Search

Recent increases in Douglas-fir (Psuedotsuga menziesii var. menziesii) mortality in the Klamath Mountains ecoregion raise concerns about the long-term resilience of Douglas-fir in the ecoregion and increased potential for uncharacteristic wildfire. We used data from the USDA Forest Service Aerial Detection Survey and ninety-six field plots to explore the relationships between physiographic and climate variables and Douglas-fir mortality. Our results provide strong evidence for a decline spiral in which Douglas-fir growing on hot, dry sites (predisposing factor) are further stressed by drought (inciting factor) and are then exploited by the flatheaded fir borer (Phaenops drummondi) and other secondary biotic agents (contributing factors), resulting in decline and mortality. At the landscape scale, Douglas-fir mortality increased as average annual precipitation declined and average climatic water deficit increased. We developed a risk score integrating several environmental variables associated with drought and heat stress to predict the likelihood and intensity of mortality at the stand scale.

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.

Swiss needle cast (SNC) is an important foliage disease of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) caused by the native pathogen Nothophaeocryptopus gaeumannii (T. Rohde) Videira, C. Nakash., U. Braun & Crous, which has been present in epidemic proportions in coastal Douglas-fir forests since the 1990s. Under conducive environmental and stand conditions, the fungal fruiting bodies emerge on young needles, inhibiting gas exchange and causing premature needle casting and subsequent growth losses. Using a new regional plot network, which extends and approximately doubles the area of SNC-susceptible coastal forest sampled, we investigated the distribution of SNC disease indices across the region and throughout individual tree crowns. Foliage retention varied from 1.15 to 3.9 years, and disease severity index (incidence × % occluded stomata) ranged from 0.05% to 52.11%. Foliage retention was positively correlated with distance from the coast and elevation, while foliage retention and disease severity were negatively associated across the study area. Within crowns, disease severity was negatively associated with crown depth, and foliage retention was positively associated with crown depth, regardless of distance from coast. Across the entire study area, foliage retention was found to decrease and disease severity increase with latitude, all else being equal. Tree growth metrics are positively associated with increasing foliage retention, and normal growth occurs greater than ∼3.2 years.

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.

The horticulturist Luther Burbank introduced the European mistletoe (Viscum album L.) to Sebastopol, Sonoma County, California, USA, around 1900 to grow as a Christmas ornament crop and tincture for medicinal use. The mistletoe has since spread from the point of introduction on apple to other hardwood trees, especially non-native hardwoods in yards and farms of the region. Mistletoe surveys were previously conducted in 1971, 1986, and 1991. We re-surveyed the region in 2019, with emphasis on the 1991 perimeter, and documented the current farthest distribution of V. album. This represents a 120-year record of spread. We observed infected trees up to 24.6 km (15.3 miles) from the point of introduction, doubling the farthest distance reported in 1991. The estimated area encompassed by mistletoe-infected hosts increased from 184 km2 (71 miles2) in 1991 to 606 km2 (234 miles2) in 2019. We also updated the host species list including both native and non-native mistletoe hosts. Viscum album spread appears to be limited by available habitat and hosts, but within the survey area it is intensifying, concentrated in urban and semi-urban yards, streets, and farms. However, V. album was also commonly found in riparian areas, which have a large number of native hosts.

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.

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.

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.

Swiss needle cast (SNC) is a foliage disease of Douglas-fir (Pseudotsuga menziesii) caused by Nothophaeocryptopus gaeumannii, an ascomycete fungus (Mycosphaerellaceae) that causes tree growth reductions in the Pacific Northwest. The epidemiology of the fungus is generally well known, but the relations between disease expression and foliar nutrition are unclear. In this study, we used data from the Swiss Needle Cast Cooperative research and monitoring plot network in western Oregon and SW Washington to assess associations between SNC severity, carbon, and nine foliage nutrients (nitrogen, Na, K, P, Ca, Mg, Mn, Al, S). Foliage samples were collected from midcrown of selected Douglas-firs from each plot. SNC severity was determined on 2-year-old needles by multiplying disease incidence and fungal reproductive (pseudothecia) density. Disease severity and nutrient relations were determined using linear mixed models. SNC severity showed statistically significant positive trends with concentrations of carbon, nitrogen, Na, K, and S, no relation with concentrations of Ca, Mg, or Al, and slightly negative trends that were not significant for P and Mn. This is the first such analysis of associations between a conifer foliage disease and foliage nutrients across a landscape; subsequently, there is little published literature on how or why these nutrients may interact with disease.

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.

Swiss needle cast (SNC) is a fungal disease of Douglas-fir (Pseudotsuga menziesii) that is having important consequences on tree growth in the Pacific Northwest (PNW) region of the USA. Once considered innocuous in PNW forests, SNC symptom expression has increased rapidly in extent and intensity in recent decades. Previous research has linked the disease epidemiology of SNC to climate, and observations indicate a link with forestry practices of the 20th century as well, particularly with the conversion of old growth and mature mixed-conifer forests to young monocultures of Douglas-fir on private forestlands. Given the sensitivity of SNC intensification to forestry practices and changing climatic conditions, it seems plausible that disease behavior response to short-term directional climate changes could differ between contrasting forest management regimes. We compared the relationship between trends in canopy energy and water flux parameters detected during the spring and summer months (May–August) along the Pacific Coast of Oregon from 2003 to 2012, and the distribution of SNC symptoms in 2012 on private and public lands. Canopy energy and water exchange parameters were calculated with MODIS Land Surface Temperature (LST), and evapotranspiration (ET) data, and with Parameter-elevation Relationships on Independent Slopes Model (PRISM) precipitation data. We found that a higher level of deviance in SNC presence/absence could be explained on private land compared with public land. Proximity to coast explained 9.3% of the deviance on private land and 6.7% on public land. Trends in LST during May and August emerged as important and explained 7% of the deviance in SNC symptom distribution on private land compared with 2% on public land. When combined with proximity to coast, May and August LST explained 14% of the deviance in SNC symptom expression on private land and 8.7% on public land. We found a significant difference between public and private ownership for the proportion of SNC (p = 0.0006), and a significant interaction between ownership and distance to coast (p = 0.0019), such that across public and private ownership, distance to coast has a different effect. LST may provide valuable information on leaf wetness, or thermal properties of the canopy, possibly capturing both early season and late season dynamics important to SNC epidemiology. We find evidence that recent short-term directional climate changes may have contributed to differences in symptom development in Douglas-fir forests on private and public land, with symptoms more prevalent on private land.

In the northwest, White-headed Woodpeckers (Picoides albolarvatus) are an uncommon species restricted to interior, dry coniferous forests. This species forages primarily by probing in bark crevices, surface gleaning, and excavating below the bark. Here we report on an observation from Mt. Ashland in southern Oregon of a White-headed Woodpecker feeding on insects in and around the sporophores of Veiled Polypore (Cryptoporus volvatus). Inspection of other sporophores with signs of woodpecker feeding revealed abundant fly larvae and beetles (adults and larvae). This widespread decay fungus colonizes the sapwood of recently dead trees where it is frequently associated with bark beetles. A suite of other insects is attracted to the sporophore, many using the enclosed pouch as a pupation chamber. In addition to exploring the significance of C. volvatus as a foraging substrate for White-headed Woodpeckers, we discuss the potential role of White-headed Woodpeckers as dispersal agents for C. volvatus and the possible interplay between C. volvatus-induced decay, insect availability, and habitat selection by White-headed Woodpeckers.

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.

Dwarf mistletoes (Viscaceae: Arceuthobium spp.) and fire interact in important ways in the coniferous forests of western North America. Fire directly affects dwarf mistletoes by killing the host, host branch, or heating/smoking the aerial shoots and fruits. Fire is a primary determinant of dwarf mistletoe distribution on the landscape, and time since fire controls many aspects of dwarf mistletoe epidemiology. Conversely, dwarf mistletoes can influence fire by causing changes in forest composition, structure, and fuels. Prescribed fire is important for management of dwarf mistletoes, while fire suppression is thought to have increased dwarf mistletoe abundance in western forests. Two dwarf mistletoes are compared in order to illustrate fire interactions in Oregon and Washington, USA: Arceuthobium americanum Nutt. ex Engelm. (lodgepole pine dwarf mistletoe) and Arceuthobium tsugense (Rosendahl)(western hemlock dwarf mistletoe). Arceuthobium persists on the landscape where the host is not killed by fire. Arceuthobium americanum spreads directly into the regenerating Pinus contorta Dougl. ex Loud., while A. tsugense persists in refugia for 200 years or more following fire. Host successional status is a driver of fire – dwarf mistletoe interactions, but forest disturbance agents also play a role. Given the importance of these interactions to the ecology of fire-prone forests, dwarf mistletoes warrant inclusion in disturbance ecology research.

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.

Mountain pine beetle (Dendroctonus ponderosae Hopkins; MPB), a bark beetle native to western North America, has caused vast areas of tree mortality over the last several decades. The majority of this mortality has been in lodgepole pine (Pinus contorta Douglas ex Loudon) forests and has heightened concerns over the potential for extreme fire behavior across large landscapes. Although considerable research has emerged concerning influence of MPB on forest fuels, there has been little work in the climax lodgepole pine forests of south-central Oregon, USA. Specifically, we assessed changes in forest structure and crown and surface fuels across a chronosequence of time since mountain pine beetle (TSB) epidemics in south-central Oregon (1979 to 2008). We classified four distinct periods in which significant changes occur in fuels: overstory mortality stage (2 to 4 years TSB), standing snag and snag fall stage (5 to 13 years TSB), regeneration stage (14 to 25 years TSB), and overstory recovery stage (26 to 32 years TSB). Multivariate analyses indicated changes in crown fuels and forest structure following MPB epidemics were driven primarily by basal area of live and standing dead trees, canopy bulk density, canopy base height, and canopy height. Substantial declines in canopy bulk density occurred early (2 to 5 yrs) following beetle activity and slowly recovered over time. The pattern of succession of surface fuels following a MPB epidemic was largely determined by changes over time in 10-h, 100-h, and 1000-h fuel loads, in addition to increasing fuel bed depth. The 100-h fuel load increased over the entire 30-year period, while 1000-h fuel load reached an asymptote 14 to 26 years following epidemic initiation. Live woody fuels increased through the initial overstory mortality stage and began to decrease during the overstory recovery stage. Our key findings concerning changing fuels and forest structure following a MPB epidemic in south-central Oregon lodgepole pine forests include: 1-h fuels and litter changed little over time, surface fuel loads changed dramatically between the standing snag and the regeneration stages, lodgepole pine remained dominant, and canopy bulk density was low throughout the chronosequence. These factors point to the perpetuation of a lodgepole pine dominated system with a mixed-severity fire regime well into the future.

Phytophthora ramorum, cause of “sudden oak death” or SOD, has had significant impacts on composition and structure in coastal forests of central and northern coastal California and southwestern Oregon. Despite the proximity of susceptible coast live oak (Quercus agrifolia) forests to densely populated urban areas, the impacts of SOD on their fuels have not been studied. We sampled surface fuels and vegetation structure in 16 plots in both SOD-caused gaps and intact stands (32 plots total) across two parks in the East Bay Regional Park District, east of San Francisco Bay. Plots were selected from a set of randomly placed pre-existing locations used in determining the disease distribution and intensity across the park system. Among the vegetation characteristics examined, only coast live oak basal area and live and dead tree density, canopy cover and maximum forb height differed between SOD created gap plots and adjacent intact forest plots. However, surface fuels such as vegetation cover, litter cover, wood cover, duff depth, fuels height, ladder fuels abundance, 1 h, 10 h, 100 h, and 1000 h fuels were greater in gap plots than intact forest plots. Although no fire behavior models were run, surface fuels suggest SOD created gaps may facilitate passive crown fire due to increased ladder and other fuels. This study represents a spatially explicit (gap focused) point-in-time estimate of surface fuels that will continue to change through time as disease progresses through these stands causing vegetation changes as fuels accumulate and decompose.

We investigated the forest health of red fir (Abies magnifica) and how it compared with commonly-associated species Jeffrey pine (Pinus jeffreyi), lodgepole pine (Pinus contorta) and white fir (Abies concolor) in the upper montane forests of California. We evaluated tree mortality rates, changes in the density of recently-dead trees and the amount of insect and disease damage on live trees from comprehensive forest inventories. The annual mortality rate for red fir was 1.8%, while the rates for Jeffrey pine, lodgepole pine and white fir were 1.9, 1.1 and 3.0%, respectively. The proportion of recently-dead red fir trees increased over time, suggesting an increase in mortality; however, stage-transition models suggested the current population structure of red fir is stable. Dwarf mistletoe and drought-stressed sites were significant predictors of red fir mortality. Trees with substantial damage had a higher probability of experiencing mortality in five years. Our results are consistent with others, but the timeframe is too short to make conclusions about long-term declines. Our results suggest that the most significant of multiple biotic factors involved in red fir mortality processes is dwarf mistletoe, while tree age and the proportion of forest in old-growth may also influence mortality rates.

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 cooccurring Douglas-fi r trees expressing a gradient of Swiss needle cast symptom severity quantified as previous-year func tional 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.

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.

We investigated the vertical pattern of foliage retention of Douglas-fir (Pseudotsuga menziesii) in the western Oregon Coast Range where Swiss needle cast, a foliage disease caused by Phaeocryptopus gaeumannii, is causing foliage loss and growth impacts. Swiss needle cast reduced foliage retention more in the upper crown than the lower crown within the epidemic area, which is unusual as foliage diseases usually reduce foliage retention most in the lower crown. We hypothesized that as foliage retention increased across environmental gradients that it would also increase in the upper crown at a greater rate than the lower crown. We randomly selected 72 sites from a population of Douglas-fir plantations in the northwest Oregon Coast Range. We estimated foliage retention from the lower, mid and upper crown of 10 trees per plot. We fitted a two-level hierarchical model with tree and stand level predictors to model changes in foliage retention with changing environmental gradient for foliage in each of three vertical crown positions. We found that the vertical pattern of foliage retention was generally similar throughout the study area with lowest retention in the upper crown, and highest retention in the lower crown. Foliage retention increased with increasing distance from the coast, which is correlated with increased elevation and decreased temperature and site productivity. These findings are consistent with our current understanding of conifer foliage retention. No apparent shift occurs from whole crown to lower crown impacts in our study area as foliage retention increases and approaches normal.

Swiss needle cast (SNC), an important fungal disease of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), has increased in severity throughout its natural and introduced range over the last half century. The role of climate change and forest management practices in the increase is unclear. We analyzed tree-ring chronologies from six late-successional Douglas-fir stands in the western Oregon Coast Range using time-series intervention analysis (TSIA) to address how climate relates to the impact of SNC on tree growth. Tree-ring chronologies of western hemlock (Tsuga heterophylla (Raf.) Sarg.), a species not susceptible to the fungus Phaeocryptopus gaeumannii (Rhode) Petrak, were used as a climate proxy in the TSIA. We found that growth reductions associated with SNC dated back to the 1590s, the earliest record in our dendritic data. Growth reductions were synchronous across the six sites, indicating that the disease severity was largely influenced by climatic conditions. SNC impact peaked in 1984–1986 at all six study sites, followed by unprecedented disease impacts of 100% in 1996 and 2004 at one site, while decreasing to previous levels at the other five sites. Our SNC index of impact significantly correlated with winter and summer temperatures and summer precipitation. Winter conditions were more strongly associated with disease impact at wetter, cooler sites, whereas summer conditions were more important at less humid, warmer sites. With climate change, SNC impacts are likely to increase in coastal areas where June–July precipitation is much higher than the P. gaeumannii-limiting threshold of ∼110 mm, and decrease where summer precipitation is at or below the threshold. Warmer winters will increase disease severity at higher elevation, north along the coast from northern Oregon to British Columbia, and at inland sites where current winter temperatures limit fungal growth.

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.

The influence of fertilization on disease severity is unknown in most forest pathosystems. Fertilization treatments were randomly applied to 0.01 ha plots centered on individual dominant or co-dominant Douglas-fir trees in ten Douglas-fir stands from coastal Oregon to the foothills of the Oregon Cascade Range, USA. This region is affected by Swiss needle cast, caused by the fungal pathogen Phaeocryptopus gaeumannii. Selected stands represented a range of Swiss needle cast disease severity, and 10 replications of each fertilization treatment were applied in each stand. The six treatments included nitrogen (urea), calcium as lime (calcium carbonate), calcium as calcium chloride, phosphorus (monosodium phosphate), a site-specific blend (Kinsey) and an unfertilized control. Fertilization took place from February–April 2007, and single branches were collected from treated trees for disease severity assessment of foliage in May 2010. Disease severity of 1- and 2-year-old needles was evaluated by counting the frequency of infected needles and the density of P. gaeumannii fruiting bodies (pseudothecia) on a random subset of needles from each tree and needle age class. Fertilization treatment effects on infection index (mean fruiting body density) were tested by mixed-effects models that accounted for site as a blocking factor. Treatment effects on infection index at each of the study locations were also tested by ten separate ANOVAs. Across and within sites, fertilization treatment did not significantly affect infection index of 1- or 2-year old needles (p > 0.05). Small differences in mean fruiting body density (⩽3%) between fertilization and control treatments across sites were not statistically significant, nor are they believed to be biologically or economically significant. Decisions regarding fertilization should be based on site-specific attributes, such as soil chemistry. There is no evidence that fertilization directly ameliorates or exacerbates Swiss needle cast severity in western Oregon.

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.

Climate change resulting from increased concentrations of atmospheric carbon dioxide ([CO2]) is expected to result in warmer temperatures and changed precipitation regimes during this century. In the northwestern U.S., these changes will likely decrease snowpack, cause earlier snowmelt, increase summer evapotranspiration, and increase the frequency and severity of droughts. Elevated [CO2] and warmer temperatures may have positive effects on growth and productivity where there is adequate moisture or growth is currently limited by cold. However, the effects of climate change are generally expected to reduce growth and survival, predispose forests to disturbance by wildfire, insects, and disease; and ultimately change forest structure and composition at the landscape scale. Substantial warming will likely decrease winter chilling resulting in delayed bud burst, and adversely affect flowering and seed germination for some species. The extent of these effects will depend on the magnitude of climate change, the abilities of individual trees to acclimate, and for tree populations to adapt in situ, or to migrate to suitable habitats. These coping mechanisms may be insufficient to maintain optimal fitness of tree populations to rapidly changing climate. Physiological responses to climatic stresses are relatively well-understood at the organ or whole-plant scale but not at the stand or landscape scale. In particular, the interactive effects of multiple stressors is not well known. Genetic and silvicultural approaches to increase adaptive capacities and to decrease climate-related vulnerabilities of forests can be based on ecophysiological knowledge. Effective approaches to climate adaptation will likely include assisted migration of species and populations, and density management. Use of these approaches to increase forest resistance and resilience at the landscape scale requires a better understanding of species adaptations, within-species

Tree-ring analysis was applied to assess the impacts of the fungal disease Swiss needle cast on the radial growth of mature Douglas-fir (Pseudotsuga menziesii) forests in the western Oregon Coast Range. Although considered endemic to the Pacific Northwest, Swiss needle cast has significantly lowered productivity in Douglas-fir forests only in the past 20–30 years. To date, studies on Swiss needle cast impacts have almost exclusively involved young ( 80 years) trees, we extracted tree cores from dominant and codominant Douglas-fir and western hemlock (Tsuga heterophylla) in three even-aged stands in western Oregon. In the least affected stand growth rates of both species did not significantly differ, while at the most severely diseased site Douglas-fir radial growth was reduced by as much as 85%. Growth reductions likely associated with Swiss needle cast were dated to as early as 1950, though the most severe impacts occurred after 1984. An index of Swiss needle cast severity significantly (p < 0.01) related to instrumental records of air temperatures such that warm conditions from March through August were associated with reduced radial growth at the most severely affected site. Overall, this study demonstrates that even mature forests of natural origin are susceptible to severe growth reductions by Swiss needle cast, that warmer spring and summer temperatures are associated with Swiss needle cast impacts, and that the disease appears to be increasing in severity.

Summary Our objective was to test ground penetrating radar (GPR) to non-destructively estimate decay volumes in living coniferous trees. GPR is geophysical tool which uses an antenna to propagate short bursts of electromagnetic energy in solid materials and measure the two-way travel time and amplitude of reflected signals. We compared estimates of bole decay from data collected with a SIR 3000 GPR system equipped with a 900 MHz antenna to measurements of decay from stem cross sections and increment cores for three conifer species (Pseudotsuga menziesii, Thuja plicata and Tsuga heterophylla). We found that near-surface decay, air-filled voids and desiccated boles had unique electromagnetic signatures, which could be separated from other defects. GPR successfully estimated the percent area of air-filled cavities and was not significantly different than results from destructive sampling. However, separation of incipient to severe decay from benign reflectors (e.g. moisture gradient between sapwood and heartwood) in conifers was much less diagnostic than with angiosperms. A limited assessment of Acer saccharum showed that GPR has potential to detect defects in angiosperms; however, more research is needed to outline the full range of detectable defects. Based on the trees in this study, the potential for GPR to detect decay-related defects in conifers seems limited. Despite problems detecting decay, reflections originating from the sapwood : heartwood boundary may prove useful to determine thickness of functional sapwood in conifers, but accurate quantification will require further technical development.

We investigated the effect of western hemlock dwarf mistletoe ( Arceuthobium tsugense (Rosend.) G.N. Jones ssp. tsugense ) on the 13 year basal area growth of large (>45.7 cm diameter at breast height) western hemlock ( Tsuga heterophylla (Raf.) Sarg.) trees in an old-growth forest in southwestern Washington state. We controlled for spatial effects on tree growth by utilizing twenty-seven 0.4 ha plots that had uninfected, lightly, moderately, and severely infected classes of trees present on each plot. Basal area growth was analyzed using analysis of covariance of a randomized block design that was a balanced design with four treatments (infection classes) and 27 replicates (blocks or 0.4 ha plots), with initial diameter as the covariate. Basal area growth was found to be significantly different among the infection classes (F[3,77] = 10.09 and p < 0.0001). Growth of the light and moderate infection classes was not detectably different from growth of uninfected trees. However, severely infected trees grew from 16% to 46% (mean = 36%) less than uninfected trees of the same initial diameter over the period of study (1991–2004). The large trees in this stand did not show growth impacts until they were severely infected.

Herbivory is an important ecological process in forest canopies but is difficult to measure, especially for whole stands. We used the Wind River Canopy Crane in Washington State to access 101 randomly-located sample points throughout the forest canopy. This provided a relatively quick and convenient way to estimate herbivory for a whole stand. The overall level of herbivory was estimated at 1.6% of leaf area. The distribution was strongly skewed to the lower canopy where broad-leafed species experienced higher levels of herbivory. Herbivory averaged 0.3% in conifers and 13.5% in broad-leafed species. Fully half of the sample points had no detectable herbivory. Herbivory in this old-growth conifer forest is among the lowest levels published for forests around the globe and may reflect the general levels of herbivory in temperate coniferous forests during nonoutbreak conditions. Our whole-stand estimate is the first attempt at measuring herbivory for an entire forest stand in the Pacific Northwest.

The volume and complexity of their vascular systems make the dynamics of long-distance water transport in large trees difficult to study. We used heat and deuterated water (D 2 O) as tracers to characterize whole-tree water transport and storage properties in individual trees belonging to the coniferous species Pseudotsuga menziesii (Mirb.) Franco and Tsuga heterophylla (Raf.) Sarg. The trees used in this study spanned a broad range of height (13.5-58 m) and diameter (0.14-1.43 m). Sap flow was monitored continuously with heat dissipation probes near the base of the trunk prior to, during and following injec- tion of D 2 O. The transit time for D 2 O transport from the base of the trunk to the upper crown and the tracer resi- dence time were determined by measuring hydrogen iso- tope ratios in water extracted from leaves sampled at regular intervals. Transit times for arrival of D 2 O in the upper crown ranged from 2.5 to 21 d and residence times ranged from 36 to 79 d. Estimates of maximum sap veloc- ity derived from tracer transit times and path length ranged from 2.4 to 5.4 m d − 1 . Tracer residence time and half-life increased as tree diameter increased, independent of species. Species-independent scaling of tracer velocity with sapwood-specific conductivity was also observed. When data from this study were combined with similar data from an earlier study of four tropical angiosperm trees, species-independent scaling of tracer velocity and residence time with sapwood hydraulic capacitance was observed. Sapwood capacitance is an intrinsic tissue-level property that appears to govern whole-tree water trans- port in a similar manner among both tracheid- and vessel- bearing species.

We investigated the distribution and severity of trees infected with western hemlock dwarf mistletoe (Arceuthobium tsugense (Rosendahl) G.N. Jones subsp. tsugense) in an old-growth Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) – western hemlock (Tsuga heterophylla (Raf.) Sarg.) forest. With the use of Hawksworth six-class dwarf mistletoe rating system, infection status was assessed for 3516 hemlock and true firs ≥5 cm diameter on a 12-ha stem-mapped plot located in the Cascade Mountains of southwest Washington State. Within the plot, 33% of the area had some level of infection and 25% (719) of western hemlocks, 2.2% (12) of Pacific silver fir (Abies amabilis (Dougl.) Forbes), and 29% (2) of noble fir (Abies procera Rehd.) trees were infected. Infected trees are larger than uninfected trees, on average, and within the infected tree population, the severely infected trees averaged larger than lightly infected trees. Abundant dwarf mistletoe in larger trees definitely positions the dwarf mistletoe population for future spread. Ripley's K analysis indicates a negative association between infected and uninfected hemlock trees, confirming that the infected trees form distinct dwarf mistletoe infection centers. The infection centers are actively spreading at their margins, which was confirmed by nearest neighbor analysis. Heavily infected trees had a negative association with uninfected trees, while lightly infected trees had a positive association with uninfected trees.

Forest managers need a comprehensive scientific understanding of natural stand development processes when designing silvicultural systems that integrate ecological and economic objectives, including a better appreciation of the nature of disturbance regimes and the biological legacies, such as live trees, snags, and logs, that they leave behind. Most conceptual forest development models do not incorporate current knowledge of the: (1) complexity of structures (including spatial patterns) and developmental processes; (2) duration of development in long-lived forests; (3) complex spatial patterns of stands that develop in later stages of seres; and particularly (4) the role of disturbances in creating structural legacies that become key elements of the post-disturbance stands. We elaborate on existing models for stand structural development using natural stand development of the Douglas-fir—western hemlock sere in the Pacific Northwest as our primary example; most of the principles are broadly applicable while some processes (e.g. role of epicormic branches) are related to specific species. We discuss the use of principles from disturbance ecology and natural stand development to create silvicultural approaches that are more aligned with natural processes. Such approaches provide for a greater abundance of standing dead and down wood and large old trees, perhaps reducing short-term commercial productivity but ultimately enhancing wildlife habitat, biodiversity, and ecosystem function, including soil protection and nutrient retention. # 2002 Elsevier Science B.V. All rights reserved.