Sierra Nevada reference conditions: A dataset of contemporary reference sites and corresponding remote sensing-derived forest structure metrics for yellow pine and mixed-conifer forests.

Contemporary reference sites in California's Sierra Nevada represent areas where a frequent, low-intensity fire regime - an integral ecological process in temperate dry forests - has been reintroduced after several decades of fire suppression. Produced by an intact fire regime, forest structural patterns in these sites are likely more resilient to future disturbances and climate, and thus can provide reference conditions to guide management and ecological research. In this paper, we present a set of 119 delineated contemporary reference sites in the Sierra Nevada yellow pine and mixed-conifer zone along with a suite of key remote sensing-derived forest structure metrics representing conditions within these sites. We also provide a set of summary figures for individual reference sites and sites grouped by dominant climate class. We identified restored frequent-fire landscapes using a combination of fire history, burn severity, management history, and forest type datasets and we delineated individual polygons using catchment basins, fire perimeters, and imagery. Reference sites ranged in size from 101-966 ha with a mean size of 240 ha. Where available (for 59 sites), we used airborne lidar datasets to characterize a suite of key forest structure metrics within reference sites. Across all 119 sites, we provide a set of forest structure metrics produced by the California Forest Observatory. Reference sites were categorized based on their dominant climate class to assist users in identifying the most climatically relevant reference conditions for their project or study area. We encourage the use of the reference sites and associated forest structure datasets for guiding ecologically focused forest management and research in the Sierra Nevada.

Conifer water-use patterns across temporal and topographic gradients in the southern Sierra Nevada.

Climate change is increasing the severity and duration of drought events experienced by forest ecosystems. Because water is essential for tree physiological processes, the ability of trees to survive prolonged droughts will largely depend on whether they have access to reliable water sources. While many woody plant species exhibit the ability to shift water sources between different depths of soil and rock water in response to changes in climate and water availability, it is unclear if Sierra Nevada conifers exhibit this plasticity. Here we analysed the δ18O and δ13C values of annual tree rings to determine the water-use patterns of large Sierra Nevada conifers during the 2012-16 California drought and 4 years before this drought event (2004-07). We analysed four species (Pinus jeffreyi Grev. & Balf. (Jeffrey pine), Pinus lambertiana Dougl. (sugar pine), Abies concolor (Gord. & Glend.) Lindl. Ex Hilderbr (white fir) and Calocedrus decurrens (Torr.) Florin (incense-cedar)) across a range of topographic positions to investigate differences in water-use patterns by species and position on the landscape. We found no significant differences in δ18O and δ13C values for the pre-drought and drought periods. This stability in δ18O values suggests that trees did not shift their water-use patterns in response to the 2012-16 drought. We did find species-specific differences in water-use patterns, with incense-cedar exhibiting more depleted δ18O values than all other species. We also found trends that suggest the water source used by a tree may depend on topographic and growing environment attributes such as topographic wetness and the surrounding basal area. Overall, our results suggest that the water source used by trees varies by the species and topographic position, but that Sierra Nevada conifers do not switch their water-use patterns in response to the drought. This lack of plasticity could make Sierra Nevada conifers particularly vulnerable to drought mortality as their historically reliable water sources begin to dry out with climate change.

Adapting western North American forests to climate change and wildfires: 10 common questions.

We review science-based adaptation strategies for western North American (wNA) forests that include restoring active fire regimes and fostering resilient structure and composition of forested landscapes. As part of the review, we address common questions associated with climate adaptation and realignment treatments that run counter to a broad consensus in the literature. These include the following: (1) Are the effects of fire exclusion overstated? If so, are treatments unwarranted and even counterproductive? (2) Is forest thinning alone sufficient to mitigate wildfire hazard? (3) Can forest thinning and prescribed burning solve the problem? (4) Should active forest management, including forest thinning, be concentrated in the wildland urban interface (WUI)? (5) Can wildfires on their own do the work of fuel treatments? (6) Is the primary objective of fuel reduction treatments to assist in future firefighting response and containment? (7) Do fuel treatments work under extreme fire weather? (8) Is the scale of the problem too great? Can we ever catch up? (9) Will planting more trees mitigate climate change in wNA forests? And (10) is post-fire management needed or even ecologically justified? Based on our review of the scientific evidence, a range of proactive management actions are justified and necessary to keep pace with changing climatic and wildfire regimes and declining forest heterogeneity after severe wildfires. Science-based adaptation options include the use of managed wildfire, prescribed burning, and coupled mechanical thinning and prescribed burning as is consistent with land management allocations and forest conditions. Although some current models of fire management in wNA are averse to short-term risks and uncertainties, the long-term environmental, social, and cultural consequences of wildfire management primarily grounded in fire suppression are well documented, highlighting an urgency to invest in intentional forest management and restoration of active fire regimes.

Cross-scale interaction of host tree size and climatic water deficit governs bark beetle-induced tree mortality.

The recent Californian hot drought (2012-2016) precipitated unprecedented ponderosa pine (Pinus ponderosa) mortality, largely attributable to the western pine beetle (Dendroctonus brevicomis; WPB). Broad-scale climate conditions can directly shape tree mortality patterns, but mortality rates respond non-linearly to climate when local-scale forest characteristics influence the behavior of tree-killing bark beetles (e.g., WPB). To test for these cross-scale interactions, we conduct aerial drone surveys at 32 sites along a gradient of climatic water deficit (CWD) spanning 350 km of latitude and 1000 m of elevation in WPB-impacted Sierra Nevada forests. We map, measure, and classify over 450,000 trees within 9 km2, validating measurements with coincident field plots. We find greater size, proportion, and density of ponderosa pine (the WPB host) increase host mortality rates, as does greater CWD. Critically, we find a CWD/host size interaction such that larger trees amplify host mortality rates in hot/dry sites. Management strategies for climate change adaptation should consider how bark beetle disturbances can depend on cross-scale interactions, which challenge our ability to predict and understand patterns of tree mortality.

Changing climate reallocates the carbon debt of frequent-fire forests.

Ongoing climate change will alter the carbon carrying capacity of forests as they adjust to climatic extremes and changing disturbance regimes. In frequent-fire forests, increasing drought frequency and severity are already causing widespread tree mortality events, which can exacerbate the carbon debt that has developed as a result of fire exclusion. Forest management techniques that reduce tree density and surface fuels decrease the risk of high-severity wildfire and may also limit drought-induced mortality by reducing competition. We used a long-term thinning and burning experiment in a mixed-conifer forest to investigate the effects of the 2012-2015 California drought on forest carbon dynamics in each treatment, including the carbon emissions from a second-entry prescribed fire that followed the drought. We assessed differences in carbon stability and tree survival across treatments, expecting that both carbon stability and survival probability would increase with increasing treatment intensity (decreasing basal area). Additionally, we analyzed the effects of drought- mortality on second-entry burn emissions and compared emissions for the first- and second-entry burns. We found a non-linear relationship between treatment intensity and carbon stability, which was in part driven by varying relationships between individual tree growing space and survival across treatments. Drought mortality increased dead tree and surface fuel carbon in all treatments, which contributed to higher second-entry burn emissions for two of the three burn treatments when compared to the first burn. Our findings suggest that restoration treatments will not serve as a panacea for ongoing climate change and that the carbon debt of these forests will become increasingly unstable as the carbon carrying capacity adjusts to severe drought events. Managing the carbon debt with prescribed fire will help reduce the risk of additional mortality from wildfire, but at an increasing carbon cost for forest management.

Wildfire-Driven Forest Conversion in Western North American Landscapes.

Changing disturbance regimes and climate can overcome forest ecosystem resilience. Following high-severity fire, forest recovery may be compromised by lack of tree seed sources, warmer and drier postfire climate, or short-interval reburning. A potential outcome of the loss of resilience is the conversion of the prefire forest to a different forest type or nonforest vegetation. Conversion implies major, extensive, and enduring changes in dominant species, life forms, or functions, with impacts on ecosystem services. In the present article, we synthesize a growing body of evidence of fire-driven conversion and our understanding of its causes across western North America. We assess our capacity to predict conversion and highlight important uncertainties. Increasing forest vulnerability to changing fire activity and climate compels shifts in management approaches, and we propose key themes for applied research coproduced by scientists and managers to support decision-making in an era when the prefire forest may not return.

Local forest structure variability increases resilience to wildfire in dry western U.S. coniferous forests.

A 'resilient' forest endures disturbance and is likely to persist. Resilience to wildfire may arise from feedback between fire behaviour and forest structure in dry forest systems. Frequent fire creates fine-scale variability in forest structure, which may then interrupt fuel continuity and prevent future fires from killing overstorey trees. Testing the generality and scale of this phenomenon is challenging for vast, long-lived forest ecosystems. We quantify forest structural variability and fire severity across >30 years and >1000 wildfires in California's Sierra Nevada. We find that greater variability in forest structure increases resilience by reducing rates of fire-induced tree mortality and that the scale of this effect is local, manifesting at the smallest spatial extent of forest structure tested (90 × 90 m). Resilience of these forests is likely compromised by structural homogenisation from a century of fire suppression, but could be restored with management that increases forest structural variability.

Extreme mid-winter drought weakens tree hydraulic-carbohydrate systems and slows growth.

Rising temperatures and extended periods of drought compromise tree hydraulic and carbohydrate systems, threatening forest health globally. Despite winter's biological significance to many forests, the effects of warmer and dryer winters on tree hydraulic and carbohydrate status have largely been overlooked. Here we report a sharp and previously unknown decline in stem water content of three conifer species during California's anomalous 2015 mid-winter drought that was followed by dampened spring starch accumulation. Recent precipitation and seasonal vapor pressure deficit (VPD) anomaly, not absolute VPD, best predicted the hydraulic patterns observed. By linking relative water content and hydraulic conductivity (Kh ), we estimated that stand-level Kh declined by 52% during California's 2015 mid-winter drought, followed by a 50% reduction in spring starch accumulation. Further examination of tree increment records indicated a concurrent decline of growth with rising mid-winter, but not summer, VPD anomaly. Thus, our findings suggest a seasonality to tree hydraulic and carbohydrate declines, with consequences for annual growth rates, raising novel physiological and ecological questions about how rising winter temperatures will affect forest vitality as climate changes.

Leap frog in slow motion: Divergent responses of tree species and life stages to climatic warming in Great Basin subalpine forests.

In response to climate warming, subalpine treelines are expected to move up in elevation since treelines are generally controlled by growing season temperature. Where treeline is advancing, dispersal differences and early life stage environmental tolerances are likely to affect how species expand their ranges. Species with an establishment advantage will colonize newly available habitat first, potentially excluding species that have slower establishment rates. Using a network of plots across five mountain ranges, we described patterns of upslope elevational range shift for the two dominant Great Basin subalpine species, limber pine and Great Basin bristlecone pine. We found that the Great Basin treeline for these species is expanding upslope with a mean vertical elevation shift of 19.1 m since 1950, which is lower than what we might expect based on temperature increases alone. The largest advances were on limber pine-dominated granitic soils, on west aspects, and at lower latitudes. Bristlecone pine juveniles establishing above treeline share some environmental associations with bristlecone adults. Limber pine above-treeline juveniles, in contrast, are prevalent across environmental conditions and share few environmental associations with limber pine adults. Strikingly, limber pine is establishing above treeline throughout the region without regard to site characteristic such as soil type, slope, aspect, or soil texture. Although limber pine is often rare at treeline where it coexists with bristlecone pine, limber pine juveniles dominate above treeline even on calcareous soils that are core bristlecone pine habitat. Limber pine is successfully "leap-frogging" over bristlecone pine, probably because of its strong dispersal advantage and broader tolerances for establishment. This early-stage dominance indicates the potential for the species composition of treeline to change in response to climate change. More broadly, it shows how species differences in dispersal and establishment may result in future communities with very different specific composition.

Online versus face-to-face pulmonary rehabilitation for patients with chronic obstructive pulmonary disease: randomised controlled trial.

OBJECTIVE: To obtain evidence whether the online pulmonary rehabilitation(PR) programme 'my-PR' is non-inferior to a conventional face-to-face PR in improving physical performance and symptom scores in patients with COPD. DESIGN: A two-arm parallel single-blind, randomised controlled trial. SETTING: The online arm carried out pulmonary rehabilitation in their own homes and the face to face arm in a local rehabilitation facility. PARTICIPANTS: 90 patients with a diagnosis of chronic obstructive pulmonary disease (COPD), modified Medical Research Council score of 2 or greater referred for pulmonary rehabilitation (PR), randomised in a 2:1 ratio to online (n=64) or face-to-face PR (n=26). Participants unable to use an internet-enabled device at home were excluded. MAIN OUTCOME MEASURES: Coprimary outcomes were 6 min walk distance test and the COPD assessment test (CAT) score at completion of the programme. INTERVENTIONS: A 6-week PR programme organised either as group sessions in a local rehabilitation facility, or online PR via log in and access to 'myPR'. RESULTS: The adjusted mean difference for the 6 min walk test (6MWT) between groups for the intention-to-treat (ITT) population was 23.8 m with the lower 95% CI well above the non-inferiority threshold of -40.5 m at -4.5 m with an upper 95% CI of +52.2 m. This result was consistent in the per-protocol (PP) population with a mean adjusted difference of 15 m (-13.7 to 43.8). The CAT score difference in the ITT was -1.0 in favour of the online intervention with the upper 95% CI well below the non-inferiority threshold of 1.8 at 0.86 and the lower 95% CI of -2.9. The PP analysis was consistent with the ITT. CONCLUSION: PR is an evidenced-based and guideline-mandated intervention for patients with COPD with functional limitation. A 6-week programme of online-supported PR was non-inferior to a conventional model delivered in face-to-face sessions in terms of effects on 6MWT distance, and symptom scores and was safe and well tolerated.

Evaluating a new method for reconstructing forest conditions from General Land Office survey records.

Historical forest conditions are often used to inform contemporary management goals because historical forests are considered to be resilient to ecological disturbances. The General Land Office (GLO) surveys of the late 19th and early 20th centuries provide regionally quasi-contiguous data sets of historical forests across much of the Western United States. Multiple methods exist for estimating tree density from point-based sampling such as the GLO surveys, including distance-based and area-based approaches. Area-based approaches have been applied in California mixed-conifer forests but their estimates have not been validated. To assess the accuracy and precision of plotless density estimators with potential for application to GLO data in this region, we imposed a GLO sampling scheme on six mapped forest stands of known densities (159-784 trees/ha) in the Sierra Nevada in California, USA, and Baja California Norte, Mexico. We compared three distance-based plotless density estimators (Cottam, Pollard, and Morisita) as well as two Voronoi area (VA) estimators, the Delincé and mean harmonic Voronoi density (MHVD), to the true densities. We simulated sampling schemes of increasing intensity to assess sampling error. The relative error (RE) of density estimates for the GLO sampling scheme ranged from 0.36 to 4.78. The least biased estimate of tree density in every stand was obtained with the Morisita estimator and the most biased was obtained with the MHVD estimator. The MHVD estimates of tree density were 1.2-3.8 times larger than the true densities and performed best in stands subject to fire exclusion for 100 yr. The Delincé approach obtained accurate estimates of density, implying that the Voronoi approach is theoretically sound but that its application in the MHVD was flawed. The misapplication was attributed to two causes: (1) the use of a crown scaling factor that does not correct for the number of trees sampled and (2) the persistent underestimate of the true VA due to a weak relationship between tree size and VA. The magnitude of differences between true densities and MHVD estimates suggest caution in using results based on the MHVD to inform management and restoration practices in the conifer forests of the American West.

Restoring forest structure and process stabilizes forest carbon in wildfire-prone southwestern ponderosa pine forests.

Changing climate and a legacy of fire-exclusion have increased the probability of high-severity wildfire, leading to an increased risk of forest carbon loss in ponderosa pine forests in the southwestern USA. Efforts to reduce high-severity fire risk through forest thinning and prescribed burning require both the removal and emission of carbon from these forests, and any potential carbon benefits from treatment may depend on the occurrence of wildfire. We sought to determine how forest treatments alter the effects of stochastic wildfire events on the forest carbon balance. We modeled three treatments (control, thin-only, and thin and burn) with and without the occurrence of wildfire. We evaluated how two different probabilities of wildfire occurrence, 1% and 2% per year, might alter the carbon balance of treatments. In the absence of wildfire, we found that thinning and burning treatments initially reduced total ecosystem carbon (TEC) and increased net ecosystem carbon balance (NECB). In the presence of wildfire, the thin and burn treatment TEC surpassed that of the control in year 40 at 2%/yr wildfire probability, and in year 51 at 1%/yr wildfire probability. NECB in the presence of wildfire showed a similar response to the no-wildfire scenarios: both thin-only and thin and burn treatments increased the C sink. Treatments increased TEC by reducing both mean wildfire severity and its variability. While the carbon balance of treatments may differ in more productive forest types, the carbon balance benefits from restoring forest structure and fire in southwestern ponderosa pine forests are clear.

Average Stand Age from Forest Inventory Plots Does Not Describe Historical Fire Regimes in Ponderosa Pine and Mixed-Conifer Forests of Western North America.

Quantifying historical fire regimes provides important information for managing contemporary forests. Historical fire frequency and severity can be estimated using several methods; each method has strengths and weaknesses and presents challenges for interpretation and verification. Recent efforts to quantify the timing of historical high-severity fire events in forests of western North America have assumed that the "stand age" variable from the US Forest Service Forest Inventory and Analysis (FIA) program reflects the timing of historical high-severity (i.e. stand-replacing) fire in ponderosa pine and mixed-conifer forests. To test this assumption, we re-analyze the dataset used in a previous analysis, and compare information from fire history records with information from co-located FIA plots. We demonstrate that 1) the FIA stand age variable does not reflect the large range of individual tree ages in the FIA plots: older trees comprised more than 10% of pre-stand age basal area in 58% of plots analyzed and more than 30% of pre-stand age basal area in 32% of plots, and 2) recruitment events are not necessarily related to high-severity fire occurrence. Because the FIA stand age variable is estimated from a sample of tree ages within the tree size class containing a plurality of canopy trees in the plot, it does not necessarily include the oldest trees, especially in uneven-aged stands. Thus, the FIA stand age variable does not indicate whether the trees in the predominant size class established in response to severe fire, or established during the absence of fire. FIA stand age was not designed to measure the time since a stand-replacing disturbance. Quantification of historical "mixed-severity" fire regimes must be explicit about the spatial scale of high-severity fire effects, which is not possible using FIA stand age data.

Bark water uptake promotes localized hydraulic recovery in coastal redwood crown.

Coastal redwood (Sequoia sempervirens), the world's tallest tree species, rehydrates leaves via foliar water uptake during fog/rain events. Here we examine if bark also permits water uptake in redwood branches, exploring potential flow mechanisms and biological significance. Using isotopic labelling and microCT imaging, we observed that water entered the xylem via bark and reduced tracheid embolization. Moreover, prolonged bark wetting (16 h) partially restored xylem hydraulic conductivity in isolated branch segments and whole branches. Partial hydraulic recovery coincided with an increase in branch water potential from about -5.5 ± 0.4 to -4.2 ± 0.3 MPa, suggesting localized recovery and possibly hydraulic isolation. As bark water uptake rate correlated with xylem osmotic potential (R(2) = 0.88), we suspect a symplastic role in transferring water from bark to xylem. Using historical weather data from typical redwood habitat, we estimated that bark and leaves are wet more than 1000 h per year on average, with over 30 events being sufficiently long (>24 h) to allow for bark-assisted hydraulic recovery. The capacity to uptake biologically meaningful volumes of water via bark and leaves for localized hydraulic recovery throughout the crown during rain/fog events might be physiologically advantageous, allowing for relatively constant transpiration.

A novel lung explant model for the ex vivo study of efficacy and mechanisms of anti-influenza drugs.

Influenza A virus causes considerable morbidity and mortality largely because of a lack of effective antiviral drugs. Viral neuraminidase inhibitors, which inhibit viral release from the infected cell, are currently the only approved drugs for influenza, but have recently been shown to be less effective than previously thought. Growing resistance to therapies that target viral proteins has led to increased urgency in the search for novel anti-influenza compounds. However, discovery and development of new drugs have been restricted because of differences in susceptibility to influenza between animal models and humans and a lack of translation between cell culture and in vivo measures of efficacy. To circumvent these limitations, we developed an experimental approach based on ex vivo infection of human bronchial tissue explants and optimized a method of flow cytometric analysis to directly quantify infection rates in bronchial epithelial tissues. This allowed testing of the effectiveness of TVB024, a vATPase inhibitor that inhibits viral replication rather than virus release, and to compare efficacy with the current frontline neuraminidase inhibitor, oseltamivir. The study showed that the vATPase inhibitor completely abrogated epithelial cell infection, virus shedding, and the associated induction of proinflammatory mediators, whereas oseltamivir was only partially effective at reducing these mediators and ineffective against innate responses. We propose, therefore, that this explant model could be used to predict the efficacy of novel anti-influenza compounds targeting diverse stages of the viral replication cycle, thereby complementing animal models and facilitating progression of new drugs into clinical trials.

Wildfire and drought dynamics destabilize carbon stores of fire-suppressed forests.

Widespread fire suppression and thinning have altered the structure and composition of many forests in the western United States, making them more susceptible to the synergy of large-scale drought and fire events. We examine how these changes affect carbon storage and stability compared to historic fire-adapted conditions. We modeled carbon dynamics under possible drought and fire conditions over a 300-year simulation period in two mixed-conifer conditions common in the western United States: (1) pine-dominated with an active fire regime and (2) fir-dominated, fire suppressed forests. Fir-dominated stands, with higher live- and dead-wood density, had much lower carbon stability as drought and fire frequency increased compared to pine-dominated forest. Carbon instability resulted from species (i.e., fir's greater susceptibility to drought and fire) and stand (i.e., high density of smaller trees) conditions that develop in the absence of active management. Our modeling suggests restoring historic species composition and active fire regimes can significantly increase carbon stability in fire-suppressed, mixed-conifer forests. Long-term management of forest carbon should consider the relative resilience of stand structure and composition to possible increases in disturbance frequency and intensity under changing climate.

Contrasting spatial patterns in active-fire and fire-suppressed Mediterranean climate old-growth mixed conifer forests.

In Mediterranean environments in western North America, historic fire regimes in frequent-fire conifer forests are highly variable both temporally and spatially. This complexity influenced forest structure and spatial patterns, but some of this diversity has been lost due to anthropogenic disruption of ecosystem processes, including fire. Information from reference forest sites can help management efforts to restore forests conditions that may be more resilient to future changes in disturbance regimes and climate. In this study, we characterize tree spatial patterns using four-ha stem maps from four old-growth, Jeffrey pine-mixed conifer forests, two with active-fire regimes in northwestern Mexico and two that experienced fire exclusion in the southern Sierra Nevada. Most of the trees were in patches, averaging six to 11 trees per patch at 0.007 to 0.014 ha(-1), and occupied 27-46% of the study areas. Average canopy gap sizes (0.04 ha) covering 11-20% of the area were not significantly different among sites. The putative main effects of fire exclusion were higher densities of single trees in smaller size classes, larger proportion of trees (≥ 56%) in large patches (≥ 10 trees), and decreases in spatial complexity. While a homogenization of forest structure has been a typical result from fire exclusion, some similarities in patch, single tree, and gap attributes were maintained at these sites. These within-stand descriptions provide spatially relevant benchmarks from which to manage for structural heterogeneity in frequent-fire forest types.