Isoscapes of remnant and restored Hawaiian montane forests reveal differences in biological nitrogen fixation and carbon inputs.

Deforestation and subsequent land-use conversion has altered ecosystems and led to negative effects on biodiversity. To ameliorate these effects, nitrogen-fixing (N2-fixing) trees are frequently used in the reforestation of degraded landscapes, especially in the tropics; however, their influence on ecosystem properties such as nitrogen (N) availability and carbon (C) stocks are understudied. Here, we use a 30-y old reforestation site of outplanted native N2-fixing trees (Acacia koa) dominated by exotic grass understory, and a neighboring remnant forest dominated by A. koa canopy trees and native understory, to assess whether restoration is leading to similar N and C biogeochemical landscapes and soil and plant properties as a target remnant forest ecosystem. We measured nutrient contents and isotope values (δ15N, δ13C) in soils, A. koa, and non-N2-fixing understory plants (Rubus spp.) and generated δ15N and δ13C isoscapes of the two forests to test for (1) different levels of biological nitrogen fixation (BNF) and its contribution to non-N2-fixing understory plants, and (2) the influence of historic land conversion and more recent afforestation on plant and soil δ13C. In the plantation, A. koa densities were higher and foliar δ15N values for A. koa and Rubus spp. were lower than in the remnant forest. Foliar and soil isoscapes also showed a more homogeneous distribution of low δ15N values in the plantation and greater influence of A. koa on neighboring plants and soil, suggesting greater BNF. Foliar δ13C also indicated higher water use efficiency (WUEi) in the plantation, indicative of differences in plant-water relations or soil water status between the two forest types. Plantation soil δ13C was higher than the remnant forest, consistent with greater contributions of exotic C4-pasture grasses to soil C pools, possibly due to facilitation of non-native grasses by the dense A. koa canopy. These findings are consequential for forest restoration, as they contribute to the mounting evidence that outplanting N2-fixing trees produces different biogeochemical landscapes than those observed in reference ecosystems, thereby influencing plant-soil interactions which can influence restoration outcomes.

Patterns and Distribution of Botryosphaeriaceae Fungi Related to Dieback in Big Berry Manzanita.

Dieback and mortality in wildland plant species due to climate change have been on the rise in recent decades, and latent fungal pathogens might play a significant role in these events. During a severe multiyear drought, canopy dieback associated with latent pathogens in the Botryosphaeriaceae (Bot) family was observed in stands of a dominant shrub species, big berry manzanita (Arctostaphylos glauca), across chaparral landscapes in California. These fungi are significant pathogens of woody agricultural species, especially in hosts experiencing stress, and have become a threat to economically important crops worldwide. However, little is known regarding their occurrence, distribution, and impact in wildland systems. We conducted a field survey of 300 A. glauca shrubs across an elevational gradient to identify Bot species infection as it relates to (i) A. glauca dieback severity and (ii) landscape variables associated with plant drought stress. Our results show that Bots are widely infecting A. glauca across the landscape, and there is a significant correlation between elevation and dieback severity. Dieback severity was significantly higher at lower elevations, suggesting that infected shrubs at lower elevations are at greater risk than those at higher elevations. Furthermore, two Bot species, Neofusicoccum australe and Botryosphaeria dothidea, were most frequently isolated, with N. australe being the most common and, based on haplotype analysis, likely the most recently introduced of the two. Our results confirm the wide distribution of latent Bot fungi in a wild shrubland system and provide valuable insight into areas of greatest risk for future shrub dieback and mortality. These findings could be particularly useful for informing future wildlands management strategies with regard to introduced latent pathogens.

Can the impact of canopy trees on soil and understory be altered using litter additions?

Trees can have large effects on soil nutrients in ways that alter succession, particularly in the case of nitrogen-(N)-fixing trees. In Hawai'i, forest restoration relies heavily on use of a native N-fixing tree, Acacia koa (koa), but this species increases soil-available N and likely facilitates competitive dominance of exotic pasture grasses. In contrast, Metrosideros polymorpha ('ohi'a), the dominant native tree in Hawai'i, is less often planted because it is slow growing; yet it is typically associated with lower soil N and grass biomass, and greater native understory recruitment. We experimentally tested whether it is possible to reverse high soil N under koa by adding 'ohi'a litter, using additions of koa litter or no litter as controls, over 2.5 yr. We then quantified natural litterfall and decomposition rates of 'ohi'a and koa litter to place litter additions in perspective. Finally, we quantified whether litter additions altered grass biomass and if this had effects on native outplants. Adding 'ohi'a litter increased soil carbon, but increased rather than decreased inorganic soil N pools. Contrary to expectations, koa litter decomposed more slowly than 'ohi'a, although it released more N per unit of litter. We saw no reduction in grass biomass due to 'ohi'a litter addition, and no change in native outplanted understory survival or growth. We conclude that the high N soil conditions under koa are difficult to reverse. However, we also found that outplanted native woody species were able to decrease exotic grass biomass over time, regardless of the litter environment, making this a better strategy for lowering exotic species impacts.

Mechanisms of severe dieback and mortality in a classically drought-tolerant shrubland species (Arctostaphylos glauca).

PREMISE: Mortality events involving drought and pathogens in natural plant systems are on the rise due to global climate change. In Santa Barbara, California, United States, big berry manzanita (Arctostaphylos glauca) has experienced canopy dieback related to a multi-year drought and infection from fungal pathogens in the Botryosphaeriaceae family. A greenhouse experiment was conducted using Neofusicoccum australe to test the specific influences of drought and fungal infection on A. glauca. METHODS: A full factorial design was used to compare four treatment groups (drought + inoculation; drought - inoculation; watering + inoculation; and control: watering - inoculation). Data were collected for 10 weeks on stress symptoms, changes in leaf fluorescence and photosynthesis, and mortality. RESULTS: Results indicated significant effects of watering and inoculation treatments on net photosynthesis, dark-adapted fluorescence, and disease symptom severity (P < 0.05), and a strong correlation was found between physiological decline and visible stress (P < 0.0001). Mortality differed between treatments, with all groups except for the control experiencing mortality (43% mortality in drought - inoculation, 83% in watering - inoculation, and 100% in drought + inoculation). A Kaplan-Meier survival analysis showed drought + inoculation to have the least estimated survivorship compared to all other treatment groups. CONCLUSIONS: In addition to a possible synergistic interaction between drought and fungal infection in disease onset and mortality rates in A. glauca, these results indicate that young, non-drought-stressed plants are susceptible to mortality from N. australe infection, with important implications for the future of wildland shrub communities.

Change in dominance determines herbivore effects on plant biodiversity.

Herbivores alter plant biodiversity (species richness) in many of the world's ecosystems, but the magnitude and the direction of herbivore effects on biodiversity vary widely within and among ecosystems. One current theory predicts that herbivores enhance plant biodiversity at high productivity but have the opposite effect at low productivity. Yet, empirical support for the importance of site productivity as a mediator of these herbivore impacts is equivocal. Here, we synthesize data from 252 large-herbivore exclusion studies, spanning a 20-fold range in site productivity, to test an alternative hypothesis-that herbivore-induced changes in the competitive environment determine the response of plant biodiversity to herbivory irrespective of productivity. Under this hypothesis, when herbivores reduce the abundance (biomass, cover) of dominant species (for example, because the dominant plant is palatable), additional resources become available to support new species, thereby increasing biodiversity. By contrast, if herbivores promote high dominance by increasing the abundance of herbivory-resistant, unpalatable species, then resource availability for other species decreases reducing biodiversity. We show that herbivore-induced change in dominance, independent of site productivity or precipitation (a proxy for productivity), is the best predictor of herbivore effects on biodiversity in grassland and savannah sites. Given that most herbaceous ecosystems are dominated by one or a few species, altering the competitive environment via herbivores or by other means may be an effective strategy for conserving biodiversity in grasslands and savannahs globally.

Boom-bust dynamics in biological invasions: towards an improved application of the concept.

Boom-bust dynamics - the rise of a population to outbreak levels, followed by a dramatic decline - have been associated with biological invasions and offered as a reason not to manage troublesome invaders. However, boom-bust dynamics rarely have been critically defined, analyzed, or interpreted. Here, we define boom-bust dynamics and provide specific suggestions for improving the application of the boom-bust concept. Boom-bust dynamics can arise from many causes, some closely associated with invasions, but others occurring across a wide range of ecological settings, especially when environmental conditions are changing rapidly. As a result, it is difficult to infer cause or predict future trajectories merely by observing the dynamic. We use tests with simulated data to show that a common metric for detecting and describing boom-bust dynamics, decline from an observed peak to a subsequent trough, tends to severely overestimate the frequency and severity of busts, and should be used cautiously if at all. We review and test other metrics that are better suited to describe boom-bust dynamics. Understanding the frequency and importance of boom-bust dynamics requires empirical studies of large, representative, long-term data sets that use clear definitions of boom-bust, appropriate analytical methods, and careful interpretations.

Can local adaptation explain varying patterns of herbivory tolerance in a recently introduced woody plant in North America?

Patterns of woody-plant mortality have been linked to global-scale environmental changes, such as extreme drought, heat stress, more frequent and intense fires, and episodic outbreaks of insects and pathogens. Although many studies have focussed on survival and mortality in response to specific physiological stresses, little attention has been paid to the role of genetic heritability of traits and local adaptation in influencing patterns of plant mortality, especially in non-native species. Tamarix spp. is a dominant, non-native riparian tree in western North America that is experiencing dieback in some areas of its range due to episodic herbivory by the recently introduced northern tamarisk leaf beetle (Diorhabda carinulata). We propose that genotype × environment interactions largely underpin current and future patterns of Tamarix mortality. We anticipate that (i) despite its recent introduction, and the potential for significant gene flow, Tamarix in western North America is generally adapted to local environmental conditions across its current range in part due to hybridization of two species; (ii) local adaptation to specific climate, soil and resource availability will yield predictable responses to episodic herbivory; and (iii) the ability to cope with a combination of episodic herbivory and increased aridity associated with climate change will be largely based on functional tradeoffs in resource allocation. This review focusses on the potential heritability of plant carbon allocation patterns in Tamarix, focussing on the relative contribution of acquired carbon to non-structural carbohydrate (NSC) pools versus other sinks as the basis for surviving episodic disturbance. Where high aridity and/or poor edaphic position lead to chronic stress, NSC pools may fall below a minimum threshold because of an imbalance between the supply of carbon and its demand by various sinks. Identifying patterns of local adaptation of traits related to resource allocation will improve forecasting of Tamarix population susceptibility to episodic herbivory.

Impact of fog drip versus fog immersion on the physiology of Bishop pine saplings.

Fog-drip to the soil is the most obvious contribution of fog to the water budget of an ecosystem, but several studies provide convincing evidence that foliar absorption of fog water through leaf wetting events is also possible. The focus of our research was to assess the relative importance of fog drip and fog immersion (foliar wetting) on leaf gas-exchange rates and photosynthetic capacity of a coastal pine species, Bishop pine (Pinus muricata D.Don), a drought-sensitive species restricted to the fog belt of coastal California and offshore islands. In a controlled experiment, we manipulated fog water inputs to potted Bishop pine saplings during a 3 week dry-down period. Ten saplings were randomly assigned one of two fog treatments: (1) fog drip to the soil and canopy fog immersion, or (2) fog immersion alone. Five saplings were assigned the 'control' group and received no fog water inputs. We found that fog immersion alone significantly increased carbon assimilation rates and photosynthetic capacity of saplings as soil moisture declined compared with those that received no fog at all. The highest carbon assimilation rates were observed in saplings that also received fog drip. Soil moisture was 40% higher in the fog immersion compared with the control group during the dry-down, indicating a reduced demand for soil water in saplings that had only leaves wetted by canopy interception of fog. Leaf-level physiology is more strongly enhanced by fog drip compared with fog immersion, although the results of this study provide evidence that foliar absorption is a viable mechanism by which Bishop pines use fog water and that it can enhance instantaneous plant carbon gain and potentially whole plant productivity.

Climate modifies response of non-native and native species richness to nutrient enrichment.

Ecosystem eutrophication often increases domination by non-natives and causes displacement of native taxa. However, variation in environmental conditions may affect the outcome of interactions between native and non-native taxa in environments where nutrient supply is elevated. We examined the interactive effects of eutrophication, climate variability and climate average conditions on the success of native and non-native plant species using experimental nutrient manipulations replicated at 32 grassland sites on four continents. We hypothesized that effects of nutrient addition would be greatest where climate was stable and benign, owing to reduced niche partitioning. We found that the abundance of non-native species increased with nutrient addition independent of climate; however, nutrient addition increased non-native species richness and decreased native species richness, with these effects dampened in warmer or wetter sites. Eutrophication also altered the time scale in which grassland invasion responded to climate, decreasing the importance of long-term climate and increasing that of annual climate. Thus, climatic conditions mediate the responses of native and non-native flora to nutrient enrichment. Our results suggest that the negative effect of nutrient addition on native abundance is decoupled from its effect on richness, and reduces the time scale of the links between climate and compositional change.

Coastal fog during summer drought improves the water status of sapling trees more than adult trees in a California pine forest.

Fog water inputs can offset seasonal drought in the Mediterranean climate of coastal California and may be critical to the persistence of many endemic plant species. The ability to predict plant species response to potential changes in the fog regime hinges on understanding the ways that fog can impact plant physiological function across life stages. Our study uses a direct metric of water status, namely plant water potential, to understand differential responses of adult versus sapling trees to seasonal drought and fog water inputs. We place these measurements within a water balance framework that incorporates the varying climatic and soil property impacts on water budgets and deficit. We conducted our study at a coastal and an inland site within the largest stand of the regionally endemic bishop pine (Pinus muricata D. Don) on Santa Cruz Island. Our results show conclusively that summer drought negatively affects the water status of sapling more than adult trees and that sapling trees are also more responsive to changes in shallow soil moisture inputs from fog water deposition. Moreover, between the beginning and end of a large, late-season fog drip event, water status increased more for saplings than for adults. Relative to non-foggy conditions, we found that fog water reduces modeled peak water deficit by 80 and 70 % at the inland and coastal sites, respectively. Results from our study inform mechanistically based predictions of how population dynamics of this and other coastal species may be affected by a warmer, drier, and potentially less foggy future.

The ghosts of trees past: savanna trees create enduring legacies in plant species composition.

Isolated trees in savannas worldwide are known to modify their local environment and interact directly with neighboring plants. Less is known about how related tree species differ in their impacts on surrounding communities, how the effects of trees vary between years, and how composition might change following loss of the tree. To address these knowledge gaps, we explored the following questions: How do savanna trees influence the surrounding composition of herbaceous plants? Is the influence of trees consistent across different species and years? How does this change following the death of the tree? We surveyed herbaceous species composition and environmental attributes surrounding living and dead evergreen and deciduous Quercus trees in California (USA) savannas across several years that differed in their total precipitation. Oak trees of all species created distinct, homogenous understory communities dominated by exotic grasses across several sites. The composition of the low-diversity understory communities showed less interannual variation than open grassland, despite a two-fold difference in precipitation between the driest and wettest year. Vegetation composition was correlated with variation in soil properties, which were strongly affected by trees. Oaks also influenced the communities beyond the edge of the crown, but this depended on site and oak species. Low-diversity understory communities persisted up to 43 years following the death of the tree. A gradual decline in the effect of trees on the physical, environment following death did not result in vegetation becoming more similar to open grassland over time. The presence of long-lasting legacies of past tree crowns highlights the difficulty of assigning control of the current distribution of herbaceous species in grassland to their contemporary environment.

Evaluating nurse plants for restoring native woody species to degraded subtropical woodlands.

Harsh habitats dominated by invasive species are difficult to restore. Invasive grasses in arid environments slow succession toward more desired composition, yet grass removal exacerbates high light and temperature, making the use of "nurse plants" an appealing strategy. In this study of degraded subtropical woodlands dominated by alien grasses in Hawai'i, we evaluated whether individuals of two native (Dodonaea viscosa, Leptocophylla tameiameia) and one non-native (Morella faya) woody species (1) act as natural nodes of recruitment for native woody species and (2) can be used to enhance survivorship of outplanted native woody species. To address these questions, we quantified the presence and persistence of seedlings naturally recruiting beneath adult nurse shrubs and compared survival and growth of experimentally outplanted seedlings of seven native woody species under the nurse species compared to intact and cleared alien-grass plots. We found that the two native nurse shrubs recruit their own offspring, but do not act as establishment nodes for other species. Morella faya recruited even fewer seedlings than native shrubs. Thus, outplanting will be necessary to increase abundance and diversity of native woody species. Outplant survival was the highest under shrubs compared to away from them with few differences between nurse species. The worst habitat for native seedling survival and growth was within the unmanaged invasive grass matrix. Although the two native nurse species did not differentially affect outplant survival, D. viscosa is the most widespread and easily propagated and is thus more likely to be useful as an initial nurse species. The outplanted species showed variable responses to nurse habitats that we attribute to resource requirements resulting from their typical successional stage and nitrogen fixation capability.

Remote sensing analysis of vegetation recovery following short-interval fires in Southern California shrublands.

Increased fire frequency has been shown to promote alien plant invasions in the western United States, resulting in persistent vegetation type change. Short interval fires are widely considered to be detrimental to reestablishment of shrub species in southern California chaparral, facilitating the invasion of exotic annuals and producing "type conversion". However, supporting evidence for type conversion has largely been at local, site scales and over short post-fire time scales. Type conversion has not been shown to be persistent or widespread in chaparral, and past range improvement studies present evidence that chaparral type conversion may be difficult and a relatively rare phenomenon across the landscape. With the aid of remote sensing data covering coastal southern California and a historical wildfire dataset, the effects of short interval fires (<8 years) on chaparral recovery were evaluated by comparing areas that burned twice to adjacent areas burned only once. Twelve pairs of once- and twice-burned areas were compared using normalized burn ratio (NBR) distributions. Correlations between measures of recovery and explanatory factors (fire history, climate and elevation) were analyzed by linear regression. Reduced vegetation cover was found in some lower elevation areas that were burned twice in short interval fires, where non-sprouting species are more common. However, extensive type conversion of chaparral to grassland was not evident in this study. Most variables, with the exception of elevation, were moderately or poorly correlated with differences in vegetation recovery.

Integrated assessment of biological invasions.

As the main witnesses of the ecological and economic impacts of invasions on ecosystems around the world, ecologists seek to provide the relevant science that informs managers about the potential for invasion of specific organisms in their region(s) of interest. Yet, the assorted literature that could inform such forecasts is rarely integrated to do so, and further, the diverse nature of the data available complicates synthesis and quantitative prediction. Here we present a set of analytical tools for synthesizing different levels of distributional and/or demographic data to produce meaningful assessments of invasion potential that can guide management at multiple phases of ongoing invasions, from dispersal to colonization to proliferation. We illustrate the utility of data-synthesis and data-model assimilation approaches with case studies of three well-known invasive species--a vine, a marine mussel, and a freshwater crayfish--under current and projected future climatic conditions. Results from the integrated assessments reflect the complexity of the invasion process and show that the most relevant climatic variables can have contrasting effects or operate at different intensities across habitat types. As a consequence, for two of the study species climate trends will increase the likelihood of invasion in some habitats and decrease it in others. Our results identified and quantified both bottlenecks and windows of opportunity for invasion, mainly related to the role of human uses of the landscape or to disruption of the flow of resources. The approach we describe has a high potential to enhance model realism, explanatory insight, and predictive capability, generating information that can inform management decisions and optimize phase-specific prevention and control efforts for a wide range of biological invasions.

Self-reinforcing impacts of plant invasions change over time.

Returning native species to habitats degraded by biological invasions is a critical conservation goal. A leading hypothesis poses that exotic plant dominance is self-reinforced by impacts on ecosystem processes, leading to persistent stable states. Invaders have been documented to modify fire regimes, alter soil nutrients or shift microbial communities in ways that feed back to benefit themselves over competitors. However, few studies have followed invasions through time to ask whether ecosystem impacts and feedbacks persist. Here we return to woodland sites in Hawai'i Volcanoes National Park that were invaded by exotic C4 grasses in the 1960s, the ecosystem impacts of which were studied intensively in the 1990s. We show that positive feedbacks between exotic grasses and soil nitrogen cycling have broken down, but rather than facilitating native vegetation, the weakening feedbacks facilitate new exotic species. Data from the 1990s showed that exotic grasses increased nitrogen-mineralization rates by two- to fourfold, but were nitrogen-limited. Thus, the impacts of the invader created a positive feedback early in the invasion. We now show that annual net soil nitrogen mineralization has since dropped to pre-invasion levels. In addition, a seedling outplanting experiment that varied soil nitrogen and grass competition demonstrates that the changing impacts of grasses do not favour native species re-establishment. Instead, decreased nitrogen availability most benefits another aggressive invader, the nitrogen-fixing tree Morella faya. Long-term studies of invasions may reveal that ecosystem impacts and feedbacks shift over time, but that this may not benefit native species recovery.

Predicting invasion in grassland ecosystems: is exotic dominance the real embarrassment of richness?

Invasions have increased the size of regional species pools, but are typically assumed to reduce native diversity. However, global-scale tests of this assumption have been elusive because of the focus on exotic species richness, rather than relative abundance. This is problematic because low invader richness can indicate invasion resistance by the native community or, alternatively, dominance by a single exotic species. Here, we used a globally replicated study to quantify relationships between exotic richness and abundance in grass-dominated ecosystems in 13 countries on six continents, ranging from salt marshes to alpine tundra. We tested effects of human land use, native community diversity, herbivore pressure, and nutrient limitation on exotic plant dominance. Despite its widespread use, exotic richness was a poor proxy for exotic dominance at low exotic richness, because sites that contained few exotic species ranged from relatively pristine (low exotic richness and cover) to almost completely exotic-dominated ones (low exotic richness but high exotic cover). Both exotic cover and richness were predicted by native plant diversity (native grass richness) and land use (distance to cultivation). Although climate was important for predicting both exotic cover and richness, climatic factors predicting cover (precipitation variability) differed from those predicting richness (maximum temperature and mean temperature in the wettest quarter). Herbivory and nutrient limitation did not predict exotic richness or cover. Exotic dominance was greatest in areas with low native grass richness at the site- or regional-scale. Although this could reflect native grass displacement, a lack of biotic resistance is a more likely explanation, given that grasses comprise the most aggressive invaders. These findings underscore the need to move beyond richness as a surrogate for the extent of invasion, because this metric confounds monodominance with invasion resistance. Monitoring species' relative abundance will more rapidly advance our understanding of invasions.

Non-Additive effects on decomposition from mixing litter of the invasive Mikania micrantha H.B.K. with native plants.

A common hypothesis to explain the effect of litter mixing is based on the difference in litter N content between mixed species. Although many studies have shown that litter of invasive non-native plants typically has higher N content than that of native plants in the communities they invade, there has been surprisingly little study of mixing effects during plant invasions. We address this question in south China where Mikania micrantha H.B.K., a non-native vine, with high litter N content, has invaded many forested ecosystems. We were specifically interested in whether this invader accelerated decomposition and how the strength of the litter mixing effect changes with the degree of invasion and over time during litter decomposition. Using litterbags, we evaluated the effect of mixing litter of M. micrantha with the litter of 7 native resident plants, at 3 ratios: M1 (1∶4, = exotic:native litter), M2 (1∶1) and M3 (4∶1, = exotic:native litter) over three incubation periods. We compared mixed litter with unmixed litter of the native species to identify if a non-additive effect of mixing litter existed. We found that there were positive significant non-additive effects of litter mixing on both mass loss and nutrient release. These effects changed with native species identity, mixture ratio and decay times. Overall the greatest accelerations of mixture decay and N release tended to be in the highest degree of invasion (mix ratio M3) and during the middle and final measured stages of decomposition. Contrary to expectations, the initial difference in litter N did not explain species differences in the effect of mixing but overall it appears that invasion by M. micrantha is accelerating the decomposition of native species litter. This effect on a fundamental ecosystem process could contribute to higher rates of nutrient turnover in invaded ecosystems.

Bounded ranges of variation as a framework for future conservation and fire management.

Alterations in natural fire patterns have negatively affected fire-prone ecosystems in many ways. The historical range of variability (HRV) concept evolved as a management target for natural vegetation composition and fire regimes in fire-prone ecosystems. HRV-based management inherently assumes that ecosystem resilience is reflected in observed ranges of past vegetation and fire dynamics, typically without knowledge of where thresholds exist beyond these dynamics. Given uncertainty in future conditions, some have argued that HRV may not adequately reflect ecosystem resilience to future fire activity. We suggest a refinement that includes concepts from the thresholds of potential concern (TPC) framework, which emphasizes tipping points at the extremes of ecosystem dynamics and other socially unacceptable outcomes. We propose bounded ranges of variation (BRV), an approach focused on building resilience by using historical information, but also by identifying socio-ecological thresholds to avoid and associated management action triggers. Here, we examine nonnative species and carbon sequestration as examples of how the BRV framework could be used in the context of conservation and fire management.

Introduced annual grass increases regional fire activity across the arid western USA (1980-2009).

Non-native, invasive grasses have been linked to altered grass-fire cycles worldwide. Although a few studies have quantified resulting changes in fire activity at local scales, and many have speculated about larger scales, regional alterations to fire regimes remain poorly documented. We assessed the influence of large-scale Bromus tectorum (hereafter cheatgrass) invasion on fire size, duration, spread rate, and interannual variability in comparison to other prominent land cover classes across the Great Basin, USA. We compared regional land cover maps to burned area measured using the Moderate Resolution Imaging Spectroradiometer (MODIS) for 2000-2009 and to fire extents recorded by the USGS registry of fires from 1980 to 2009. Cheatgrass dominates at least 6% of the central Great Basin (650 000 km(2) ). MODIS records show that 13% of these cheatgrass-dominated lands burned, resulting in a fire return interval of 78 years for any given location within cheatgrass. This proportion was more than double the amount burned across all other vegetation types (range: 0.5-6% burned). During the 1990s, this difference was even more extreme, with cheatgrass burning nearly four times more frequently than any native vegetation type (16% of cheatgrass burned compared to 1-5% of native vegetation). Cheatgrass was also disproportionately represented in the largest fires, comprising 24% of the land area of the 50 largest fires recorded by MODIS during the 2000s. Furthermore, multi-date fires that burned across multiple vegetation types were significantly more likely to have started in cheatgrass. Finally, cheatgrass fires showed a strong interannual response to wet years, a trend only weakly observed in native vegetation types. These results demonstrate that cheatgrass invasion has substantially altered the regional fire regime. Although this result has been suspected by managers for decades, this study is the first to document recent cheatgrass-driven fire regimes at a regional scale.