z indexes). For composition, we used non‐metric multidimensional scaling. We then used PERMANOVA models to examine how disturbance history, temperature, precipitation, and aridity around the time of the fire affected subsequent community composition and diversity. One fire fundamentally changed community composition and reduced species richness, and each subsequent fire reduced richness further. Alpha diversity decreased after one fire. Beta diversity declined after the third fire. Cover of exotics was 10% higher in all burned plots, and native cover was 20% lower than in unburned plots, regardless of frequency. PERMANOVA models showed fire frequency and antecedent precipitation as the strongest predictors of beta diversity, while time since fire and vapor pressure deficit for the year of the fire were the strongest predictors of community composition. Given that a single fire has such a marked effect on species composition, and repeated fires reduce richness and beta diversity, we suggest that in lower elevation big sagebrush systems fire should be minimized as much as possible, perhaps even prescribed fire. Restoration efforts should be focused on timing with wet years on cooler, wetter sites.

Repeated perturbations, both biotic and abiotic, can lead to fundamental changes in the nature of ecosystems, including changes in state. Sagebrush steppe communities provide important habitat for wildlife and grazing for livestock. Fire is an integral part of these systems, but there is concern that increased ignition frequencies and invasive species are fundamentally altering them. Despite these issues, the majority of studies of fire effects in systems dominated by Artemisia tridentata wyomingensis have focused on the effects of single burns. The Arid Lands Ecology Reserve (ALE), in south-central Washington (USA), was one of the largest contiguous areas of sagebrush steppe habitat in the state until large wildfires burned the majority of it in 2000 and 2007. We analyzed data from permanent vegetation transects established in 1996 and resampled in 2002 and 2009. Our objective was to describe how the fires, and subsequent postfire restoration efforts, affected communities' successional pathways. Plant communities differed in response to repeated fire and restoration; these differences could largely be ascribed to the functional traits of the dominant species. Low-elevation communities, previously dominated by obligate seeders, moved furthest from their initial composition and were dominated by weedy, early-successional species in 2009. Higher-elevation sites with resprouting shrubs, native bunchgrasses, and few invasive species were generally more resilient to the effects of repeated disturbances. Shrub cover has been almost entirely removed from ALE, although there was some recovery where communities were dominated by resprouters. Bromus tectorum dominance was reduced by herbicide application in areas where it was previously abundant, but it increased significantly in untreated areas. Several resprouting species, notably Phlox longifolia and Poa secunda , expanded remarkably following competitive release from shrub canopies and/or abundant B. tectorum . Our results suggest that community dynamics can be understood through a state and transition model with two axes (shrub/grass and native/invasive abundance), although such models also need to account for differences in plant functional traits and disturbance regimes. We use our results to develop a conceptual model that will be validated with further research.

We assessed plant community succession following prescribed fire on ungrazed Wyoming big sagebrush steppe, eastern Oregon. Treatments were burned (Burn; September and October, 2002) and unburned (Control) sagebrush steppe. Herbaceous yield, vegetation canopy cover and density were compared between treatments after fire (2003–18). Herbaceous yield in the Burn treatment was about double the Control for most of the study period. Prior to fire, native perennials comprised 90–95% of herbaceous yield. After fire, native perennials represented 78% (range 67–93%) and exotic annuals 22% (range 7–33%) of total yield. Exotic annuals increased after fire and responded in two stages. In the first 8 years after fire, desert alyssum dominated the annual plant composition. In the last half of the study, cheatgrass co-dominated the annual component with alyssum. Sagebrush recovery was slow and we estimated sagebrush cover would return to pre-burn levels, at the earliest, in 115 years. Burning Wyoming big sagebrush steppe would be detrimental to sagebrush-obligate wildlife for an extended time period, because of lost cover and structure provided by sagebrush. The additional forage provided on burned areas may give livestock manager’s greater flexibility to rest or defer unburned habitat for wildlife species of critical concern.

Fire historically occurred across the sagebrush steppe, but little is known about how patterns of post-fire fuel accumulation influence future fire in Wyoming big sagebrush (Artemisia tridentata ssp. wyomingensis) communities. To quantify change in fuel composition and structure in intact sagebrush ecosystems, we sampled 17 years following prescribed fire in eight approximately 400 ha plots (4 burned, 4 unburned control) at Hart Mountain National Antelope Refuge, OR, USA. Fuels data were used to model potential fire behavior in burn and control plots across four environmental scenarios that mimic drying of fuels through the fire season. Seventeen years after fire total fuel loads were 7 × higher in controls (6015 kg ha⁻¹) than burned plots (831 kg ha⁻¹; P < 0.01). Herbaceous fuels were 5 times greater in burns (P < 0. 01). Shrub fuel was nearly 10 times higher in unburned plots (P < 0.01), and litter under shrubs in controls was 3.75 times greater than in burns (P < 0.01). Potential fire behavior was lower in burned plots than in unburned controls across all environmental scenarios. In the driest scenario, potential rate of spread ranged from 0.4 to 1.5 m min⁻¹ in burns and 2.7 to 5.5 m min⁻¹ in controls (P < 0.01). Maintaining resilience in these ecosystems at multiple spatial and temporal scales may include a consideration of the natural role of fire in good condition Wyoming big sagebrush ecosystems. This study shows that under these conditions, fire can promote good condition mid-successional ecosystems and can act as a fuel break, slowing the spread and decreasing the intensity of a future wildfire.

Repeated perturbations, both biotic and abiotic, can lead to fundamental changes in the nature of ecosystems, including changes in state. Sagebrush steppe communities provide important habitat for wildlife and grazing for livestock. Fire is an integral part of these systems, but there is concern that increased ignition frequencies and invasive species are fundamentally altering them. Despite these issues, the majority of studies of fire effects in systems dominated by Artemisia tridentata wyomingensis have focused on the effects of single burns. The Arid Lands Ecology Reserve (ALE), in south-central Washington (USA), was one of the largest contiguous areas of sagebrush steppe habitat in the state until large wildfires burned the majority of it in 2000 and 2007. We analyzed data from permanent vegetation transects established in 1996 and resampled in 2002 and 2009. Our objective was to describe how the fires, and subsequent postfire restoration efforts, affected communities' successional pathways. Plant communities differed in response to repeated fire and restoration; these differences could largely be ascribed to the functional traits of the dominant species. Low-elevation communities, previously dominated by obligate seeders, moved furthest from their initial composition and were dominated by weedy, early-successional species in 2009. Higher-elevation sites with resprouting shrubs, native bunchgrasses, and few invasive species were generally more resilient to the effects of repeated disturbances. Shrub cover has been almost entirely removed from ALE, although there was some recovery where communities were dominated by resprouters. Bromus tectorum dominance was reduced by herbicide application in areas where it was previously abundant, but it increased significantly in untreated areas. Several resprouting species, notably Phlox longifolia and Poa secunda , expanded remarkably following competitive release from shrub canopies and/or abundant B. tectorum . Our results suggest that community dynamics can be understood through a state and transition model with two axes (shrub/grass and native/invasive abundance), although such models also need to account for differences in plant functional traits and disturbance regimes. We use our results to develop a conceptual model that will be validated with further research.

An increase in mega-fires and wildfires is a global issue that is expected to become worse with climate change. Fuel treatments are often recommended to moderate behaviour and decrease severity of wildfires; however, the extensive nature of rangelands limits the use of many treatments. Dormant-season grazing has been suggested as a rangeland fuel treatment, but its effects on fire characteristics are generally unknown. We investigated the influence of dormant-season (winter) grazing by cattle (Bos taurus) on fuel characteristics, fire behaviour and area burned in Wyoming big sagebrush (Artemisia tridentata subsp. wyomingensis) shrub-grassland communities in south-eastern Oregon, USA. Winter grazing was applied for 5 years before burning and compared with ungrazed areas. Winter grazing decreased fine fuels and increased fine fuel moisture, which reduced flame height and depth, rate of spread and area burned. Winter-grazed areas also had lower maximum temperature and heat loading during fires than ungrazed areas, and thereby decreased risk of fire-induced mortality of important herbaceous functional groups. These results suggest that winter grazing may be a fuel management treatment that can be applied across vast shrub-grasslands to decrease wildfire risk and fire intensity to mediate climate change effects on wildfire activity.

Background Wildfire is a major proximate cause of historical and ongoing losses of intact big sagebrush ( Artemisia tridentata Nutt.) plant communities and declines in sagebrush obligate wildlife species. In recent decades, fire return intervals have shortened and area burned has increased in some areas, and habitat degradation is occurring where post-fire re-establishment of sagebrush is hindered by invasive annual grasses. In coming decades, the changing climate may accelerate these wildfire and invasive feedbacks, although projecting future wildfire dynamics requires a better understanding of long-term wildfire drivers across the big sagebrush region. Here, we integrated wildfire observations with climate and vegetation data to derive a statistical model for the entire big sagebrush region that represents how annual wildfire probability is influenced by climate and fine fuel characteristics. Results Wildfire frequency varied significantly across the sagebrush region, and our statistical model represented much of that variation. Biomass of annual and perennial grasses and forbs, which we used as proxies for fine fuels, influenced wildfire probability. Wildfire probability was highest in areas with high annual forb and grass biomass, which is consistent with the well-documented phenomenon of increased wildfire following annual grass invasion. The effects of annuals on wildfire probability were strongest in places with dry summers. Wildfire probability varied with the biomass of perennial grasses and forbs and was highest at intermediate biomass levels. Climate, which varies substantially across the sagebrush region, was also predictive of wildfire probability, and predictions were highest in areas with a low proportion of precipitation received in summer, intermediate precipitation, and high temperature. Conclusions We developed a carefully validated model that contains relatively simple and biologically plausible relationships, with the goal of adequate performance under novel conditions so that useful projections of average annual wildfire probability can be made given general changes in conditions. Previous studies on the impacts of vegetation and climate on wildfire probability in sagebrush ecosystems have generally used more complex machine learning approaches and have usually been applicable to only portions of the sagebrush region. Therefore, our model complements existing work and forms an additional tool for understanding future wildfire and ecological dynamics across the sagebrush region.