Arbuscular Mycorrhizae Alter Photosynthetic Responses to Drought in Seedlings of Artemisia tridentata.

The establishment of Artemisia tridentata, a keystone species of the sagebrush steppe, is often limited by summer drought. Symbioses with arbuscular mycorrhizal fungi (AMF) can help plants to cope with drought. We investigated this possible effect on A. tridentata seedlings inoculated with native AMF and exposed to drought in greenhouse and field settings. In greenhouse experiments, AMF colonization increased intrinsic water use efficiency under water stress and delayed the decrease in photosynthesis caused by drought, or this decrease occurred at a lower soil water content. In the field, we evaluated the effect of AMF inoculation on colonization, leaf water potential, survival, and inflorescence development. Inoculation increased AMF colonization, and the seedlings experienced water stress, as evidenced by water potentials between -2 and -4 MPa and reduced stomatal conductance. However, survival remained high, and no differences in water potentials or survival occurred between treatments. Only the percentage of plants with inflorescence was higher in inoculated than non-inoculated seedlings. Overall, the greenhouse results support that AMF colonization enhances drought tolerance in A. tridentata seedlings. Yet, the significance of these results in increasing survival in nature remains to be tested under more severe drought than the plants experienced in our field experiment.

A genotype × environment experiment reveals contrasting response strategies to drought between populations of a keystone species (Artemisia tridentata; Asteraceae).

Western North America has been experiencing persistent drought exacerbated by climate change for over two decades. This extreme climate event is a clear threat to native plant communities. Artemisia tridentata is a keystone shrub species in western North America and is threatened by climate change, urbanization, and wildfire. A drought Genotype × Environment (G × E) experiment was conducted to assess phenotypic plasticity and differential gene expression in A. tridentata. The G × E experiment was performed on diploid A. tridentata seedlings from two populations (one from Idaho, USA and one from Utah, USA), which experience differing levels of drought stress during the summer months. Photosynthetic data, leaf temperature, and gene expression levels were compared between treatments and populations. The Utah population maintained higher photosynthetic rates and photosynthetic efficiency than the Idaho population under drought stress. The Utah population also exhibited far greater transcriptional plasticity than the Idaho population and expressed genes of response pathways distinct from those of the Idaho population. Populations of A. tridentata differ greatly in their drought response pathways, likely due to differences in response pathways that have evolved under distinct climatic regimes. Epigenetic processes likely contribute to the observed differences between the populations.

Acclimation and hardening of a slow-growing woody species emblematic to western North America from in vitro plantlets.

Premise: Determining the tolerance of plant populations to climate change requires the development of biotechnological protocols producing genetically identical individuals used for genotype-by-environment experiments. Such protocols are missing for slow-growth, woody plants; to address this gap, this study uses Artemisia tridentata, a western North American keystone shrub, as model. Methods and Results: The production of individual lines is a two-step process: in vitro propagation under aseptic conditions followed by ex vitro acclimation and hardening. Due to aseptic growth conditions, in vitro plantlets exhibit maladapted phenotypes, and this protocol focuses on presenting an approach promoting morphogenesis for slow-growth, woody species. Survival was used as the main criterion determining successful acclimation and hardening. Phenotypic changes were confirmed by inspecting leaf anatomy, and shoot water potential was used to ensure that plantlets were not water stressed. Conclusions: Although our protocol has lower survival rates (11-41%) compared to protocols developed for herbaceous, fast-growing species, it provides a benchmark for slow-growth, woody species occurring in dry ecosystems.

Fuel connectivity, burn severity, and seed bank survivorship drive ecosystem transformation in a semiarid shrubland.

A key challenge in ecology is understanding how multiple drivers interact to precipitate persistent vegetation state changes. These state changes may be both precipitated and maintained by disturbances, but predicting whether the state change will be fleeting or persistent requires an understanding of the mechanisms by which disturbance affects the alternative communities. In the sagebrush shrublands of the western United States, widespread annual grass invasion has increased fuel connectivity, which increases the size and spatial contiguity of fires, leading to postfire monocultures of introduced annual grasses (IAG). The novel grassland state can be persistent and is more likely to promote large fires than the shrubland it replaced. But the mechanisms by which prefire invasion and fire occurrence are linked to higher postfire flammability are not fully understood. A natural experiment to explore these interactions presented itself when we arrived in northern Nevada immediately after a 50,000 ha wildfire was extinguished. We hypothesized that the novel grassland state is maintained via a reinforcing feedback where higher fuel connectivity increases burn severity, which subsequently increases postfire IAG dispersal, seed survivorship, and fuel connectivity. We used a Bayesian joint species distribution model and structural equation model framework to assess the strength of the support for each element in this feedback pathway. We found that prefire fuel connectivity increased burn severity and that higher burn severity had mostly positive effects on the occurrence of IAG and another nonnative species and mostly negative or neutral relationships with all other species. Finally, we found that the abundance of IAG seeds in the seed bank immediately after a fire had a positive effect on the fuel connectivity 3 years after the fire, completing a positive feedback promoting IAG. These results demonstrate that the strength of the positive feedback is controlled by measurable characteristics of ecosystem structure, composition, and disturbance. Further, each node in the loop is affected independently by multiple global change drivers. It is possible that these characteristics can be modeled to predict threshold behavior and inform management actions to mitigate or slow the establishment of the grass-fire cycle, perhaps via targeted restoration applications or prefire fuel treatments.

Herbivory Amplifies Adverse Effects of Drought on Seedling Recruitment in a Keystone Species of Western North American Rangelands.

Biotic interactions can affect a plant's ability to withstand drought. Such an effect may impact the restoration of the imperiled western North American sagebrush steppe, where seedlings are exposed to summer drought. This study investigated the impact of herbivory on seedlings' drought tolerance for a keystone species in this steppe, the shrub Artemisia tridentata. Herbivory effects were investigated in two field experiments where seedlings were without tree protectors or within plastic or metal-mesh tree protectors. Treatment effects were statistically evaluated on herbivory, survival, leaf water potential, and inflorescence development. Herbivory occurrence was 80% higher in seedlings without protectors. This damage occurred in early spring and was likely caused by ground squirrels. Most plants recovered, but herbivory was associated with higher mortality during the summer when seedlings experienced water potentials between -2.5 and -7 MPa. However, there were no differences in water potential between treatments, suggesting that the browsed plants were less tolerant of the low water potentials experienced. Twenty months after outplanting, the survival of plants without protectors was 40 to 60% lower than those with protectors. The percentage of live plants developing inflorescences was approximately threefold higher in plants with protectors. Overall, spring herbivory amplified susceptibility to drought and delayed reproductive development.

Small effects of livestock grazing intensification on diversity, abundance, and composition in a dryland plant community.

Livestock grazing is a globally important land use and has the potential to significantly influence plant community structure and ecosystem function, yet several critical knowledge gaps remain on the direction and magnitude of grazing impacts. Furthermore, much of our understanding of the long-term effects on plant community composition and structure are based on grazer exclusion experiments, which explicitly avoid characterizing effects along grazing intensity gradients. We sampled big sagebrush plant communities using 68 plots located along grazing intensity gradients to determine how grazing intensity influences multiple aspects of plant community structure over time. This was accomplished by sampling plant communities at different distances from 17 artificial watering sources, using distance from water and cow dung density as proxies for grazing intensity at individual plots. Total vegetation cover and total grass cover were negatively related to grazing intensity, and cover of annual forbs, exotic cover, and exotic richness were positively related to grazing intensity. In contrast, species richness and composition, bunchgrass biomass, shrub density and size, percentage cover of bare ground, litter, and biological soil crusts did not vary along our grazing intensity gradients, in spite of our expectations to the contrary. Our results suggest that the effects of livestock grazing over multiple decades (mean = 46 years) in our sites are relatively small, especially for native perennial species, and that the big sagebrush plant communities we sampled are somewhat resistant to livestock grazing. Collectively, our findings are consistent with existing evidence that indicates the stability of the big sagebrush plant functional type composition under current grazing management regimes.

Essential Oil Yield, Composition, and Bioactivity of Sagebrush Species in the Bighorn Mountains.

Sagebrush (Artemisia spp.) are dominant wild plants in large areas of the U.S., Canada and Mexico, and they include several species and subspecies. The aim was to determine if there are significant differences in essential oil (EO) yield, composition, and biological activity of sagebrush within the Bighorn Mountains, U.S. The EO yield in fresh herbage varied from 0.15 to 1.69% for all species, including 0.25-1.69% in A. tridentata var. vaseyana, 0.64-1.44% in A. tridentata var. tridentata, 1% in A. tridentata var. wyomingensis, 0.8-1.2% in A. longifolia, 0.8-1% in A. cana, and 0.16% in A. ludoviciana. There was significant variability in the EO profile between species, and subspecies. Some EO constituents, such as α-pinene (0-35.5%), camphene (0-21.5%), eucalyptol (0-30.8%), and camphor (0-45.5%), were found in most species and varied with species and subspecies. The antioxidant capacity of the EOs varied between the species and subspecies. None of the sagebrush EOs had significant antimicrobial, antimalarial, antileishmanial activity, or contained podophyllotoxin. Some accessions yielded EO with significant concentrations of compounds including camphor, eucalyptol, cis-thujone, α-pinene, α-necrodol-acetate, fragranol, grandisol, para-cymene, and arthole. Therefore, chemotypes can be selected and possibly introduced into culture and be grown for commercial production of these compounds to meet specific industry needs.

A haploid pseudo-chromosome genome assembly for a keystone sagebrush species of western North American rangelands.

Increased ecological disturbances, species invasions, and climate change are creating severe conservation problems for several plant species that are widespread and foundational. Understanding the genetic diversity of these species and how it relates to adaptation to these stressors are necessary for guiding conservation and restoration efforts. This need is particularly acute for big sagebrush (Artemisia tridentata; Asteraceae), which was once the dominant shrub over 1,000,000 km2 in western North America but has since retracted by half and thus has become the target of one of the largest restoration seeding efforts globally. Here, we present the first reference-quality genome assembly for an ecologically important subspecies of big sagebrush (A. tridentata subsp. tridentata) based on short and long reads, as well as chromatin proximity ligation data analyzed using the HiRise pipeline. The final 4.2-Gb assembly consists of 5,492 scaffolds, with nine pseudo-chromosomal scaffolds (nine scaffolds comprising at least 90% of the assembled genome; n = 9). The assembly contains an estimated 43,377 genes based on ab initio gene discovery and transcriptional data analyzed using the MAKER pipeline, with 91.37% of BUSCOs being completely assembled. The final assembly was highly repetitive, with repeat elements comprising 77.99% of the genome, making the Artemisia tridentata subsp. tridentata genome one of the most highly repetitive plant genomes to be sequenced and assembled. This genome assembly advances studies on plant adaptation to drought and heat stress and provides a valuable tool for future genomic research.

Interannual climate variability mediates changes in carbon and nitrogen pools caused by annual grass invasion in a semiarid shrubland.

Exotic plant invasions alter ecosystem properties and threaten ecosystem functions globally. Interannual climate variability (ICV) influences both plant community composition (PCC) and soil properties, and interactions between ICV and PCC may influence nitrogen (N) and carbon (C) pools. We asked how ICV and non-native annual grass invasion covary to influence soil and plant N and C in a semiarid shrubland undergoing widespread ecosystem transformation due to invasions and altered fire regimes. We sampled four progressive stages of annual grass invasion at 20 sites across a large (25,000 km2 ) landscape for plant community composition, plant tissue N and C, and soil total N and C in 2013 and 2016, which followed 2 years of dry and wet conditions, respectively. Multivariate analyses and ANOVAs showed that in invasion stages where native shrub and perennial grass and forb communities were replaced by annual grass-dominated communities, the ecosystem lost more soil N and C in wet years. Path analysis showed that high water availability led to higher herbaceous cover in all invasion stages. In stages with native shrubs and perennial grasses, higher perennial grass cover was associated with increased soil C and N, while in annual-dominated stages, higher annual grass cover was associated with losses of soil C and N. Also, soil total C and C:N ratios were more homogeneous in annual-dominated invasion stages as indicated by within-site standard deviations. Loss of native shrubs and perennial grasses and forbs coupled with annual grass invasion may lead to long-term declines in soil N and C and hamper restoration efforts. Restoration strategies that use innovative techniques and novel species to address increasing temperatures and ICV and emphasize maintaining plant community structure-shrubs, grasses, and forbs-will allow sagebrush ecosystems to maintain C sequestration, soil fertility, and soil heterogeneity.

Risk of herbivory negatively correlates with the diversity of volatile emissions involved in plant communication.

Plant-to-plant volatile-mediated communication and subsequent induced resistance to insect herbivores is common. Less clear is the adaptive significance of these interactions; what selective mechanisms favour plant communication and what conditions allow individuals to benefit by both emitting and responding to cues? We explored the predictions of two non-exclusive hypotheses to explain why plants might emit cues, the kin selection hypothesis (KSH) and the mutual benefit hypothesis (MBH). We examined 15 populations of sagebrush that experience a range of naturally occurring herbivory along a 300 km latitudinal transect. As predicted by the KSH, we found several uncommon chemotypes with some chemotypes occurring only within a single population. Consistent with the MBH, chemotypic diversity was negatively correlated with herbivore pressure; sites with higher levels of herbivory were associated with a few common cues broadly recognized by most individuals. These cues varied among different populations. Our results are similar to those reported for anti-predator signalling in vertebrates.

The role of genome duplication in big sagebrush growth and fecundity.

PREMISE: Adaptive traits can be dramatically altered by genome duplication. The study of interactions among traits, ploidy, and the environment are necessary to develop an understanding of how polyploidy affects niche differentiation and to develop restoration strategies for resilient native ecosystems. METHODS: Growth and fecundity were measured in common gardens for 39 populations of big sagebrush (Artemisia tridentata) containing two subspecies and two ploidy levels. General linear mixed-effect models assessed how much of the trait variation could be attributed to genetics (i.e., ploidy and climatic adaptation), environment, and gene-environment interactions. RESULTS: Growth and fecundity variation were explained well by the mixed models (80% and 91%, respectively). Much of the trait variation was attributed to environment, and 15% of variation in growth and 34% of variation in seed yield were attributed to genetics. Genetic trait variation was mostly attributable to ploidy, with much higher growth and seed production in diploids, even in a warm-dry environment typically dominated by tetraploids. Population-level genetic variation was also evident and was related to the climate of each population's origin. CONCLUSIONS: Ploidy is a strong predictor growth and seed yield, regardless of common-garden environment. The superior growth and fecundity of diploids across environments raises the question as to how tetraploids can be more prevalent than diploids, especially in warm-dry environments. Two hypotheses that may explain the abundance of tetraploids on the landscape include selection for drought resistance at the seedling stage, and greater competitive ability in water uptake in the upper soil horizon.

Intraspecific variation mediates density dependence in a genetically diverse plant species.

Interactions between neighboring plants are critical for biodiversity maintenance in plant populations and communities. Intraspecific trait variation and genome duplication are common in plant species and can drive eco-evolutionary dynamics through genotype-mediated plant-plant interactions. However, few studies have examined how species-wide intraspecific variation may alter interactions between neighboring plants. We investigate how subspecies and ploidy variation in a genetically diverse species, big sagebrush (Artemisia tridentata), can alter the demographic outcomes of plant interactions. Using a replicated, long-term common garden experiment that represents range-wide diversity of A. tridentata, we ask how intraspecific variation, environment, and stand age mediate neighbor effects on plant growth and survival. Spatially explicit models revealed that ploidy variation and subspecies identity can mediate plant-plant interactions but that the effect size varied in time and across experimental sites. We found that demographic impacts of neighbor effects were strongest during early stages of stand development and in sites with greater growth rates. Within subspecies, tetraploid populations showed greater tolerance to neighbor crowding compared to their diploid variants. Our findings provide evidence that intraspecific variation related to genome size and subspecies identity impacts spatial demography in a genetically diverse plant species. Accounting for intraspecific variation in studies of conspecific density dependence will improve our understanding of how local populations will respond to novel genotypes and biotic interaction regimes. As introduction of novel genotypes into local populations becomes more common, quantifying demographic processes in genetically diverse populations will help predict long-term consequences of plant-plant interactions.

Divergent climate change effects on widespread dryland plant communities driven by climatic and ecohydrological gradients.

Plant community response to climate change will be influenced by individual plant responses that emerge from competition for limiting resources that fluctuate through time and vary across space. Projecting these responses requires an approach that integrates environmental conditions and species interactions that result from future climatic variability. Dryland plant communities are being substantially affected by climate change because their structure and function are closely tied to precipitation and temperature, yet impacts vary substantially due to environmental heterogeneity, especially in topographically complex regions. Here, we quantified the effects of climate change on big sagebrush (Artemisia tridentata Nutt.) plant communities that span 76 million ha in the western United States. We used an individual-based plant simulation model that represents intra- and inter-specific competition for water availability, which is represented by a process-based soil water balance model. For dominant plant functional types, we quantified changes in biomass and characterized agreement among 52 future climate scenarios. We then used a multivariate matching algorithm to generate fine-scale interpolated surfaces of functional type biomass for our study area. Results suggest geographically divergent responses of big sagebrush to climate change (changes in biomass of -20% to +27%), declines in perennial C3 grass and perennial forb biomass in most sites, and widespread, consistent, and sometimes large increases in perennial C4 grasses. The largest declines in big sagebrush, perennial C3 grass and perennial forb biomass were simulated in warm, dry sites. In contrast, we simulated no change or increases in functional type biomass in cold, moist sites. There was high agreement among climate scenarios on climate change impacts to functional type biomass, except for big sagebrush. Collectively, these results suggest divergent responses to warming in moisture-limited versus temperature-limited sites and potential shifts in the relative importance of some of the dominant functional types that result from competition for limiting resources.

Time and Chance: Using Age Partitioning to Understand How Luck Drives Variation in Reproductive Success.

AbstractOver the course of individual lifetimes, luck usually explains a large fraction of the between-individual variation in life span or lifetime reproductive output (LRO) within a population, while variation in individual traits or "quality" explains much less. To understand how, where in the life cycle, and through which demographic processes luck trumps trait variation, we show how to partition by age the contributions of luck and trait variation to LRO variance and how to quantify three distinct components of luck. We apply these tools to several empirical case studies. We find that luck swamps effects of trait variation at all ages, primarily because of randomness in individual state dynamics ("state trajectory luck"). Luck early in life is most important. Very early state trajectory luck generally determines whether an individual ever breeds, likely by ensuring that they are not dead or doomed quickly. Less early luck drives variation in success among those breeding at least once. Consequently, the importance of luck often has a sharp peak early in life or it has two peaks. We suggest that ages or stages where the importance of luck peaks are potential targets for interventions to benefit a population of concern, different from those identified by eigenvalue elasticity analysis.

Soil and stand structure explain shrub mortality patterns following global change-type drought and extreme precipitation.

The probability of extreme weather events is increasing, with the potential for widespread impacts to plants, plant communities, and ecosystems. Reports of drought-related tree mortality are becoming more frequent, and there is increasing evidence that drought accompanied by high temperatures is especially detrimental. Simultaneously, extreme large precipitation events have become more frequent over the past century. Water-limited ecosystems may be more vulnerable to these extreme events than other ecosystems, especially when pushed outside of their historical range of variability. However, drought-related mortality of shrubs-an important component of dryland vegetation-remains understudied relative to tree mortality. In 2014, a landscape-scale die-off of the widespread shrub, big sagebrush (Artemisia tridentata Nutt.), was reported in southwest Wyoming, following extreme hot and dry conditions in 2012 and extremely high precipitation in September of 2013. Here we examine how severe drought, extreme precipitation, soil texture and salinity, and shrub-stand characteristics contributed to this die-off event. At 98 plots within and around the die-off, we quantified big sagebrush mortality, characterized soil texture and salinity, and simulated soil-water conditions from 1916 to 2016 using an ecosystem water-balance model. We found that the extreme weather conditions alone did not explain patterns of big sagebrush mortality and did not result in extreme (historically unprecedented) soil-water conditions during the drought. Instead, plots with chronically dry soil conditions experienced greatest mortality following the global change-type (hot) drought in 2012. Furthermore, mortality was greater in locations with high potential run-on and low potential run-off where saturated soil conditions were simulated in September 2013, suggesting that extreme precipitation also played an important role in the die-off in these locations. In locations where drought alone contributed to mortality, stem density negatively impacted big sagebrush. In locations that may have been affected by both drought and saturation, however, mortality was greatest where stem density was lowest, suggesting that these locations may have already been less favorable to big sagebrush. Paradoxically, vulnerability to both extreme events (drought and saturation) was associated with finer-textured soils, and our results highlight the importance of soils in determining local variation of the vulnerability of dryland plants to extreme events.

Freezing resistance, safety margins, and survival vary among big sagebrush populations across the western United States.

PREMISE: Physiological responses to temperature extremes are considered strong drivers of species' demographic responses to climate variability. Plants are typically classified as either avoiders or tolerators in their freezing-resistance mechanism, but a gradient of physiological-threshold freezing responses may exist among individuals of a species. Moreover, adaptive significance of physiological freezing responses is poorly characterized, particularly under warming conditions that relax selection on cold hardiness. METHODS: Freezing responses were measured in winter and again for new foliage in spring for 14 populations of Artemisia tridentata collected throughout its range and planted in a warm common garden. The relationships of the freezing responses to survival were evaluated in the warm garden and in two colder gardens. RESULTS: Winter and spring freezing resistance were not correlated and appeared to be under differing selection regimes, as evident in correlations with different population climate of origin variables. All populations resisted considerably lower temperatures in winter than in spring, with populations from more continental climates showing narrower freezing safety margins (difference in temperatures at which ice-nucleation occurs and 50% reduction in chlorophyll fluorescence occurs) in spring. Populations with greater winter freezing resistance had lower survivorship in the warmest garden, while populations with greater spring freezing resistance had lower survivorship in a colder garden. CONCLUSIONS: These survivorship patterns relative to physiological thresholds suggest excess freezing resistance may incur a survival cost that likely relates to a trade-off between carbon gain and freezing resistance during critical periods of moisture availability. This cost has implications for seed moved from cooler to warmer environments and for plants growing in warming environments.

Soil texture and precipitation seasonality influence plant community structure in North American temperate shrub steppe.

In drylands, the coexistence of grasses and woody plants has been attributed to soil-water resource partitioning. Soil texture and precipitation seasonality can influence the amount and distribution of water in the soil, and their interaction may play an important role in determining the relative importance of grasses and woody plants. We investigated the influence of this interaction on plant functional types across a broad range of precipitation regimes and soil textures in western North America by analyzing plant-cover data collected at 2,084 plots that included the widespread shrub big sagebrush (Artemisia tridentata Nutt.). We characterized how the significance of the inverse-texture effect varies across soil conditions by quantifying relationships between precipitation and foliar cover on finer- vs. coarser-textured soils across a range of potential texture divisions represented by sand content. We found evidence of the inverse-texture effect for every plant functional type (except for cheatgrass) that we examined with at least one component of precipitation (annual, warm, or cold season), and provide the first evidence for this effect in locations with cold-season-dominated precipitation regimes. The texture and precipitation combinations that exhibited the inverse-texture effect varied with plant functional type, presumably because of effects of soil texture on water availability at different soil depths with season. Furthermore, we found an inverse-texture effect that was remarkably similar for shrub cover with cold-season precipitation and grass cover with warm-season precipitation. These results provide new insight into how the inverse-texture effect interacts with precipitation seasonality to influence plant functional type composition in drylands, and further suggest that quantifying the soil-texture division at which the inverse-texture effect is relevant under a given set of environmental conditions may provide support for the effect across dryland plant communities.

Integrating anthropogenic factors into regional-scale species distribution models-A novel application in the imperiled sagebrush biome.

Species distribution models (SDMs) that rely on regional-scale environmental variables will play a key role in forecasting species occurrence in the face of climate change. However, in the Anthropocene, a number of local-scale anthropogenic variables, including wildfire history, land-use change, invasive species, and ecological restoration practices can override regional-scale variables to drive patterns of species distribution. Incorporating these human-induced factors into SDMs remains a major research challenge, in part because spatial variability in these factors occurs at fine scales, rendering prediction over regional extents problematic. Here, we used big sagebrush (Artemisia tridentata Nutt.) as a model species to explore whether including human-induced factors improves the fit of the SDM. We applied a Bayesian hurdle spatial approach using 21,753 data points of field-sampled vegetation obtained from the LANDFIRE program to model sagebrush occurrence and cover by incorporating fire history metrics and restoration treatments from 1980 to 2015 throughout the Great Basin of North America. Models including fire attributes and restoration treatments performed better than those including only climate and topographic variables. Number of fires and fire occurrence had the strongest relative effects on big sagebrush occurrence and cover, respectively. The models predicted that the probability of big sagebrush occurrence decreases by 1.2% (95% CI: -6.9%, 0.6%) when one fire occurs and cover decreases by 44.7% (95% CI: -47.9%, -41.3%) if at least one fire occurred over the 36 year period of record. Restoration practices increased the probability of big sagebrush occurrence but had minimal effect on cover. Our results demonstrate the potential value of including disturbance and land management along with climate in models to predict species distributions. As an increasing number of datasets representing land-use history become available, we anticipate that our modeling framework will have broad relevance across a range of biomes and species.

Shrub facilitation of tree establishment varies with ontogenetic stage across environmental gradients.

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.

Polyphenols from the sagebrush Artemisia tridentata ssp. tridentata affect the redox state of cultured hepatocytes by direct and indirect mechanisms.

Basin big sagebrush (Artemisia tridentata Nutt. ssp. tridentata (Asteraceae)), is a widespread North American shrub which produces a variety of polyphenolic compounds. Although sagebrush has been used as a traditional remedy by natives and settlers to the region, the polyphenols in Artemisia tridentata ssp. tridentata have not been highly investigated for their bioactive properties. To determine whether these polyphenols affect the intracellular redox state, we measured their ability to neutralize radicals in vitro and in a human liver carcinoma cell line (HepG2), and their effects on intracellular glutathione levels. Extracts from Artemisia tridentata ssp. tridentata decreased the oxidation of 2'7'-dichlorofluorescin in vitro and in cultured cells. Cells treated with polyphenolic extracts showed increased levels of glutathione in a time and dose-dependent manner. Approximately 48 polyphenolic compounds were distinguishable in extracts, by HPLC/UV absorbance detection. Mass spectroscopy was used to identify thirteen compounds as aesculin, aesculetin, apigenin, apigenin-7-O-glucoside, axillarin, casticin, chlorogenic acid, isoscopoletin, kaempferol, luteolin, methyl-axillarin, quercetin, and scopoletin. These results indicate that polyphenols produced in Artemisia tridentata ssp. tridentata affect the redox state of living cells by multiple mechanisms.