Genetic architecture of disease resistance and tolerance in Douglas-fir trees.

Understanding the genetic basis of how plants defend against pathogens is important to monitor and maintain resilient tree populations. Swiss needle cast (SNC) and Rhabdocline needle cast (RNC) epidemics are responsible for major damage of forest ecosystems in North America. Here we investigate the genetic architecture of tolerance and resistance to needle cast diseases in Douglas-fir (Pseudotsuga menziesii) caused by two fungal pathogens: SNC caused by Nothophaeocryptopus gaeumannii, and RNC caused by Rhabdocline pseudotsugae. We performed case-control genome-wide association analyses and found disease resistance and tolerance in Douglas-fir to be polygenic and under strong selection. We show that stomatal regulation as well as ethylene and jasmonic acid pathways are important for resisting SNC infection, and secondary metabolite pathways play a role in tolerating SNC once the plant is infected. We identify a major transcriptional regulator of plant defense, ERF1, as the top candidate for RNC resistance. Our findings shed light on the highly polygenic architectures underlying fungal disease resistance and tolerance and have important implications for forestry and conservation as the climate changes.

Phylogenetic analyses allow species-level recognition of Leptographium wageneri varieties that cause black stain root disease of conifers in western North America.

Leptographium wageneri is a native fungal pathogen in western North America that causes black stain root disease (BSRD) of conifers. Three host-specialized varieties of this pathogen were previously described: L. wageneri var. wageneri on pinyon pines (Pinus monophylla and P. edulis); L. wageneri var. ponderosum, primarily on hard pines (e.g., P. ponderosa, P. jeffreyi); and L. wageneri var. pseudotsugae on Douglas-fir (Pseudotsuga menziesii). Morphological, physiological, and ecological differences among the three pathogen varieties have been previously determined; however, DNA-based characterization and analyses are needed to determine the genetic relationships among these varieties. The objective of this study was to use DNA sequences of 10 gene regions to assess phylogenetic relationships among L. wageneri isolates collected from different hosts. The multigene phylogenetic analyses, based on maximum likelihood and Bayesian inference, strongly supported species-level separation of the three L. wageneri varieties. These results, in conjunction with previously established phenotypic differences, support the elevation of L. wageneri var. ponderosum and L. wageneri var. pseudotsugae to the species level as L. ponderosum comb. nov. and L. pseudotsugae comb. nov., respectively, while maintaining L. wageneri var. wageneri as Leptographium wageneri. Characterization of the three Leptographium species, each with distinct host ranges, provides a baseline to further understand the ecological interactions and evolutionary relationships of these forest pathogens, which informs management of black stain root disease.

Distribution of a Foliage Disease Fungus Within Canopies of Mature Douglas-Fir in Western Oregon.

Nothophaeocryptopus gaeumannii is a common native, endophytic fungus of Douglas-fir foliage, which causes Swiss needle cast, an important foliage disease that is considered a threat to Douglas-fir plantations in Oregon. Disease expression is influenced by fungal fruiting bodies (pseudothecia), which plug the stomata and inhibit gas exchange. Trees are impacted when pseudothecia plug stomates on 1-year-old and older needles resulting in early needle abscission. Mature (100 years+) trees appear to be less impacted from disease, and we hypothesize this is due to the greater emergence of pseudothecia on older than younger needles, which allows for more needle retention. We measured the density of pseudothecia occluding stomates across 2- to 5-year-old needles from upper, middle, and lower canopy positions of mature trees at three sites in the Oregon Coast Range and two sites in the western Oregon Cascade Mountains. Binomial generalized linear mixed model (GLMM) was used to test for the effects of canopy position (upper, middle, and lower), sites, needle age (2-5 years old), and years (2016 and 2017), and their interactions on the pseudothecia density. Pseudothecia density varied annually depending on sites, needle age and canopy positions. Pseudothecia density peaked on 3-, and 4-year-old needles, however, needles emerging from the same year, like 2-year-old needles in 2016 and 3-year-old needles in 2017 both emerged in 2014, had consistently similar patterns of pseudothecia density for both years, across site and canopy positions. Canopy position was important for 3-, and 4-year-old needles, showing less pseudothecia in the lower canopy. This research confirms that N. gaeumannii pseudothecia density is greatest in 3- and 4-year old needles in mature trees in contrast to plantations where pseudothecia density usually peaks on 2-year-old needles, and that pseudothecia density (disease severity) is generally lower in mature trees. Something about mature forest canopies and foliage appears to increase the time it takes for pseudothecia to emerge from the needles, in contrast to younger plantations, thus allowing the mature trees to have greater needle retention.

Tree species diversity increases with conspecific negative density dependence across an elevation gradient.

Elevational and latitudinal gradients in species diversity may be mediated by biotic interactions that cause density-dependent effects of conspecifics on survival or growth to differ from effects of heterospecifics (i.e. conspecific density dependence), but limited evidence exists to support this. We tested the hypothesis that conspecific density dependence varies with elevation using over 40 years of data on tree survival and growth from 23 old-growth temperate forest stands across a 1,000-m elevation gradient. We found that conspecific-density-dependent effects on survival of small-to-intermediate-sized focal trees were negative in lower elevation, higher diversity forest stands typically characterised by warmer temperatures and greater relative humidity. Conspecific-density-dependent effects on survival were less negative in higher elevation stands and ridges than in lower elevation stands and valley bottoms for small-to-intermediate-sized trees, but were neutral for larger trees across elevations. Conspecific-density-dependent effects on growth were negative across all tree size classes and elevations. These findings reveal fundamental differences in biotic interactions that may contribute to relationships between species diversity, elevation and climate.

The complexity of biological disturbance agents, fuels heterogeneity, and fire in coniferous forests of the western United States.

Forest biological disturbance agents (BDAs) are insects, pathogens, and parasitic plants that affect tree decline, mortality, and forest ecosystems processes. BDAs are commonly thought to increase the likelihood and severity of fire by converting live standing trees to more flammable, dead and downed fuel. However, recent research indicates that BDAs do not necessarily increase, and can reduce, the likelihood or severity of fire. This has led to confusion regarding the role of BDAs in influencing fuels and fire in fire-prone western United States forests. Here, we review the existing literature on BDAs and their effects on fuels and fire in the western US and develop a conceptual framework to better understand the complex relationships between BDAs, fuels and fire. We ask: 1) What are the major BDA groups in western US forests that affect fuels? and 2) How do BDA-affected fuels influence fire risk and outcomes? The conceptual framework is rooted in the spatiotemporal aspects of BDA life histories, which drive forest impacts, fuel characteristics and if ignited, fire outcomes. Life histories vary among BDAs from episodic, landscape-scale outbreaks (bark beetles, defoliators), to chronic, localized disturbance effects (dwarf mistletoes, root rots). Generally, BDAs convert aboveground live biomass to dead biomass, decreasing canopy fuels and increasing surface fuels. However, the rate of conversion varies with time-since-event and among BDAs and forest types, resulting in a wide range of effects on the amount of dead fuels at any given time and place, which interacts with the structure and composition of the stand before and subsequent to BDA events. A major influence on fuels may be that BDAs have emerged as dominant agents of forest heterogeneity creation. Because BDAs play complex roles in fuels and fire heterogeneity across the western US which are further complicated by interactions with climate change, drought, and forest management (fire suppression), their impacts on fuels, fire and ecological consequences cannot be categorized simply as positive or negative but need to be evaluated within the context of BDA life histories and ecosystem dynamics.

An Ecological Perspective on Living with Fire in Ponderosa Pine Forests of Oregon and Washington: Resistance, Gone but not Forgotten.

Wildland fires (WLF) have become more frequent, larger, and severe with greater impacts to society and ecosystems and dramatic increases in firefighting costs. Forests throughout the range of ponderosa pine in Oregon and Washington are jeopardized by the interaction of anomalously dense forest structure, a warming and drying climate, and an expanding human population. These forests evolved with frequent interacting disturbances including low-severity surface fires, droughts, and biological disturbance agents (BDAs). Chronic low-severity disturbances were, and still are, critical to maintaining disturbance resistance, the property of an ecosystem to withstand disturbance while maintaining its structure and ecological function. Restoration of that historical resistance offers multiple social and ecological benefits. Moving forward, we need a shared understanding of the ecology of ponderosa pine forests to appreciate how restoring resistance can reduce the impacts of disturbances. Given contemporary forest conditions, a warming climate, and growing human populations, we predict continued elevation of tree mortality from drought, BDAs, and the large high-severity WLFs that threaten lives and property as well as ecosystem functions and services. We recommend more comprehensive planning to promote greater use of prescribed fire and management of reported fires for ecological benefits, plus increased responsibility and preparedness of local agencies, communities and individual homeowners for WLF and smoke events. Ultimately, by more effectively preparing for fire in the wildland urban interface, and by increasing the resistance of ponderosa pine forests, we can greatly enhance our ability to live with fire and other disturbances.

Persistence of the Swiss Needle Cast Outbreak in Oregon Coastal Douglas-Fir and New Insights from Research and Monitoring.

Swiss needle cast (SNC), caused by Nothophaeocryptopus gaeumannii, is a foliage disease of Douglas-fir (Pseudotsuga menziesii), that reduces growth in native stands and exotic plantations worldwide. An outbreak of SNC began in coastal Oregon in the mid-1990s and has persisted since that time. Here we review the current state of knowledge after 24 years of research and monitoring, with a focus on Oregon, although the disease is significant in coastal Washington and has recently emerged in southwestern British Columbia. We present new insights into SNC distribution, landscape patterns, disease epidemiology and ecology, host-pathogen interactions, trophic and hydrologic influences, and the challenges of Douglas-fir plantation management in the presence of the disease. In Oregon, the SNC outbreak has remained geographically contained but has intensified. Finally, we consider the implications of climate change and other recently emerged foliage diseases on the future of Douglas-fir plantation management.

Tree growth declines and mortality were associated with a parasitic plant during warm and dry climatic conditions in a temperate coniferous forest ecosystem.

Insects and pathogens are widely recognized as contributing to increased tree vulnerability to the projected future increasing frequency of hot and dry conditions, but the role of parasitic plants is poorly understood even though they are common throughout temperate coniferous forests in the western United States. We investigated the influence of western hemlock dwarf mistletoe (Arceuthobium tsugense) on large (≥45.7 cm diameter) western hemlock (Tsuga heterophylla) growth and mortality in a 500 year old coniferous forest at the Wind River Experimental Forest, Washington State, United States. We used five repeated measurements from a long-term tree record for 1,395 T. heterophylla individuals. Data were collected across a time gradient (1991-2014) capturing temperature increases and precipitation decreases. The dwarf mistletoe rating (DMR), a measure of infection intensity, varied among individuals. Our results indicated that warmer and drier conditions amplified dwarf mistletoe effects on T. heterophylla tree growth and mortality. We found that heavy infection (i.e., high DMR) resulted in reduced growth during all four measurement intervals, but during warm and dry intervals (a) growth declined across the entire population regardless of DMR level, and (b) both moderate and heavy infections resulted in greater growth declines compared to light infection levels. Mortality rates increased from cooler-wetter to warmer-drier measurement intervals, in part reflecting increasing mortality with decreasing tree growth. Mortality rates were positively related to DMR, but only during the warm and dry measurement intervals. These results imply that parasitic plants like dwarf mistletoe can amplify the impact of climatic stressors of trees, contributing to the vulnerability of forest landscapes to climate-induced productivity losses and mortality events.

Increased streamflow in catchments affected by a forest disease epidemic.

Natural disturbances help maintain healthy forested and aquatic ecosystems. However, biotic and abiotic disturbance regimes are changing rapidly. For example, the Swiss needle cast (SNC) epidemic in the Coast Range of Oregon in the U.S. Pacific Northwest has increased in area from 53,050 to 238,705ha over the 1996-2015 period. We investigated whether the hydrologic regime (i.e., annual streamflow, runoff ratio, and magnitude and timing of peak flows and low flows) was affected by SNC in 12 catchments in western Oregon. The catchments ranged in size from 183 to 1834km2 and area affected by SNC from 0 to 90.5%. To maximize the number of catchments included in the study, we analyzed 20years of SNC aerial survey data and 15-26years of stream discharge (Q) and PRISM precipitation (P) and air temperature (Tair) data to test for trends in hydrologic variables for each catchment. As expected, we found that runoff ratios (Q/P) increased in five catchments, all with an area impacted by SNC >10%. This was likely due to the effects of SNC on the hydraulic architecture (i.e., needle retention, sapwood area, sapwood permeability) of affected trees, leading to decreased canopy interception and transpiration losses. Interestingly, two catchments with the greatest area affected by SNC showed no changes in hydrologic regime. The lack of hydrologic response could either be due to compensatory transpiration by vegetation unaffected by the disease or sub-canopy abiotic evaporation, which counteracted reductions in transpiration. This study is the first to illustrate that chronic canopy disturbance from a foliage pathogen can influence catchment scale hydrology.

Severity of Swiss needle cast in young and mature Douglas-fir forests in western Oregon, USA.

Swiss needle cast (SNC), caused by Nothophaeocryptopus gaeumannii, is an important foliage disease of Douglas-fir (Pseudotsuga menziesii) forests of the Pacific Northwest. The fungus lives endophytically within the foliage, until forming reproductive structures (pseudothecia) that plug stomates and cause carbon starvation. When pseudothecia appear on one- and two-year-old foliage, significant needle abscission can occur, which reduces productivity of the tree. While there is considerable evidence of SNC disease in coastal Douglas-fir plantations, the severity of SNC in mature and old-growth forests is poorly understood. We compared tree crowns of mature and old-growth conifer forests and nearby young forests at three locations in the Oregon Coast Range and four locations in the western Cascade Range of Oregon. We assessed disease severity for N. gaeumannii on two-year-old foliage, incidence by presence of N. gaeumannii on all foliage, foliage retention for the first four years, and foliar nitrogen of one-year-old foliage. We also compared leaf wetness at three heights in one mature and one young tree at five of the seven sites. Disease severity was greater in young forests than mature forests at all sites except for high elevation Cascade Range areas. Incidence of disease was highest for two-year-old needles in young trees and 3-5 year-old needles in mature trees, except for one coastal site. Retention of 1-4 year-old needle cohorts differed between young and mature trees, and mature trees had much larger complements of > four-year-old needles. Total foliar nitrogen (TN) concentration did not differ in needles of young and mature trees, but at some locations total N differed between canopy positions. Leaf wetness differences were not consistent between young and mature tree crowns. However, at one study site in the core epidemic area, the younger stand had longer periods of wetness in the upper crowns than a nearby old stand. Leaf wetness and foliar N were hypothesized to play a role in SNC disease severity, but they do not explain differences in adjacent young and mature trees. Although the fungus is present in old and young trees, the likelihood of disease expression and lower foliage retention appears to be greater in younger plantation trees than mature and older trees in western Oregon Douglas-fir forests.

Interactions of predominant insects and diseases with climate change in Douglas-fir forests of western Oregon and Washington, U.S.A.

Forest disturbance regimes are beginning to show evidence of climate-mediated changes, such as increasing severity of droughts and insect outbreaks. We review the major insects and pathogens affecting the disturbance regime for coastal Douglas-fir forests in western Oregon and Washington State, USA, and ask how future climate changes may influence their role in disturbance ecology. Although the physiological constraints of light, temperature, and moisture largely control tree growth, episodic and chronic disturbances interacting with biological factors have substantial impacts on the structure and functioning of forest ecosystems in this region. Understanding insect and disease interactions is critical to predicting forest response to climate change and the consequences for ecosystem services, such as timber, clean water, fish and wildlife. We focused on future predictions for warmer wetter winters, hotter drier summers, and elevated atmospheric CO2 to hypothesize the response of Douglas-fir forests to the major insects and diseases influencing this forest type: Douglas-fir beetle, Swiss needle cast, black stain root disease, and laminated root rot. We hypothesize that 1) Douglas-fir beetle and black stain root disease could become more prevalent with increasing, fire, temperature stress, and moisture stress, 2) future impacts of Swiss needle cast are difficult to predict due to uncertainties in May-July leaf wetness, but warmer winters could contribute to intensification at higher elevations, and 3) laminated root rot will be influenced primarily by forest management, rather than climatic change. Furthermore, these biotic disturbance agents interact in complex ways that are poorly understood. Consequently, to inform management decisions, insect and disease influences on disturbance regimes must be characterized specifically by forest type and region in order to accurately capture these interactions in light of future climate-mediated changes.

Climate Risk Modelling of Balsam Woolly Adelgid Damage Severity in Subalpine Fir Stands of Western North America.

The balsam woolly adelgid (Adelges piceae (Ratzeburg) (Homoptera: Adelgidae)) (BWA) is a nonnative, invasive insect that threatens Abies species throughout North America. It is well established in the Pacific Northwest, but continues to move eastward through Idaho and into Montana and potentially threatens subalpine fir to the south in the central and southern Rocky Mountains. We developed a climatic risk model and map that predicts BWA impacts to subalpine fir using a two-step process. Using 30-year monthly climate normals from sites with quantitatively derived BWA damage severity index values, we built a regression model that significantly explained insect damage. The sites were grouped into two distinct damage categories (high damage and mortality versus little or no mortality and low damage) and the model estimates for each group were used to designate distinct value ranges for four climatic risk categories: minimal, low, moderate, and high. We then calculated model estimates for each cell of a 4-kilometer resolution climate raster and mapped the risk categories over the entire range of subalpine fir in the western United States. The spatial variation of risk classes indicates a gradient of climatic susceptibility generally decreasing from the Olympic Peninsula in Washington and the Cascade Range in Oregon and Washington moving eastward, with the exception of some high risk areas in northern Idaho and western Montana. There is also a pattern of decreasing climatic susceptibility from north to south in the Rocky Mountains. Our study provides an initial step for modeling the relationship between climate and BWA damage severity across the range of subalpine fir. We showed that September minimum temperature and a metric calculated as the maximum May temperature divided by total May precipitation were the best climatic predictors of BWA severity. Although winter cold temperatures and summer heat have been shown to influence BWA impacts in other locations, these variables were not as predictive as spring and fall conditions in the Pacific Northwest.

Effects of dwarf mistletoe on stand structure of lodgepole pine forests 21-28 years post-mountain pine beetle epidemic in central Oregon.

Lodgepole pine (Pinus contorta) forests are widely distributed throughout North America and are subject to mountain pine beetle (Dendroctonus ponderosae) epidemics, which have caused mortality over millions of hectares of mature trees in recent decades. Mountain pine beetle is known to influence stand structure, and has the ability to impact many forest processes. Dwarf mistletoe (Arceuthobium americanum) also influences stand structure and occurs frequently in post-mountain pine beetle epidemic lodgepole pine forests. Few studies have incorporated both disturbances simultaneously although they co-occur frequently on the landscape. The aim of this study is to investigate the stand structure of lodgepole pine forests 21-28 years after a mountain pine beetle epidemic with varying levels of dwarf mistletoe infection in the Deschutes National Forest in central Oregon. We compared stand density, stand basal area, canopy volume, proportion of the stand in dominant/codominant, intermediate, and suppressed cohorts, average height and average diameter of each cohort, across the range of dwarf mistletoe ratings to address differences in stand structure. We found strong evidence of a decrease in canopy volume, suppressed cohort height, and dominant/codominant cohort diameter with increasing stand-level dwarf mistletoe rating. There was strong evidence that as dwarf mistletoe rating increases, proportion of the stand in the dominant/codominant cohort decreases while proportion of the stand in the suppressed cohort increases. Structural differences associated with variable dwarf mistletoe severity create heterogeneity in this forest type and may have a significant influence on stand productivity and the resistance and resilience of these stands to future biotic and abiotic disturbances. Our findings show that it is imperative to incorporate dwarf mistletoe when studying stand productivity and ecosystem recovery processes in lodgepole pine forests because of its potential to influence stand structure.

Impacts of dwarf mistletoe on the physiology of host Tsuga heterophylla trees as recorded in tree-ring C and O stable isotopes.

Dwarf mistletoes, obligate, parasitic plants with diminutive aerial shoots, have long-term effects on host tree water relations, hydraulic architecture and photosynthetic gas exchange and can eventually induce tree death. To investigate the long-term (1886-2010) impacts of dwarf mistletoe on the growth and gas exchange characteristics of host western hemlock, we compared the diameter growth and tree-ring cellulose stable carbon (C) and oxygen (O) isotope ratios (δ(13)Ccell, δ(18)Ocell) of heavily infected and uninfected trees. The relative basal area growth of infected trees was significantly greater than that of uninfected trees in 1886-90, but declined more rapidly in infected than uninfected trees through time and became significantly lower in infected than uninfected trees in 2006-10. Infected trees had significantly lower δ(13)Ccell and δ(18)Ocell than uninfected trees. Differences in δ(18)Ocell between infected and uninfected trees were unexpected given that stomatal conductance and environmental variables that were expected to influence the δ(18)O values of leaf water were similar for both groups. However, estimates of mesophyll conductance (gm) were significantly lower and estimates of effective path length for water movement (L) were significantly higher in leaves of infected trees, consistent with their lower values of δ(18)Ocell. This study reconstructs the long-term physiological responses of western hemlock to dwarf mistletoe infection. The long-term diameter growth and δ(13)Ccell trajectories suggested that infected trees were growing faster than uninfected trees prior to becoming infected and subsequently declined in growth and leaf-level photosynthetic capacity compared with uninfected trees as the dwarf mistletoe infection became severe. This study further points to limitations of the dual-isotope approach for identifying sources of variation in δ(13)Ccell and indicates that changes in leaf internal properties such as gm and L that affect δ(18)Ocell must be considered.

Seasonal carbohydrate dynamics and growth in Douglas-fir trees experiencing chronic, fungal-mediated reduction in functional leaf area.

Stored non-structural carbohydrates (NSCs) could play an important role in tree survival in the face of a changing climate and associated stress-related mortality. We explored the effects of the stomata-blocking and defoliating fungal disease called Swiss needle cast on Douglas-fir carbohydrate reserves and growth to evaluate the extent to which NSCs can be mobilized under natural conditions of low water stress and restricted carbon supply in relation to potential demands for growth. We analyzed the concentrations of starch, sucrose, glucose and fructose in foliage, twig wood and trunk sapwood of 15 co-occurring Douglas-fir trees expressing a gradient of Swiss needle cast symptom severity quantified as previous-year functional foliage mass. Growth (mean basal area increment, BAI) decreased by ∼80% and trunk NSC concentration decreased by 60% with decreasing functional foliage mass. The ratio of relative changes in NSC concentration and BAI, an index of the relative priority of storage versus growth, more than doubled with increasing disease severity. In contrast, twig and foliage NSC concentrations remained nearly constant with decreasing functional foliage mass. These results suggest that under disease-induced reductions in carbon supply, Douglas-fir trees retain NSCs (either actively or due to sequestration) at the expense of trunk radial growth. The crown retains the highest concentrations of NSC, presumably to maintain foliage growth and shoot extension in the spring, partially compensating for rapid foliage loss in the summer and fall.

Tree-ring stable isotopes record the impact of a foliar fungal pathogen on CO(2) assimilation and growth in Douglas-fir.

Swiss needle cast (SNC) is a fungal disease of Douglas-fir (Pseudotsuga menziesii) that has recently become prevalent in coastal areas of the Pacific Northwest. We used growth measurements and stable isotopes of carbon and oxygen in tree-rings of Douglas-fir and a non-susceptible reference species (western hemlock, Tsuga heterophylla) to evaluate their use as proxies for variation in past SNC infection, particularly in relation to potential explanatory climate factors. We sampled trees from an Oregon site where a fungicide trial took place from 1996 to 2000, which enabled the comparison of stable isotope values between trees with and without disease. Carbon stable isotope discrimination (Δ(13)C) of treated Douglas-fir tree-rings was greater than that of untreated Douglas-fir tree-rings during the fungicide treatment period. Both annual growth and tree-ring Δ(13)C increased with treatment such that treated Douglas-fir had values similar to co-occurring western hemlock during the treatment period. There was no difference in the tree-ring oxygen stable isotope ratio between treated and untreated Douglas-fir. Tree-ring Δ(13)C of diseased Douglas-fir was negatively correlated with relative humidity during the two previous summers, consistent with increased leaf colonization by SNC under high humidity conditions that leads to greater disease severity in following years.

Ethanol attracts scolytid beetles to Phytophthora ramorum cankers on coast live oak.

Ethanol in sapwood was analyzed along vertical transects, through small spot cankers and larger basal cankers, of Phytophthora ramorum-infected stems of Quercus agrifolia at three sites in California. Trees with large basal cankers, known to attract scolytid beetles, had a 4.3 times higher ethanol level than trees with spot cankers that attract fewer beetles. Ethanol concentrations inside cankers, where scolytid beetles preferentially attack, varied by about four orders of magnitude among samples, with a median level of 16.0 µg.g(-1) fresh mass. This concentration was 4.3 and 15.5 times greater, respectively, than the concentrations at 1 cm or 15-30 cm outside the canker boundaries. In the laboratory, we demonstrated that ethanol escaped through the bark of a Q. garryana log just 3 days after it was added to the sapwood. At the three study sites, traps baited with ethanol captured more Xyleborinus saxesenii, Pseudopityophthorus pubipennis, and Monarthrum dentiger (all Coleoptera: Curculionidae: Scolytinae) than traps baited with ethanol plus (-)-α-pinene, or ethanol plus 4-allylanisole (4AA). Logs of Q. agrifolia with a 50 % ethanol solution added to the sapwood were placed at the study sites, with or without additional bark treatments above the ethanol. The number of scolytid beetle gallery holes above the ethanol-infused sapwood was 4.4 times greater than that on the opposite side of the log where no ethanol was added. Attachment of ultra-high release (-)-α-pinene pouches to the bark surface above the 50 % ethanol solution reduced scolytid attacks to a density of 19.1 % that of logs without this treatment. We conclude that ethanol in P. ramorum cankers functions as a primary host attractant for scolytid beetles and is an important link in colonization of these cankers and accelerated mortality of Q. agrifolia. The results of this research shed light on the chemical ecology behind the focused scolytid attacks on P. ramorum-infected coast live oaks, and lay the groundwork for future efforts to prolong the survival of individual trees of this keystone species.