Trajectories and tipping points of piñon-juniper woodlands after fire and thinning.

Piñon-juniper (PJ) woodlands are a dominant community type across the Intermountain West, comprising over a million acres and experiencing critical effects from increasing wildfire. Large PJ mortality and regeneration failure after catastrophic wildfire have elevated concerns about the long-term viability of PJ woodlands. Thinning is increasingly used to safeguard forests from fire and in an attempt to increase climate resilience. We have only a limited understanding of how fire and thinning will affect the structure and function of PJ ecosystems. Here, we examined vegetation structure, microclimate conditions, and PJ regeneration dynamics following ~20 years post-fire and thinning treatments. We found that burned areas had undergone a state shift that did not show signs of returning to their previous state. This shift was characterized by (1) distinct plant community composition dominated by grasses; (2) a lack of PJ recruitment; (3) a decrease in the sizes of interspaces in between plants; (4) lower abundance of late successional biological soil crusts; (5) lower mean and minimum daily soil moisture values; (6) lower minimum daily vapor pressure deficit; and (7) higher photosynthetically active radiation. Thinning created distinct plant communities and served as an intermediate between intact and burned communities. More intensive thinning decreased PJ recruitment and late successional biocrust cover. Our results indicate that fire has the potential to create drier and more stressful microsite conditions, and that, in the absence of active management following fire, there may be shifts to persistent ecological states dominated by grasses. Additionally, more intensive thinning had a larger impact on community structure and recruitment than less intensive thinning, suggesting that careful consideration of goals could help avoid unintended consequences. While our results indicate the vulnerability of PJ ecosystems to fire, they also highlight management actions that could be adapted to create conditions that promote PJ re-establishment.

Where can managers effectively resist climate-driven ecological transformation in pinyon-juniper woodlands of the US Southwest?

Pinyon-juniper (PJ) woodlands are an important component of dryland ecosystems across the US West and are potentially susceptible to ecological transformation. However, predicting woodland futures is complicated by species-specific strategies for persisting and reproducing under drought conditions, uncertainty in future climate, and limitations to inferring demographic rates from forest inventory data. Here, we leverage new demographic models to quantify how climate change is expected to alter population demographics in five PJ tree species in the US West and place our results in the context of a climate adaptation framework to resist, accept, or direct ecological transformation. Two of five study species, Pinus edulis and Juniperus monosperma, are projected to experience population declines, driven by both rising mortality and decreasing recruitment rates. These declines are reasonably consistent across various climate futures, and the magnitude of uncertainty in population growth due to future climate is less than uncertainty due to how demographic rates will respond to changing climate. We assess the effectiveness of management to reduce tree density and mitigate competition, and use the results to classify southwest woodlands into areas where transformation is (a) unlikely and can be passively resisted, (b) likely but may be resisted by active management, and (c) likely unavoidable, requiring managers to accept or direct the trajectory. Population declines are projected to promote ecological transformation in the warmer and drier PJ communities of the southwest, encompassing 37.1%-81.1% of our sites, depending on future climate scenarios. Less than 20% of sites expected to transform away from PJ have potential to retain existing tree composition by density reduction. Our results inform where this adaptation strategy could successfully resist ecological transformation in coming decades and allow for a portfolio design approach across the geographic range of PJ woodlands.

Climate disequilibrium dominates uncertainty in long-term projections of primary productivity.

Rapid climate change may exceed ecosystems' capacities to respond through processes including phenotypic plasticity, compositional turnover and evolutionary adaption. However, consequences of the resulting climate disequilibria for ecosystem functioning are rarely considered in projections of climate change impacts. Combining statistical models fit to historical climate data and remotely-sensed estimates of herbaceous net primary productivity with an ensemble of climate models, we demonstrate that assumptions concerning the magnitude of climate disequilibrium are a dominant source of uncertainty: models assuming maximum disequilibrium project widespread decreases in productivity in the western US by 2100, while models assuming minimal disequilibrium project productivity increases. Uncertainty related to climate disequilibrium is larger than uncertainties from variation among climate models or emissions pathways. A better understanding of processes that regulate climate disequilibria is essential for improving long-term projections of ecological responses and informing management to maintain ecosystem functioning at historical baselines.

Biotic vs abiotic controls on temporal sensitivity of primary production to precipitation across North American drylands.

Dryland net primary productivity (NPP) is sensitive to temporal variation in precipitation (PPT), but the magnitude of this 'temporal sensitivity' varies spatially. Hypotheses for spatial variation in temporal sensitivity have often emphasized abiotic factors, such as moisture limitation, while overlooking biotic factors, such as vegetation structure. We tested these hypotheses using spatiotemporal models fit to remote-sensing data sets to assess how vegetation structure and climate influence temporal sensitivity across five dryland ecoregions of the western USA. Temporal sensitivity was higher in locations and ecoregions dominated by herbaceous vegetation. By contrast, much less spatial variation in temporal sensitivity was explained by mean annual PPT. In fact, ecoregion-specific models showed inconsistent associations of sensitivity and PPT; whereas sensitivity decreased with increasing mean annual PPT in most ecoregions, it increased with mean annual PPT in the most arid ecoregion, the hot deserts. The strong, positive influence of herbaceous vegetation on temporal sensitivity indicates that herbaceous-dominated drylands will be particularly sensitive to future increases in precipitation variability and that dramatic changes in cover type caused by invasions or shrub encroachment will lead to changes in dryland NPP dynamics, perhaps independent of changes in precipitation.

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

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

Landscape-scale restoration minimizes tree growth vulnerability to 21st century drought in a dry forest.

Increasing aridity is a challenge for forest managers and reducing stand density to minimize competition is a recognized strategy to mitigate drought impacts on growth. In many dry forests, the most widespread and common forest management programs currently being implemented focus on restoration of historical stand structures, primarily to minimize fire risk and enhance watershed function. The implications of these restoration projects for drought vulnerability are not well understood. Here, we examined how planned restoration treatments in the Four Forests Restoration Initiative, the largest forest restoration project in the United States, would alter landscape-scale patterns of forest growth and drought vulnerability throughout the 21st century. Using drought-growth relationships developed within the landscape, we considered a suite of climate and treatment scenarios and estimated average forest growth and the proportion of years with extremely low growth as a measure of vulnerability to long-term decline. Climatic shifts projected for this landscape include higher temperatures and shifting seasonal precipitation that promotes lower soil moisture availability in the early growing season and greater hot-dry stress, conditions negatively associated with tree growth. However, drought severity and the magnitude of future growth declines were moderated by the thinning treatments. Compared to historical conditions, proportional growth in mid-century declines by ~40% if thinning ceases or continues at the status quo pace. By comparison, proportional growth declines by only 20% if the Four Forest Restoration Initiative treatments are fully implemented, and <10% if stands are thinned even more intensively than currently planned. Furthermore, restoration treatments resulted in dramatically fewer years with extremely low growth in the future, a recognized precursor to forest decline and eventual tree mortality. Benefits from density reduction for mitigating drought-induced growth declines are more apparent in mid-century and under RCP4.5 than under RCP8.5 at the end of the century. Future climate is inherently uncertain, and our results only reflect the climate projections from the representative suite of models examined. Nevertheless, these results indicate that forest restoration projects designed for other objectives also have substantial benefits for minimizing future drought vulnerability in dry forests and provide additional incentive to accelerate the pace of restoration.

Forest density intensifies the importance of snowpack to growth in water-limited pine forests.

Warming climate and resulting declines in seasonal snowpack have been associated with drought stress and tree mortality in seasonally snow-covered watersheds worldwide. Meanwhile, increasing forest density has further exacerbated drought stress due to intensified tree-tree competition. Using a uniquely detailed data set of population-level forest growth (n = 2,495 sampled trees), we examined how inter-annual variability in growth relates to snow volume across a range of forest densities (e.g., competitive environments) in sites spanning a broad aridity gradient across the United States. Forest growth was positively related to snowpack in water-limited forests located at low latitude, and this relationship was intensified by forest density. However, forest growth was negatively related to snowpack in a higher latitude more energy-limited forest, and this relationship did not interact with forest density. Future reductions in snowpack may have contrasting consequences, as growth may respond positively in energy-limited forests and negatively in water-limited forests; however, these declines may be mitigated by reducing stand density through forest thinning.

Robust ecological drought projections for drylands in the 21st century.

Dryland ecosystems may be especially vulnerable to expected 21st century increases in temperature and aridity because they are tightly controlled by moisture availability. However, climate impact assessments in drylands are difficult because ecological dynamics are dictated by drought conditions that are difficult to define and complex to estimate from climate conditions alone. In addition, precipitation projections vary substantially among climate models, enhancing variation in overall trajectories for aridity. Here, we constrain this uncertainty by utilizing an ecosystem water balance model to quantify drought conditions with recognized ecological importance, and by identifying changes in ecological drought conditions that are robust among climate models, defined here as when >90% of models agree in the direction of change. Despite limited evidence for robust changes in precipitation, changes in ecological drought are robust over large portions of drylands in the United States and Canada. Our results suggest strong regional differences in long-term drought trajectories, epitomized by chronic drought increases in southern areas, notably the Upper Gila Mountains and South-Central Semi-arid Prairies, and decreases in the north, particularly portions of the Temperate and West-Central Semi-arid Prairies. However, we also found that exposure to hot-dry stress is increasing faster than mean annual temperature over most of these drylands, and those increases are greatest in northern areas. Robust shifts in seasonal drought are most apparent during the cool season; when soil water availability is projected to increase in northern regions and decrease in southern regions. The implications of these robust drought trajectories for ecosystems will vary geographically, and these results provide useful insights about the impact of climate change on these dryland ecosystems. More broadly, this approach of identifying robust changes in ecological drought may be useful for other assessments of climate impacts in drylands and provide a more rigorous foundation for making long-term strategic resource management decisions.

Assessing the ecological impacts of biomass harvesting along a disturbance severity gradient.

Disturbance is a central driver of forest development and ecosystem processes with variable effects within and across ecosystems. Despite the high levels of variation in disturbance severity often observed in forests following natural and anthropogenic disturbance, studies quantifying disturbance impacts often rely on categorical classifications, thus limiting opportunities to examine potential gradients in ecosystem response to a given disturbance or management regime. Given the potential increases in disturbance severity associated with global change, as well as shifts in management regimes related to procurement of biofuel feedstocks, there is an increasing need to quantitatively describe disturbance severity and associated responses of forest development, soil processes, and structural conditions. This study took advantage of two replicated large-scale studies of forest biomass harvesting in Populus tremuloides and Pinus bansksiana forests, respectively, to develop and test the utility of a continuous, quantitative, disturbance severity index (DSI) for describing postharvest response of plant communities and nutrient pools to different levels of biomass removal and legacy retention (i.e., live trees and coarse woody material). There was a high degree of variability in DSI within categorical treatments associated with different levels of legacy retention and regression models using DSI as a predictor explained a portion of the variation (>50%) for many of the ecosystem- and community-level responses to biomass harvesting examined. Nutrient losses associated with biomass harvesting were positively related to disturbance severity, particularly in P. tremuloides forests, with postharvest nutrient availability generally declining along the gradient of impacts. Consistent with expectations from ecological theory, species richness and diversity of woody plant communities were greatest at intermediate disturbance severities and regeneration densities of dominant trees species were most abundant at highest levels of disturbance. Although categorical benchmarks will continue to be the primary way through which management guidelines are conveyed to practitioners, evaluation of their effectiveness at sustaining ecosystem functioning should be through continuous analyses, such as the DSI approach used in this study, to allow for the more precise identification of thresholds that ensure a range of desirable outcomes exist across managed landscapes.

A Comparison of Sexual Arousal in Men Exposed to Visual Stimuli With and Without Facial Blurring.

The role of the facial images in arousal and attraction has been examined before but never via penile plethysmography (PPG). This retrospective chart review aimed to determine the significance and magnitude of differences in arousal measured by PPG in 1,000 men exposed to slide stimuli with or without facial blurring in subjects of various ages. Arousal in response to blurred stimuli was significantly higher than nonanonymized stimuli with modest effect sizes for slides across age and gender categories. Facial blurring increased differences in arousal between adults and adolescents with a modest effect size. Our findings support the use of facial blurring to further protect the anonymity of models and limit the ethical and legal challenges of using slide stimuli with child models.

Transient population dynamics impede restoration and may promote ecosystem transformation after disturbance.

The apparent failure of ecosystems to recover from increasingly widespread disturbance is a global concern. Despite growing focus on factors inhibiting resilience and restoration, we still know very little about how demographic and population processes influence recovery. Using inverse and forward demographic modelling of 531 post-fire sagebrush populations across the western US, we show that demographic processes during recovery from seeds do not initially lead to population growth but rather to years of population decline, low density, and risk of extirpation after disturbance and restoration, even at sites with potential to support long-term, stable populations. Changes in population structure, and resulting transient population dynamics, lead to a > 50% decline in population growth rate after disturbance and significant reductions in population density. Our results indicate that demographic processes influence the recovery of ecosystems from disturbance and that demographic analyses can be used by resource managers to anticipate ecological transformation risk.

Adapting management to a changing world: Warm temperatures, dry soil, and interannual variability limit restoration success of a dominant woody shrub in temperate drylands.

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 km2 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.

Increasing temperature seasonality may overwhelm shifts in soil moisture to favor shrub over grass dominance in Colorado Plateau drylands.

Ecosystems in the southwestern U.S. are predicted to experience continued warming and drying trends of the early twenty-first century. Climate change can shift the balance between grass and woody plant abundance in these water-limited systems, which has large implications for biodiversity and ecosystem processes. However, variability in topo-edaphic conditions, notably soil texture and depth, confound efforts to quantify specific climatic controls over grass vs. shrub dominance. Here, we utilized weather records and a mechanistic soil water model to identify the timing and depth at which soil moisture related most strongly to the balance between grass and shrub dominance in the southern Colorado Plateau. Shrubs dominate where there is high soil moisture availability during winter, and where temperature is more seasonally variable, while grasses are favored where moisture is available during summer. Climate change projections indicate consistent increases in mean temperature and seasonal temperature variability for all sites, but predictions for summer and winter soil moisture vary across sites. Together, these changes in temperature and soil moisture are expected to shift the balance towards increasing shrub dominance across the region. These patterns are strongly driven by changes in temperature, which either enhance or overwhelm effects of changes in soil moisture across sites. This approach, which incorporates local, edaphic factors at sites protected from disturbance, improves understanding of climate change impacts on grass vs. shrub abundance and may be useful in other dryland regions with high edaphic and climatic heterogeneity.

Climate change-induced vegetation shifts lead to more ecological droughts despite projected rainfall increases in many global temperate drylands.

Drylands occur worldwide and are particularly vulnerable to climate change because dryland ecosystems depend directly on soil water availability that may become increasingly limited as temperatures rise. Climate change will both directly impact soil water availability and change plant biomass, with resulting indirect feedbacks on soil moisture. Thus, the net impact of direct and indirect climate change effects on soil moisture requires better understanding. We used the ecohydrological simulation model SOILWAT at sites from temperate dryland ecosystems around the globe to disentangle the contributions of direct climate change effects and of additional indirect, climate change-induced changes in vegetation on soil water availability. We simulated current and future climate conditions projected by 16 GCMs under RCP 4.5 and RCP 8.5 for the end of the century. We determined shifts in water availability due to climate change alone and due to combined changes of climate and the growth form and biomass of vegetation. Vegetation change will mostly exacerbate low soil water availability in regions already expected to suffer from negative direct impacts of climate change (with the two RCP scenarios giving us qualitatively similar effects). By contrast, in regions that will likely experience increased water availability due to climate change alone, vegetation changes will counteract these increases due to increased water losses by interception. In only a small minority of locations, climate change-induced vegetation changes may lead to a net increase in water availability. These results suggest that changes in vegetation in response to climate change may exacerbate drought conditions and may dampen the effects of increased precipitation, that is, leading to more ecological droughts despite higher precipitation in some regions. Our results underscore the value of considering indirect effects of climate change on vegetation when assessing future soil moisture conditions in water-limited ecosystems.

Variable effects of climate on forest growth in relation to climate extremes, disturbance, and forest dynamics.

Changes in the frequency, duration, and severity of climate extremes are forecast to occur under global climate change. The impacts of climate extremes on forest productivity and health remain difficult to predict due to potential interactions with disturbance events and forest dynamics-changes in forest stand composition, density, size and age structure over time. Such interactions may lead to non-linear forest growth responses to climate involving thresholds and lag effects. Understanding how forest dynamics influence growth responses to climate is particularly important given stand structure and composition can be modified through management to increase forest resistance and resilience to climate change. To inform such adaptive management, we develop a hierarchical Bayesian state space model in which climate effects on tree growth are allowed to vary over time and in relation to past climate extremes, disturbance events, and forest dynamics. The model is an important step toward integrating disturbance and forest dynamics into predictions of forest growth responses to climate extremes. We apply the model to a dendrochronology data set from forest stands of varying composition, structure, and development stage in northeastern Minnesota that have experienced extreme climate years and forest tent caterpillar defoliation events. Mean forest growth was most sensitive to water balance variables representing climatic water deficit. Forest growth responses to water deficit were partitioned into responses driven by climatic threshold exceedances and interactions with insect defoliation. Forest growth was both resistant and resilient to climate extremes with the majority of forest growth responses occurring after multiple climatic threshold exceedances across seasons and years. Interactions between climate and disturbance were observed in a subset of years with insect defoliation increasing forest growth sensitivity to water availability. Forest growth was particularly sensitive to climate extremes during periods of high stem density following major regeneration events when average inter-tree competition was high. Results suggest the resistance and resilience of forest growth to climate extremes can be increased through management steps such as thinning to reduce competition during early stages of stand development and small-group selection harvests to maintain forest structures characteristic of older, mature stands.

Temperature drives global patterns in forest biomass distribution in leaves, stems, and roots.

Whether the fraction of total forest biomass distributed in roots, stems, or leaves varies systematically across geographic gradients remains unknown despite its importance for understanding forest ecology and modeling global carbon cycles. It has been hypothesized that plants should maintain proportionally more biomass in the organ that acquires the most limiting resource. Accordingly, we hypothesize greater biomass distribution in roots and less in stems and foliage in increasingly arid climates and in colder environments at high latitudes. Such a strategy would increase uptake of soil water in dry conditions and of soil nutrients in cold soils, where they are at low supply and are less mobile. We use a large global biomass dataset (>6,200 forests from 61 countries, across a 40 °C gradient in mean annual temperature) to address these questions. Climate metrics involving temperature were better predictors of biomass partitioning than those involving moisture availability, because, surprisingly, fractional distribution of biomass to roots or foliage was unrelated to aridity. In contrast, in increasingly cold climates, the proportion of total forest biomass in roots was greater and in foliage was smaller for both angiosperm and gymnosperm forests. These findings support hypotheses about adaptive strategies of forest trees to temperature and provide biogeographically explicit relationships to improve ecosystem and earth system models. They also will allow, for the first time to our knowledge, representations of root carbon pools that consider biogeographic differences, which are useful for quantifying whole-ecosystem carbon stocks and cycles and for assessing the impact of climate change on forest carbon dynamics.

Mid-latitude shrub steppe plant communities: climate change consequences for soil water resources.

In the coming century, climate change is projected to impact precipitation and temperature regimes worldwide, with especially large effects in drylands. We use big sagebrush ecosystems as a model dryland ecosystem to explore the impacts of altered climate on ecohydrology and the implications of those changes for big sagebrush plant communities using output from 10 Global Circulation Models (GCMs) for two representative concentration pathways (RCPs). We ask: (1) What is the magnitude of variability in future temperature and precipitation regimes among GCMs and RCPs for big sagebrush ecosystems, and (2) How will altered climate and uncertainty in climate forecasts influence key aspects of big sagebrush water balance? We explored these questions across 1980-2010, 2030-2060, and 2070-2100 to determine how changes in water balance might develop through the 21st century. We assessed ecohydrological variables at 898 sagebrush sites across the western US using a process-based soil water model, SOILWAT, to model all components of daily water balance using site-specific vegetation parameters and site-specific soil properties for multiple soil layers. Our modeling approach allowed for changes in vegetation based on climate. Temperature increased across all GCMs and RCPs, whereas changes in precipitation were more variable across GCMs. Winter and spring precipitation was predicted to increase in the future (7% by 2030-2060, 12% by 2070-2100), resulting in slight increases in soil water potential (SWP) in winter. Despite wetter winter soil conditions, SWP decreased in late spring and summer due to increased evapotranspiration (6% by 2030-2060, 10% by 2070-2100) and groundwater recharge (26% and 30% increase by 2030-2060 and 2070-2100). Thus, despite increased precipitation in the cold season, soils may dry out earlier in the year, resulting in potentially longer, drier summer conditions. If winter precipitation cannot offset drier summer conditions in the future, we expect big sagebrush regeneration and survival will be negatively impacted, potentially resulting in shifts in the relative abundance of big sagebrush plant functional groups. Our results also highlight the importance of assessing multiple GCMs to understand the range of climate change outcomes on ecohydrology, which was contingent on the GCM chosen.

Predicting tree biomass growth in the temperate-boreal ecotone: Is tree size, age, competition, or climate response most important?

As global temperatures rise, variation in annual climate is also changing, with unknown consequences for forest biomes. Growing forests have the ability to capture atmospheric CO2 and thereby slow rising CO2 concentrations. Forests' ongoing ability to sequester C depends on how tree communities respond to changes in climate variation. Much of what we know about tree and forest response to climate variation comes from tree-ring records. Yet typical tree-ring datasets and models do not capture the diversity of climate responses that exist within and among trees and species. We address this issue using a model that estimates individual tree response to climate variables while accounting for variation in individuals' size, age, competitive status, and spatially structured latent covariates. Our model allows for inference about variance within and among species. We quantify how variables influence aboveground biomass growth of individual trees from a representative sample of 15 northern or southern tree species growing in a transition zone between boreal and temperate biomes. Individual trees varied in their growth response to fluctuating mean annual temperature and summer moisture stress. The variation among individuals within a species was wider than mean differences among species. The effects of mean temperature and summer moisture stress interacted, such that warm years produced positive responses to summer moisture availability and cool years produced negative responses. As climate models project significant increases in annual temperatures, growth of species like Acer saccharum, Quercus rubra, and Picea glauca will vary more in response to summer moisture stress than in the past. The magnitude of biomass growth variation in response to annual climate was 92-95% smaller than responses to tree size and age. This means that measuring or predicting the physical structure of current and future forests could tell us more about future C dynamics than growth responses related to climate change alone.

Scale dependence of disease impacts on quaking aspen (Populus tremuloides) mortality in the southwestern United States.

Depending on how disease impacts tree exposure to risk, both the prevalence of disease and disease effects on survival may contribute to patterns of mortality risk across a species' range. Disease may accelerate tree species' declines in response to global change factors, such as drought, biotic interactions, such as competition, or functional traits, such as allometry. To assess the role of disease in mediating mortality risk in quaking aspen (Populus tremuloides), we developed hierarchical Bayesian models for both disease prevalence in live aspen stems and the resulting survival rates of healthy and diseased aspen near the species' southern range limit using 5088 individual trees on 281 United States Forest Service Forest Inventory and Analysis plots in the southwestern United States. We found that disease prevalence depended primarily on tree size, tree allometry, and spatial variation in precipitation, while mortality depended on tree size, allometry, competition, spatial variation in summer temperature, and both temporal and spatial variation in summer precipitation. Disease prevalence was highest in large trees with low slenderness found on dry sites. For healthy trees, mortality decreased with diameter, slenderness, and temporal variation in summer precipitation, but increased with competition and spatial variation in summer temperature. Mortality of diseased trees decreased with diameter and aspen relative basal area and increased with mean summer temperature and precipitation. Disease infection increased aspen mortality, especially in trees of intermediate size and trees on plots at climatic extremes (i.e., cool, wet and warm, dry climates). By examining variation in disease prevalence, mortality of healthy trees, and mortality of diseased trees, we showed that the role of disease in aspen tree mortality depended on the scale of inference. For variation among individuals in diameter, disease tended to expose intermediate-size trees experiencing moderate risk to greater risk. For spatial variation in summer temperature, disease exposed lower risk populations to greater mortality probabilities, but the magnitude of this exposure depended on summer precipitation. Furthermore, the importance of diameter and slenderness in mediating responses to climate supports the increasing emphasis on trait variation in studies of ecological responses to global change.

Desert grassland responses to climate and soil moisture suggest divergent vulnerabilities across the southwestern United States.

Climate change predictions include warming and drying trends, which are expected to be particularly pronounced in the southwestern United States. In this region, grassland dynamics are tightly linked to available moisture, yet it has proven difficult to resolve what aspects of climate drive vegetation change. In part, this is because it is unclear how heterogeneity in soils affects plant responses to climate. Here, we combine climate and soil properties with a mechanistic soil water model to explain temporal fluctuations in perennial grass cover, quantify where and the degree to which incorporating soil water dynamics enhances our ability to understand temporal patterns, and explore the potential consequences of climate change by assessing future trajectories of important climate and soil water variables. Our analyses focused on long-term (20-56 years) perennial grass dynamics across the Colorado Plateau, Sonoran, and Chihuahuan Desert regions. Our results suggest that climate variability has negative effects on grass cover, and that precipitation subsidies that extend growing seasons are beneficial. Soil water metrics, including the number of dry days and availability of water from deeper (>30 cm) soil layers, explained additional grass cover variability. While individual climate variables were ranked as more important in explaining grass cover, collectively soil water accounted for 40-60% of the total explained variance. Soil water conditions were more useful for understanding the responses of C3 than C4 grass species. Projections of water balance variables under climate change indicate that conditions that currently support perennial grasses will be less common in the future, and these altered conditions will be more pronounced in the Chihuahuan Desert and Colorado Plateau. We conclude that incorporating multiple aspects of climate and accounting for soil variability can improve our ability to understand patterns, identify areas of vulnerability, and predict the future of desert grasslands.