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Background: Seedling germination and survival is a critical control on forest ecosystem boundaries, such as at the alpine-treeline ecotone. In addition, while it is known that species respond individualistically to the same suite of environmental drivers, the potential additional effect of local adaptation on seedling success has not been evaluated. Aims: To determine whether local adaptation may influence the position and movement of forest ecosystem boundaries, we quantified conifer seedling recruitment in common gardens across a subalpine forest to alpine tundra gradient at Niwot Ridge, Colorado, USA. Methods: We studied Pinus flexilis and Picea engelmannii grown from seed collected locally at High (3400 m a.s.l.) and Low (3060 m a.s.l.) elevations. We monitored emergence and survival of seeds sown directly into plots and survival of seedlings germinated indoors and transplanted after snowmelt. Results: Emergence and survival through the first growing season was greater for P. flexilis than P. engelmannii and for Low compared with High provenances. Yet survival through the second growing season was similar for both species and provenances. Seedling emergence and survival tended to be greatest in the subalpine forest and lowest in the alpine tundra. Survival was greater for transplants than for field-germinated seedlings. Conclusions: These results suggest that survival through the first few weeks is critical to the establishment of natural germinants. In addition, even small distances between seed sources can have a significant effect on early demographic performance - a factor that has rarely been considered in previous studies of tree recruitment and species range shifts. © 2013 The work of C. Castanha, M.S. Torn, and L.M. Kueppers was conducted at Lawrence Berkeley National Laboratory under Contract No. DE-AC02-05CH11231 with the U.S. Department of Energy. The U.S. Government retains, and the publisher, by accepting the article for publication, acknowledges, that the U.S. Government retains a non-exclusive, paid-up, irrevocable, worldwide license to publish or reproduce the published form of this manuscript, or allow others to do so, for U.S. Government purposes. The work of M.J. Germino at the U.S. Geological Survey was authored as part of the Contributor's official duties as an Employee of the United States Government and is therefore a work of the United States Government. In accordance with 17 U.S.C. 105, no copyright protection is available for such works under U.S. Law. B. Weibel waives her copyright in the work but not her status as co-author of the work.

Local adaptation may facilitate range expansion during invasions, but the mechanisms promoting destructive invasions remain unclear. Cheatgrass ( Bromus tectorum ), native to Eurasia and Africa, has invaded globally, with particularly severe impacts in western North America. We sequenced 307 genotypes and conducted controlled experiments. We found that diverse lineages invaded North America, where long-distance gene flow is common. Ancestry and phenotypic clines in the native range predicted those in the invaded range, indicating pre-adapted genotypes colonized different regions. Common gardens showed directional selection on flowering time that reversed between warm and cold sites, potentially maintaining clines. In the Great Basin, genomic predictions of strong local adaptation identified sites where cheatgrass is most dominant. Preventing new introductions that may fuel adaptation is critical for managing ongoing invasions., Competing Interests: Competing interests: Authors declare that they have no competing interests.

Fuel-treatments targeting shrubs and fire-prone exotic annual grasses (EAGs) are increasingly used to mitigate increased wildfire risks in arid and semiarid environments, and understanding their response to natural factors is needed for effective landscape management. Using field-data collected over four years from fuel-break treatments in semiarid sagebrush-steppe, we asked 1) how the outcomes of EAG and sagebrush fuel treatments varied with site biophysical properties, climate, and weather, and 2) how predictions of fire behavior using the Fuel Characteristic Classification System fire model related to land-management objectives of maintaining fire behavior expected of low-load, dry-climate grasslands. Generalized linear mixed effect modeling with build-up model selection was used to determine best-fit models, and marginal effects plots to assess responses for each fuel type. EAG cover decreased as antecedent-fall precipitation increased and increased as antecedent-spring temperatures and surface soil clay contents increased. Herbicides targeting EAGs were less effective where pre-treatment EAG cover was >40 % and antecedent spring temperatures were >9.5 °C. Sagebrush cover was inversely related to soil clay content, especially where clay contents were >17 %. Predicted fire behavior exceeded management objectives under 1) average fire weather conditions when EAG or sagebrush cover was >50 % or >26 %, respectively, or 2) extreme fire weather conditions when EAG or sagebrush cover was >10 % or >8 %, respectively. Consideration of the strong effects of natural variability in site properties and antecedent weather can help in justifying, planning and implementing fuel-treatments., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Published by Elsevier Ltd.)

Field-based transplant gardens, including common and reciprocal garden experiments, are a powerful tool for studying genetic variation and gene-by-environment interactions. These experiments assume that individuals within the garden represent independent replicates growing in a homogenous environment. Plant neighborhood interactions are pervasive across plant populations and could violate assumptions of transplant garden experiments. We demonstrate how spatially explicit models for plant-plant interactions can provide novel insights on genotypes' performance in field-transplant garden designs. We used individual-based models, based on data from a sagebrush ( Artemisia spp.) common garden, to simulate the impact of spatial plant-plant interactions on between-group differences in plant growth. We found that planting densities within the range of those used in many common gardens can bias experimental outcomes. Our results demonstrate that higher planting densities can lead to inflated group differences and may confound genotypes' competitive ability and genetically underpinned variation. Synthesis. We propose that spatially explicit models can help avoid biased results by informing the design and analysis of field-based transplant garden experiments. Alternately, including neighborhood effects in post hoc analyses of transplant garden experiments is likely to provide novel insights into the roles of biotic factors and density dependence in genetic differentiation., Competing Interests: The authors declare no conflicts of interest., (© 2022 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Plant-population recovery across large disturbance areas is often seed-limited. An understanding of seed dispersal patterns is fundamental for determining natural-regeneration potential. However, forecasting seed dispersal rates across heterogeneous landscapes remains a challenge. Our objectives were to determine (i) the landscape patterning of post-disturbance seed dispersal, and underlying sources of variation and the scale at which they operate, and (ii) how the natural seed dispersal patterns relate to a seed augmentation strategy. Vertical seed trapping experiments were replicated across 2 years and five burned and/or managed landscapes in sagebrush steppe. Multi-scale sampling and hierarchical Bayesian models were used to determine the scale of spatial variation in seed dispersal. We then integrated an empirical and mechanistic dispersal kernel for wind-dispersed species to project rates of seed dispersal and compared natural seed arrival to typical post-fire aerial seeding rates. Seeds were captured across the range of tested dispersal distances, up to a maximum distance of 26 m from seed-source plants, although dispersal to the furthest traps was variable. Seed dispersal was better explained by transect heterogeneity than by patch or site heterogeneity (transects were nested within patch within site). The number of seeds captured varied from a modelled mean of ~13 m -2 adjacent to patches of seed-producing plants, to nearly none at 10 m from patches, standardized over a 49-day period. Maximum seed dispersal distances on average were estimated to be 16 m according to a novel modelling approach using a 'latent' variable for dispersal distance based on seed trapping heights. Surprisingly, statistical representation of wind did not improve model fit and seed rain was not related to the large variation in total available seed of adjacent patches. The models predicted severe seed limitations were likely on typical burned areas, especially compared to the mean 95-250 seeds per m 2 that previous literature suggested were required to generate sagebrush recovery. More broadly, our Bayesian data fusion approach could be applied to other cases that require quantitative estimates of long-distance seed dispersal across heterogeneous landscapes., (© The Author(s) 2022. Published by Oxford University Press on behalf of the Annals of Botany Company. Elements of this work were written written by employees of the US Government.)

Accurate predictions of ecological restoration outcomes are needed across the increasingly large landscapes requiring treatment following disturbances. However, observational studies often fail to account for nonrandom treatment application, which can result in invalid inference. Examining a spatiotemporally extensive management treatment involving post-fire seeding of declining sagebrush shrubs across semiarid areas of the western USA over two decades, we quantify drivers and consequences of selection biases in restoration using remotely sensed data. From following more than 1,500 wildfires, we find treatments were disproportionately applied in more stressful, degraded ecological conditions. Failure to incorporate unmeasured drivers of treatment allocation led to the conclusion that costly, widespread seedings were unsuccessful; however, after considering sources of bias, restoration positively affected sagebrush recovery. Treatment effects varied with climate, indicating prioritization criteria for interventions. Our findings revise the perspective that post-fire sagebrush seedings have been broadly unsuccessful and demonstrate how selection biases can pose substantive inferential hazards in observational studies of restoration efficacy and the development of restoration theory., (© 2022. The Author(s).)

Habitat loss is the most prevalent threat to biodiversity in North America. One of the most threatened landscapes in the United States is the sagebrush ( Artemisia spp.) ecosystem, much of which has been fragmented or converted to non-native grasslands via the cheatgrass-fire cycle. Like many sagebrush obligates, greater sage-grouse ( Centrocercus urophasianus ) depend upon sagebrush for food and cover and are affected by changes to this ecosystem. We investigated habitat selection by 28 male greater sage-grouse during each of 3 years after a 113,000-ha wildfire in a sagebrush steppe ecosystem in Idaho and Oregon. During the study period, seeding and herbicide treatments were applied for habitat restoration. We evaluated sage-grouse responses to vegetation and post-fire restoration treatments. Throughout the 3 years post-fire, sage-grouse avoided areas with high exotic annual grass cover but selected strongly for recovering sagebrush and moderately strongly for perennial grasses. By the third year post-fire, they preferred high-density sagebrush, especially in winter when sagebrush is the primary component of the sage-grouse diet. Sage-grouse preferred forb habitat immediately post-fire, especially in summer, but this selection preference was less strong in later years. They also selected areas that were intensively treated with herbicide and seeded with sagebrush, grasses, and forbs, although these responses varied with time since treatment. Wildfire can have severe consequences for sagebrush-obligate species due to loss of large sagebrush plants used for food and for protection from predators and thermal extremes. Our results show that management efforts, including herbicide application and seeding of plants, directed at controlling exotic annual grasses after a wildfire can positively affect habitat selection by sage-grouse., Competing Interests: The authors declare no conflict of interest., (© 2022 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd. This article has been contributed to by U.S. Government employees and their work is in the public domain in the USA.)

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., (© 2021 by the Ecological Society of America.)

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., (© 2021 Botanical Society of America. This article has been contributed to by US Government employees and their work is in the public domain in the USA.)

Adaptive variation among plant populations must be known for effective conservation and restoration of imperiled species and predicting their responses to a changing climate. Common-garden experiments, in which plants sourced from geographically distant populations are grown together such that genetic differences may be expressed, have provided much insight on adaptive variation. Common-garden experiments also form the foundation for climate-based seed-transfer guidelines. However, the spatial scale at which population differentiation occurs is rarely addressed, leaving a critical information gap for parameterizing seed-transfer guidelines and assessing species' climate vulnerability. We asked whether adaptation was evident among populations of a foundational perennial within a single "empirical" seed-transfer zone (based on previous common-garden findings evaluating very distant populations) but different "provisional" seed zones (groupings of areas of similar climate and are not parameterized from common-garden data). Seedlings from three populations originating from similar conditions within an intermediate elevation were planted into gardens nearby at the same elevation, or 250-450 m higher or lower in elevation and 0.4-25 km away. Substantial variation was observed between gardens in survival (ranging 2%-99%), foliar crown volume (7.8-22.6 dm 3 ), and reproductive effort (0%-65%), but not among the three transplanted populations. The between garden variation was inversely related to climatic differences between the gardens and seed-source populations, specifically the site differences in maximum-minimum annual temperatures. Results suggest that substantial site-specificity in adaptation can occur at finer scales than is accounted for in empirical seed-transfer guidance when the guidance is derived from broadscale common-garden studies. Being within the same empirical seed zone, geographic unit, and even within 10 km distance may not qualify as "local" in the context of seed transfer. Moving forward, designing common-garden experiments so that they allow for testing the scale of adaptation will help in translating the resulting seed-transfer guidance to restoration projects., Competing Interests: The authors state that there is no conflict of interest., (Published 2020. This article is a U.S. Government work and is in the public domain in the USA. Ecology and Evolution published by John Wiley & Sons Ltd.)

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., (© Published 2019. This article is a U.S. Government work and is in the public domain in the USA.)

Population-level adaptation to spatial variation in factors such as climate and soils is critical for climate-vulnerability assessments, restoration seeding, and other ecological applications in species management, and the underlying information is typically based on common-garden studies that are short duration. Here, we show >20 yr were required for adaptive differences to emerge among 13 populations of a widespread shrub (sagebrush, Artemisia tridentata ssp wyomingensis) collected from around the western United States and planted into common gardens. Additionally, >10 yr were required for greater survival of local populations, that is, local adaptation, to become evident. Variation in survival was best explained by the combination of populations' home ecoregion combined with grouping of minimum temperature and aridity. Additional reductions in survival were explained by ungrouped (i.e., continuous) measures of garden-to-population-origin separation in geographic distance (5% decrease in survival per 100 km increase in separation; R 2  = 0.22) and especially in minimum temperature in younger plants (-4% per + °C difference, R 2  = 0.56 vs. 0.29 in the 14th vs. 27th post-planting years, respectively). Longer-term common garden studies are needed. While we await them, uncertainty in adaptive variation resulting from short-term observations could be quantitatively estimated and reported with seed-transfer guidelines to reduce risks of introducing maladapted provenances in restoration., (© 2018 by the Ecological Society of America.)

Restoration and rehabilitation of native vegetation in dryland ecosystems, which encompass over 40% of terrestrial ecosystems, is a common challenge that continues to grow as wildfire and biological invasions transform dryland plant communities. The difficulty in part stems from low and variable precipitation, combined with limited understanding about how weather conditions influence restoration outcomes, and increasing recognition that one-time seeding approaches can fail if they do not occur during appropriate plant establishment conditions. The sagebrush biome, which once covered over 620,000 km 2 of western North America, is a prime example of a pressing dryland restoration challenge for which restoration success has been variable. We analyzed field data on Artemisia tridentata (big sagebrush) restoration collected at 771 plots in 177 wildfire sites across its western range, and used process-based ecohydrological modeling to identify factors leading to its establishment. Our results indicate big sagebrush occurrence is most strongly associated with relatively cool temperatures and wet soils in the first spring after seeding. In particular, the amount of winter snowpack, but not total precipitation, helped explain the availability of spring soil moisture and restoration success. We also find considerable interannual variability in the probability of sagebrush establishment. Adaptive management strategies that target seeding during cool, wet years or mitigate effects of variability through repeated seeding may improve the likelihood of successful restoration in dryland ecosystems. Given consistent projections of increasing temperatures, declining snowpack, and increasing weather variability throughout midlatitude drylands, weather-centric adaptive management approaches to restoration will be increasingly important for dryland restoration success., (Published 2018. This article is a U.S. Government work and is in the public domain in the USA.)

The spatial patterning of alpine plant communities is strongly influenced by the variation in physical factors such as temperature and moisture, which are strongly affected by snow depth and snowmelt patterns. Earlier snowmelt timing and greater soil-moisture limitations may favor wide-ranging species adapted to a broader set of ecohydrological conditions than alpine-restricted species. We asked how plant community composition, phenology, plant water relations, and photosynthetic gas exchange of alpine-restricted and wide-ranging species differ in their responses to a ca. 40-day snowmelt gradient in the Colorado Rocky Mountains ( Lewisia pygmaea , Sibbaldia procumbens , and Hymenoxys grandiflora were alpine-restricted and Artemisia scopulorum , Carex rupestris , and Geum rossii were wide-ranging species). As hypothesized, species richness and foliar cover increased with earlier snowmelt, due to a greater abundance of wide-ranging species present in earlier melting plots. Flowering initiation occurred earlier with earlier snowmelt for 12 out of 19 species analyzed, while flowering duration was shortened with later snowmelt for six species (all but one were wide-ranging species). We observed >50% declines in net photosynthesis from July to September as soil moisture and plant water potentials declined. Early-season stomatal conductance was higher in wide-ranging species, indicating a more competitive strategy for water acquisition when soil moisture is high. Even so, there were no associated differences in photosynthesis or transpiration, suggesting no strong differences between these groups in physiology. Our findings reveal that plant species with different ranges (alpine-restricted vs. wide-ranging) could have differential phenological and physiological responses to snowmelt timing and associated soil moisture dry-down, and that alpine-restricted species' performance is more sensitive to snowmelt. As a result, alpine-restricted species may serve as better indicator species than their wide-ranging heterospecifics. Overall, alpine community composition and peak % cover are strongly structured by spatio-temporal patterns in snowmelt timing. Thus, near-term, community-wide changes (or variation) in phenology and physiology in response to shifts in snowmelt timing or rates of soil dry down are likely to be contingent on the legacy of past climate on community structure.

Understanding how climate warming will affect the demographic rates of different ecotypes is critical to predicting shifts in species distributions. Here, we present results from a common garden, climate change experiment in which we measured seedling recruitment of lodgepole pine, a widespread North American conifer that is also planted globally. Seeds from a low-elevation provenance had more than three-fold greater recruitment to their third year than seeds from a high-elevation provenance across sites within and above its native elevation range and across climate manipulations. Heating halved recruitment to the third year of both low- and high-elevation seed sources across the elevation gradient, while watering more than doubled recruitment, alleviating some of the negative effects of heating. Demographic models based on recruitment data from the climate manipulations and long-term observations of adult populations revealed that heating could effectively halt modeled upslope range expansion except when combined with watering. Simulating fire and rapid postfire forest recovery at lower elevations accelerated lodgepole pine expansion into the alpine, but did not alter final abundance rankings among climate scenarios. Regardless of climate scenario, greater recruitment of low-elevation seeds compensated for longer dispersal distances to treeline, assuming colonization was allowed to proceed over multiple centuries. Our results show that ecotypes from lower elevations within a species' range could enhance recruitment and facilitate upslope range shifts with climate change., (© 2017 John Wiley & Sons Ltd.)

Widespread tree mortality associated with drought has been observed on all forested continents and global change is expected to exacerbate vegetation vulnerability. Forest mortality has implications for future biosphere-atmosphere interactions of carbon, water and energy balance, and is poorly represented in dynamic vegetation models. Reducing uncertainty requires improved mortality projections founded on robust physiological processes. However, the proposed mechanisms of drought-induced mortality, including hydraulic failure and carbon starvation, are unresolved. A growing number of empirical studies have investigated these mechanisms, but data have not been consistently analysed across species and biomes using a standardized physiological framework. Here, we show that xylem hydraulic failure was ubiquitous across multiple tree taxa at drought-induced mortality. All species assessed had 60% or higher loss of xylem hydraulic conductivity, consistent with proposed theoretical and modelled survival thresholds. We found diverse responses in non-structural carbohydrate reserves at mortality, indicating that evidence supporting carbon starvation was not universal. Reduced non-structural carbohydrates were more common for gymnosperms than angiosperms, associated with xylem hydraulic vulnerability, and may have a role in reducing hydraulic function. Our finding that hydraulic failure at drought-induced mortality was persistent across species indicates that substantial improvement in vegetation modelling can be achieved using thresholds in hydraulic function.

Climate niche models project that subalpine forest ranges will extend upslope with climate warming. These projections assume that the climate suitable for adult trees will be adequate for forest regeneration, ignoring climate requirements for seedling recruitment, a potential demographic bottleneck. Moreover, local genetic adaptation is expected to facilitate range expansion, with tree populations at the upper forest edge providing the seed best adapted to the alpine. Here, we test these expectations using a novel combination of common gardens, seeded with two widely distributed subalpine conifers, and climate manipulations replicated at three elevations. Infrared heaters raised temperatures in heated plots, but raised temperatures more in the forest than at or above treeline because strong winds at high elevation reduced heating efficiency. Watering increased season-average soil moisture similarly across sites. Contrary to expectations, warming reduced Engelmann spruce recruitment at and above treeline, as well as in the forest. Warming reduced limber pine first-year recruitment in the forest, but had no net effect on fourth-year recruitment at any site. Watering during the snow-free season alleviated some negative effects of warming, indicating that warming exacerbated water limitations. Contrary to expectations of local adaptation, low-elevation seeds of both species initially recruited more strongly than high-elevation seeds across the elevation gradient, although the low-provenance advantage diminished by the fourth year for Engelmann spruce, likely due to small sample sizes. High- and low-elevation provenances responded similarly to warming across sites for Engelmann spruce, but differently for limber pine. In the context of increasing tree mortality, lower recruitment at all elevations with warming, combined with lower quality, high-provenance seed being most available for colonizing the alpine, portends range contraction for Engelmann spruce. The lower sensitivity of limber pine to warming indicates a potential for this species to become more important in subalpine forest communities in the coming centuries., (© 2016 John Wiley & Sons Ltd.)

A genecological approach was used to explore genetic variation for survival in Artemisia tridentata (big sagebrush). Artemisia tridentata is a widespread and foundational shrub species in western North America. This species has become extremely fragmented, to the detriment of dependent wildlife, and efforts to restore it are now a land management priority. Common-garden experiments were established at three sites with seedlings from 55 source-populations. Populations included each of the three predominant subspecies, and cytotype variations. Survival was monitored for 5 years to assess differences in survival between gardens and populations. We found evidence of adaptive genetic variation for survival. Survival within gardens differed by source-population and a substantial proportion of this variation was explained by seed climate of origin. Plants from areas with the coldest winters had the highest levels of survival, while populations from warmer and drier sites had the lowest levels of survival. Survival was lowest, 36%, in the garden that was prone to the lowest minimum temperatures. These results suggest the importance of climatic driven genetic differences and their effect on survival. Understanding how genetic variation is arrayed across the landscape, and its association with climate can greatly enhance the success of restoration and conservation.

Semi-arid rangelands are important carbon (C) pools at global scales. However, the degree of net C storage or release in water-limited systems is a function of precipitation amount and timing, as well as plant community composition. In northern latitudes of western North America, C storage in cold-desert ecosystems could increase with boosts in wintertime precipitation, in which climate models predict, due to increases in wintertime soil water storage that enhance summertime productivity. However, there are few long-term, manipulative field-based studies investigating how rangelands will respond to altered precipitation amount or timing. We measured aboveground C pools and fluxes at leaf, soil, and ecosystem scales over a single growing season in plots that had 200 mm of supplemental precipitation added in either winter or summer for the past 21 years, in shrub- and exotic-bunchgrass-dominated garden plots. At our cold-desert site (298 mm precipitation during the study year), we hypothesized that increased winter precipitation would stimulate the aboveground C uptake and storage relative to ambient conditions, especially in plots containing shrubs. Our hypotheses were generally supported: ecosystem C uptake and long-term biomass accumulation were greater in winter- and summer-irrigated plots compared to control plots in both vegetation communities. However, substantial increases in the aboveground biomass occurred only in winter-irrigated plots that contained shrubs. Our findings suggest that increases in winter precipitation will enhance C storage of this widespread ecosystem, and moreso in shrub- compared to grass-dominated communities.

Application of stable isotopes of water to studies of plant-soil interactions often requires a substantial preparatory step of extracting water from samples without fractionating isotopes. Online heating is an emerging approach for this need, but is relatively untested and major questions of how to best deliver standards and assess interference by organics have not been evaluated. We examined these issues in our application of measuring woody stem xylem of sagebrush using a Picarro laser spectrometer with online induction heating. We determined (1) effects of cryogenic compared to induction-heating extraction, (2) effects of delivery of standards on filter media compared to on woody stem sections, and (3) spectral interference from organic compounds for these approaches (and developed a technique to do so). Our results suggest that matching sample and standard media improves accuracy, but that isotopic values differ with the extraction method in ways that are not due to spectral interference from organics.

Climate change is altering plant species distributions globally, and warming is expected to promote uphill shifts in mountain trees. However, at many cold-edge range limits, such as alpine treelines in the western United States, tree establishment may be colimited by low temperature and low moisture, making recruitment patterns with warming difficult to predict. We measured response functions linking carbon (C) assimilation and temperature- and moisture-related microclimatic factors for limber pine (Pinus flexilis) seedlings growing in a heating × watering experiment within and above the alpine treeline. We then extrapolated these response functions using observed microclimate conditions to estimate the net effects of warming and associated soil drying on C assimilation across an entire growing season. Moisture and temperature limitations were each estimated to reduce potential growing season C gain from a theoretical upper limit by 15-30% (c. 50% combined). Warming above current treeline conditions provided relatively little benefit to modeled net assimilation, whereas assimilation was sensitive to either wetter or drier conditions. Summer precipitation may be at least as important as temperature in constraining C gain by establishing subalpine trees at and above current alpine treelines as seasonally dry subalpine and alpine ecosystems continue to warm., (© 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.)

Survival of tree seedlings at high elevations has been shown to be limited by thermal constraints on carbon balance, but it is unknown if carbon relations also limit seedling survival at lower elevations, where water relations may be more important. We measured and modeled carbon fluxes and water relations in first-year Pinus flexilis seedlings in garden plots just beyond the warm edge of their natural range, and compared these with dry-mass gain and survival across two summers. We hypothesized that mortality in these seedlings would be associated with declines in water relations, more so than with carbon-balance limitations. Rather than gradual declines in survivorship across growing seasons, we observed sharp, large-scale mortality episodes that occurred once volumetric soil-moisture content dropped below 10%. By this point, seedling water potentials had decreased below -5 MPa, seedling hydraulic conductivity had decreased by 90% and seedling hydraulic resistance had increased by >900%. Additionally, non-structural carbohydrates accumulated in aboveground tissues at the end of both summers, suggesting impairments in phloem-transport from needles to roots. This resulted in low carbohydrate concentrations in roots, which likely impaired root growth and water uptake at the time of critically low soil moisture. While photosynthesis and respiration on a leaf area basis remained high until critical hydraulic thresholds were exceeded, modeled seedling gross primary productivity declined steadily throughout the summers. At the time of mortality, modeled productivity was insufficient to support seedling biomass-gain rates, metabolism and secondary costs. Thus the large-scale mortality events that we observed near the end of each summer were most directly linked with acute, episodic declines in plant hydraulic function that were linked with important changes in whole-seedling carbon relations., (© The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Integration of conservation partnerships across geographic, biological, and administrative boundaries is increasingly relevant because drivers of change, such as climate shifts, transcend these boundaries. We explored successes and challenges of established conservation programs that span multiple watersheds and consider both social and ecological concerns. We asked representatives from a diverse set of 11 broad-extent conservation partnerships in 29 countries 17 questions that pertained to launching and maintaining partnerships for broad-extent conservation, specifying ultimate management objectives, and implementation and learning. Partnerships invested more funds in implementing conservation actions than any other aspect of conservation, and a program's context (geographic extent, United States vs. other countries, developed vs. developing nation) appeared to substantially affect program approach. Despite early successes of these organizations and benefits of broad-extent conservation, specific challenges related to uncertainties in scaling up information and to coordination in the face of diverse partner governance structures, conflicting objectives, and vast uncertainties regarding future system dynamics hindered long-term success, as demonstrated by the focal organizations. Engaging stakeholders, developing conservation measures, and implementing adaptive management were dominant challenges. To inform future research on broad-extent conservation, we considered several challenges when we developed detailed questions, such as what qualities of broad-extent partnerships ensure they complement, integrate, and strengthen, rather than replace, local conservation efforts and which adaptive management processes yield actionable conservation strategies that account explicitly for dynamics and uncertainties regarding multiscale governance, environmental conditions, and knowledge of the system?, (© 2014 Society for Conservation Biology.)

Interacting threats to ecosystem function, including climate change, wildfire, and invasive species necessitate native plant restoration in desert ecosystems. However, native plant restoration efforts often remain unguided by ecological genetic information. Given that many ecosystems are in flux from climate change, restoration plans need to account for both contemporary and future climates when choosing seed sources. In this study we analyze vegetative responses, including mortality, growth, and carbon isotope ratios in two blackbrush (Coleogyne ramosissima) common gardens that included 26 populations from a range-wide collection. This shrub occupies ecotones between the warm and cold deserts of Mojave and Colorado Plateau ecoregions in western North America. The variation observed in the vegetative responses of blackbrush populations was principally explained by grouping populations by ecoregions and by regression with site-specific climate variables. Aridity weighted by winter minimum temperatures best explained vegetative responses; Colorado Plateau sites were usually colder and drier than Mojave sites. The relationship between climate and vegetative response was mapped within the boundaries of the species-climate space projected for the contemporary climate and for the decade surrounding 2060. The mapped ecological genetic pattern showed that genetic variation could be classified into cool-adapted and warm-adapted ecotypes, with populations often separated by steep dines. These transitions are predicted to occur in both the Mojave Desert and Colorado Plateau ecoregions. While under contemporary conditions the warm-adapted ecotype occupies the majority of climate space, climate projections predict that the cool-adapted ecotype could prevail as the dominant ecotype as the climate space of blackbrush expands into higher elevations and latitudes. This study provides the framework for delineating climate change-responsive seed transfer guidelines, which are needed to inform restoration and management planning. We propose four transfer zones in blackbrush that correspond to areas currently dominated by cool-adapted and warm-adapted ecotypes in each of the two ecoregions.

Vegetation change is expected with global climate change, potentially altering ecosystem function and climate feedbacks. However, causes of plant mortality, which are central to vegetation change, are understudied, and physiological mechanisms remain unclear, particularly the roles of carbon metabolism and xylem function. We report analysis of foliar nonstructural carbohydrates (NSCs) and associated physiology from a previous experiment where earlier drought-induced mortality of Pinus edulis at elevated temperatures was associated with greater cumulative respiration. Here, we predicted faster NSC decline for warmed trees than for ambient-temperature trees. Foliar NSC in droughted trees declined by 30% through mortality and was lower than in watered controls. NSC decline resulted primarily from decreased sugar concentrations. Starch initially declined, and then increased above pre-drought concentrations before mortality. Although temperature did not affect NSC and sugar, starch concentrations ceased declining and increased earlier with higher temperatures. Reduced foliar NSC during lethal drought indicates a carbon metabolism role in mortality mechanism. Although carbohydrates were not completely exhausted at mortality, temperature differences in starch accumulation timing suggest that carbon metabolism changes are associated with time to death. Drought mortality appears to be related to temperature-dependent carbon dynamics concurrent with increasing hydraulic stress in P. edulis and potentially other similar species., (© 2013 The Authors. New Phytologist © 2013 New Phytologist Trust.)

Continued changes in climate are projected to alter the geographic distributions of plant species, in part by affecting where individuals can establish from seed. We tested the hypothesis that warming promotes uphill redistribution of subalpine tree populations by reducing cold limitation at high elevation and enhancing drought stress at low elevation. We seeded limber pine (Pinus flexilis) into plots with combinations of infrared heating and water addition treatments, at sites positioned in lower subalpine forest, the treeline ecotone, and alpine tundra. In 2010, first-year seedlings were assessed for physiological performance and survival over the snow-free growing season. Seedlings emerged in midsummer, about 5-8 weeks after snowmelt. Low temperature was not observed to limit seedling photosynthesis or respiration between emergence and October, and thus experimental warming did not appear to reduce cold limitation at high elevation. Instead, gas exchange and water potential from all sites indicated a prevailing effect of summer moisture stress on photosynthesis and carbon balance. Infrared heaters raised soil growing degree days (base 5 °C, p < 0.001) and August-September mean soil temperature (p < 0.001). Despite marked differences in vegetation cover and meteorological conditions across sites, volumetric soil moisture content (θ) at 5-10 cm below 0.16 and 0.08 m(3) m(-3) consistently corresponded with moderate and severe indications of drought stress in midday stem water potential, stomatal conductance, photosynthesis, and respiration. Seedling survival was greater in watered plots than in heated plots (p = 0.01), and negatively related to soil growing degree days and duration of exposure to θ < 0.08 m(3) m(-3) in a stepwise linear regression model (p < 0.0001). We concluded that seasonal moisture stress and high soil surface temperature imposed a strong limitation to limber pine seedling establishment across a broad elevation gradient, including at treeline, and that these limitations are likely to be enhanced by further climate warming.

Climate change is predicted to cause upward shifts in forest tree distributions, which will require seedling recruitment beyond current forest boundaries. However, predicting the likelihood of successful plant establishment beyond current species' ranges under changing climate is complicated by the interaction of genetic and environmental controls on seedling establishment. To determine how genetics and climate may interact to affect seedling establishment, we transplanted recently germinated seedlings from high- and low-elevation provenances (HI and LO, respectively) of Pinus flexilis in common gardens arrayed along an elevation and canopy gradient from subalpine forest into the alpine zone and examined differences in physiology and morphology between provenances and among sites. Plant dry mass, projected leaf area and shoot:root ratios were 12-40% greater in LO compared with HI seedlings at each elevation. There were no significant changes in these variables among sites except for decreased dry mass of LO seedlings in the alpine site. Photosynthesis, carbon balance (photosynthesis/respiration) and conductance increased >2× with elevation for both provenances, and were 35-77% greater in LO seedlings compared with HI seedlings. There were no differences in dark-adapted chlorophyll fluorescence (Fv/Fm) among sites or between provenances. Our results suggest that for P. flexilis seedlings, provenances selected for above-ground growth may outperform those selected for stress resistance in the absence of harsh climatic conditions, even well above the species' range limits in the alpine zone. This indicates that forest genetics may be important to understanding and managing species' range adjustments due to climate change.

Understanding how green sink strength is regulated in planta poses a difficult problem because non-structural carbohydrate (NSC) levels can have integrated, simultaneous feedback effects on photosynthesis, sugar uptake, and respiration that depend on specific NSC moieties. Photosynthetic gametophytes of the fern Ceratopteris richardii provide a simple land plant model to assess how different NSCs imported from the apoplast of intact plants affect green sink strength. Sink strength was quantified as the amount of exogenous sugar that plants grown in low light depleted from their liquid media, and the relative contributions of carbon assimilation by photosynthesis and sugar uptake was estimated from stable isotope analysis of plant dry mass. Gametophytes absorbed fructose and glucose with equal affinity when cultured on either hexose alone, or in the presence of an equimolar blend of both sugars. Plants also depleted sucrose from the surrounding media, although a portion of this disaccharide that was hydrolysed into fructose and glucose by putative cell wall invertase activity remained in the media. The δ(13)C in plant dry masses harvested from sugar treatments were all close to -18‰, indicating that 25-39% of total plant carbon was from C3 photosynthesis (δ(13)C=-29‰) and 61-75% was from uptake of exogenous sugars (δ(13)C=-11‰). Carbon-use efficiency (i.e. carbon accumulated/carbon depleted) was significantly improved when plants had a blend of exogenous sugars available compared with plants grown in a single hexose alone. Plants avoided complete down-regulation of photosynthesis even though a large excess of exogenous carbon fluxed through their cells.

The invasion and spread of exotic plants following land disturbance threatens semiarid ecosystems. In sagebrush steppe, soil water is scarce and is partitioned between deep-rooted perennial shrubs and shallower-rooted native forbs and grasses. Disturbances commonly remove shrubs, leaving grass-dominated communities, and may allow for the exploitation of water resources by the many species of invasive, tap-rooted forbs that are increasingly successful in this habitat. We hypothesized that exotic forb populations would benefit from increased soil water made available by removal of sagebrush, a foundation species capable of deep-rooting, in semiarid shrub-steppe ecosystems. To test this hypothesis, we used periodic matrix models to examine effects of experimental manipulations of soil water on population growth of two exotic forb species, Tragopogon dubius and Lactuca serriola, in sagebrush steppe of southern Idaho, USA. We used elasticity analyses to examine which stages in the life cycle of T. dubius and L. serriola had the largest relative influence on population growth. We studied the demography of T. dubius and L. serriola in three treatments: (1) control, in which vegetation was not disturbed, (2) shrubs removed, or (3) shrubs removed but winter-spring recharge of deep-soil water blocked by rainout shelters. The short-term population growth rate (Lambda) of T. dubius in the shrub-removal treatment was more than double that of T. dubius in either sheltered or control treatments, both of which had limited soil water. All L. serriola individuals that emerged in undisturbed sagebrush plots died, whereas Lambda of L. serriola was high (Lambda > 2.5) in all shrub-removal plots, whether they had rainout shelters or not. Population growth of both forbs in all treatments was most responsive to flowering and seed production, which are life stages that should be particularly reliant on deep-soil water, as well as seedling establishment, which is important to most plant populations, especially during invasion. These data indicate the importance of native species, in this case the dominant shrub, in influencing soil resources and restricting population growth of exotic plants. These results argue that management of invasive plants should focus not only on removal of nonnatives, but also on reestablishment of important native species.

Nonstructural carbohydrates (NSCs) are commonly used to assess the balance of carbon sources and sinks in plants. A notable application of this approach has been tests of hypotheses on carbon limitations of trees at their upper altitudinal limits, near the alpine. How NSCs vary in time is not well known in conifers during their critical seedling stage, despite the importance of knowing the temporal variations of NSCs to use snapshot measurements of NSCs to assess carbon balance. We measured NSCs in needles, separately as soluble sugars and starch; (1) over diurnal periods in seedlings of Pseudotsuga menziesii (Mirb.) Franco (a timberline species that does not occur up to treeline), (2) throughout the growth season in the seedlings of P. menziesii and Abies lasiocarpa (Hook.) Nutt. (a species that does occur up to treeline) growing along an elevation gradient in the timberline ecotone and furthermore (3) compared seedlings and co-occurring adults to assess variation with developmental stage. We also compared NSCs in seedlings grown under field or laboratory conditions to separate environmental from intrinsic factors affecting NSCs during early emergence. Diurnal variations in NSCs were minimal, especially when compared to seasonal variation, and were detectable mainly in relatively small midday maxima of soluble sugar concentrations. Seasonal patterns of NSCs were generally (and surprisingly) similar among field and laboratory seedlings and adults. Seasonal patterns of NSCs were dominated by progressive increases in soluble sugars until winter, and by early-season peaks in starch. Nonetheless, notable differences were detectable among ages, species and environmental conditions in (1) the timing and extent of the early-season maxima of starch and (2) the extent of the late-season maxima of soluble sugars. These differences in NSCs likely correspond with ecophysiologically relevant differences in carbon balance that could affect growth and survival of trees growing in the timberline ecotone.

Low rates of growth for conifers at high elevations may relate to problems in acquiring or utilizing carbon. A traditional hypothesis for growth limits of trees at alpine treeline is that low photosynthesis (A) leads to insufficient supply of carbon for growth. Alternatively, the sink-limitation hypothesis questions the importance of low A, and suggests that trees at treeline have abundant carbon for growth as a result of greater decreases in carbon use [respiration (R) and growth] than A at higher elevations. Concentrations of carbon intermediates (e.g., nonstructural carbohydrates, NSC) have been used to support the sink-limitation hypothesis, with the assumption that NSC reflects the ratio of carbon source to sinks. Our objective was to determine elevation effects on carbon balance (whole-plant uptake, storage, and efflux of carbon) of tree seedlings during their critical establishment phase at timberline. Changes in A and R (of whole crowns), NSC (starches and simple sugars), and growth were measured in seedlings of a treeline (Abies lasiocarpa) and nontreeline species (Pseudotsuga menziesii). Seedlings were outplanted at the lower (2,450 m) and upper (3,000 m) edges of the timberline zone in the Rocky Mountains, USA. At the upper compared with lower elevation, both species had 10-20% less root and needle growth, 13-15% less A, 35-39% less R, and up to 50% greater NSC. A. lasiocarpa allocated more biomass to needles and had greater A but less NSC than P. menziesii. The greater effects of elevation on R compared with A led to a 1.3-fold increase in A:R at the upper elevation, and a corresponding increase in starch (r2 = 0.42)-a pattern consistent with the predictions of the sink-limitation hypothesis. Nevertheless, A and dry mass gain were positively correlated (r2 = 0.42), indicating that variation in photosynthesis was related to growth of seedlings at timberline.

* Plants establishing in environments that are marginal for growth could be particularly sensitive to mycorrhizal associations. We investigated ectomycorrhizal colonization and its significance for young conifers growing at, or above, their normal limits for growth, in the alpine-treeline ecotone. * Colonization of seedlings (<1 yr old) and juveniles (2- to 10-yr-old) of Picea engelmannii and Abies lasiocarpa by Cenococcum geophilum was determined in a field study, and effects of Cenococcum on Picea seedling ecophysiology were investigated in a glasshouse. * Colonization by Cenococcum was c. 20-fold greater for juveniles than seedlings, and approximately 4-fold greater adjacent compared with approximately 7 m away from trees. Juveniles of Picea were more colonized at timberline than Abies, and the opposite relationship was observed in forest. Colonization enhanced seedling water potential, but not phosphorus concentrations or photosynthesis. * These landscape and age-dependent variations in colonization correspond well with known variations in conifer physiology and establishment near timberline. Facilitation of seedling establishment by older trees at alpine-treeline may include a below-ground, mycorrhizal component that complements previously reported effects of trees on the microclimate and ecophysiology of seedlings.

Most research on the occurrence and stability of alpine timberlines has focused on correlations between adult tree growth and mean temperatures rather than on specific mechanisms. Timberline migration to higher altitude is dependent on new seedling establishment in the tree-line ecotone; however, reductions in photosynthetic carbon gain in establishing seedlings have previously been interpreted solely in terms of decreased seedling survival. Our objective was to evaluate the impact of abiotic factors (temperature, light and water) on photosynthetic carbon gain in young seedlings of the two dominant conifer tree species occurring naturally above (tree-line ecotone site, TS) and below (forest site, FS) a Rocky Mountain timberline in southeastern Wyoming, USA. Coincidentally, measurements were made during an unusually dry summer. Mean daily photosynthesis in seedlings of both Abies lasiocarpa (Hook.) Nutt. (subalpine fir) and Picea engelmannii Parry ex Engelm. (Engelmann spruce) was less at TS than at FS (19 and 29%, respectively). Minimum nighttime temperatures below 2 degrees C were more frequent at TS than at FS and were associated with reduced maximum photosynthesis the following day. Low midday water potentials were associated with a reduction in carbon gain at both sites early in the season, prior to snowmelt, as well as late in the season when soils began to dry. However, the lower photosynthetic rates at TS than at FS appeared to be unrelated to seedling water status because seedlings at both sites had similar xylem pressure potentials. Solar irradiance was highly variable at both sites as a result of uneven shading by neighboring trees, although this variation was substantially reduced on cloudy days (44% of all days observed). Compared with sunny days, cloudy days resulted in greater integrated daily carbon gain at both sites (41% increase at TS and 69% increase at FS), based on a simulated photosynthesis model. Photosynthetic responses to temperature, sunlight and water suggest that variable solar irradiance and nighttime temperatures were major abiotic factors limiting photosynthetic carbon acquisition in these young seedlings, especially for seedlings growing in the tree-line ecotone.

Recent hypotheses of timberline causation include the possibility that limitations to growth processes may be more limiting than restrictions on photosynthetic carbon gain, and that cold soil is a primary limiting factor at high altitude. However, almost all of the supporting data for timberline causation have come from studies on older trees, with little focus on the mechanisms of seedling establishment and the growth of saplings away from the forest edge into the treeline ecotone. We describe a conceptual model of timberline migration that invokes a strong dependence on ecological facilitation, beginning with seed germination and continuing through seedling establishment and sapling growth to the stage where trees with forest-like stature form new subalpine forest at a higher altitude. In addition to protection from severe mechanical damage, facilitation of photosynthetic carbon gain and carbon processing is enhanced by plasticity in plant form and microsite preference, enabling seedling survival and sapling growth inside and through the often severe boundary layer just above the ground cover. Several forms of facilitation (inanimate, interspecific, intraspecific and structural) result in substantial increases in photosynthetic carbon gain throughout the summer growth period, leading to enhanced root growth, subsequent amelioration of drought stress, and increased seedling survival. Avoidance of low temperatures and low-temperature photoinhibition of photosynthesis may be major benefits of the facilitation, enhancing photosynthetic carbon gain and respiratory-driven growth processes. We propose that the growth of vertical stems (flagged tree forms) from krummholz mats is analogous functionally to the facilitated growth of a seedling/sapling in and away from ground cover. Increasing abundance and growth of newly established trees in the treeline ecotone generates a structural and microsite facilitation characteristic of the subalpine forest below. This is followed by the formation of new subalpine forest with forest-like trees, and a new timberline at higher altitude.

• The effects of supraoptimal soil temperatures on plants are not well known, despite their importance with fluctuating temperatures in many habitats. The effects of warm soils on carbon gain and water relations were evaluated for Dichanthelium lanuginosum , a grass that can persist in soils sustained at > 45°C by geothermal heating. • Microclimate, root and shoot biomass, photosynthetic gas exchange, and soil/plant water relations were measured under different root-zone temperatures in the field and glasshouse. • Carbon gain at midsummer decreased markedly for plants in 35-50°C vs c. 30°C soils, due to foliar necrosis and stomatal constraints to photosynthesis. Daily rates of dehydration, estimated from changes in leaf water potential, were also 30% greater for plants in the warmer soils. Soil water, evapotranspiration, and ratios of root : shoot biomass and area could not explain greater plant dehydration in warmer soils. • Reduced uptake of soil water may therefore lead to water stress and stomatal limitations to photosynthesis in warm soils, and may thereby influence the local-scale distribution of D. lanuginosum along steep gradients of soil-temperature within geothermal basins.