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This study investigates competing vegetation effects on foliar and total plant-derived nutrient ratios, nutrient use efficiency (NUE), and foliar nutrient content and concentration of ecosystem components using vector analysis for 19-year-old Douglas-fir (Pseudotsuga menzeisii (Mirb.) Franco), western hemlock (Tsuga hereophylla (Raf.) Sarg.), western redcedar (Thuja plicata Donn ex D. Don), and grand fir (Abies grandis (Dougl.) Lindl.) stands in Oregon's Coast Range and for Douglas-fir and western redcedar in Oregon's Cascade foothills. Treatments included the Control, which received no spring release herbicide applications, and vegetation management (VM), which received 5 years of spring release herbicide applications, reducing competing vegetation abundance. VM increased the NUE of N, P, Mg, S, and Cu across all species when calculated with total plant-derived carbon and of all nutrients when calculated with stemwood carbon. VM often produced more harvestable and plant-derived carbon per unit nutrient fixed, improving the NUE of stands managed for carbon sequestration and timber. Species showed different stand nutrient requirements, evident through foliar and plant-derived nutrient ratios and their relationship with biomass production. Grand fir may obtain larger biomass increments for a given P:N ratio in plant-derived tissue and may be efficient in P-limited Coast Range sites. [ABSTRACT FROM AUTHOR]

Abstract Background Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco) plays a critical role in the ecology and economy of Western North America. This conifer species comprises two distinct varieties: the coastal variety (var. menziesii) along the Pacific coast, and the interior variety (var. glauca) spanning the Rocky Mountains into Mexico, with instances of inter-varietal hybridization in Washington and British Columbia. Recent investigations have focused on assessing environmental pressures shaping Douglas-fir’s genomic variation for a better understanding of its evolutionary and adaptive responses. Here, we characterize range-wide population structure, estimate inter-varietal hybridization levels, identify candidate loci for climate adaptation, and forecast shifts in species and variety distribution under future climates. Results Using a custom SNP-array, we genotyped 540 trees revealing four distinct clusters with asymmetric admixture patterns in the hybridization zone. Higher genetic diversity observed in coastal and hybrid populations contrasts with lower diversity in inland populations of the southern Rockies and Mexico, exhibiting a significant isolation by distance pattern, with less marked but still significant isolation by environment. For both varieties, we identified candidate loci associated with local adaptation, with hundreds of genes linked to processes such as stimulus response, reactions to chemical compounds, and metabolic functions. Ecological niche modeling revealed contrasting potential distribution shifts among the varieties in the coming decades, with interior populations projected to lose habitat and become more vulnerable, while coastal populations are expected to gain suitable areas. Conclusions Overall, our findings provide crucial insights into the population structure and adaptive potential of Douglas-fir, with the coastal variety being the most likely to preserve its evolutionary path throughout the present century, which carry implications for the conservation and management of this species across their range.

AbstractSeedlings of 95 greenhouse-grown interior Douglas-fir half-sibling families originating from four breeding zones were subject to overwatering and periodic underwatering and compared to an optimally watered control (15 per family = 1425 seedlings total). Averaged over all trees, annual height was reduced by 4% and 11% compared to controls for overwatering and periodic underwatering, respectively. Family overwatered seedling height was 135–75%, and underwatered height was 100–75% of their respective control heights. Shorter families under stress grew like their control compared to taller families, but this was reversed in taller families when the watering conditions were ideal. A survival analysis indicated that after 30–50 days without water, family survival varied from 25% to 90%, and most trees died after 60 days. The breeding zone had little effect on the tolerance to suboptimal water or survival during complete water withdrawal and did not interact with watering treatments. Family traits for tolerance to overwatering favoured root biomass, traits for underwatering favoured reduced stem biomass, especially in ratio to roots, and traits for survival favoured belowground biomass over aboveground biomass. No tradeoffs between tolerance to over- and underwatering were found. Some families did well in all three conditions and some in one or the other. The study results suggest that variation in family growth and survival to suboptimal watering partly relate to root and stem biomass partitioning in the interior Douglas-fir population of British Columbia. [ABSTRACT FROM AUTHOR]

The current planting of economically important timber species, such as Douglas-fir, mainly relies on genetically improved seeds from seed orchards. However, published research on the effects of climate change has largely focused on natural populations. To bridge this gap, data from 80 cooperative second-cycle coastal Douglas-fir progeny tests across eight breeding zones in western Washington and Oregon were analyzed. Climate transfer functions for age-12 growth were derived, showing significant results for the US Pacific Northwest. Region-specific transfer functions (Coast, Inland, and Cascade) displayed stronger correlations. Mean annual temperature and mean coldest month temperature were the most important climatic variables explaining growth. The study found that populations from slightly warmer areas tended to grow better but moving populations from colder to warmer areas by 2 °C (analogous to projected global warming) would result in an 8% genetic loss in age-12 height and a 25% genetic loss in age-12 volume. However, substantial diversity in climatic response was found among full-sib families within large breeding zones, suggesting that breeding and selecting suitable families for future climatic conditions within breeding zones is feasible. The study discusses potential strategies to adapt current breeding programs to address the impacts of future climate change while maintaining high population growth rates in Douglas-fir breeding programs. [ABSTRACT FROM AUTHOR]

Abstract Background Selection of climate-change adapted ecotypes of commercially valuable species to date relies on DNA-assisted screening followed by growth trials. For trees, such trials can take decades, hence any approach that supports focussing on a likely set of candidates may save time and money. We use a non-stationary statistical analysis with spatially varying coefficients to identify ecotypes that indicate first regions of similarly adapted varieties of Douglas-fir (Pseudotsuga menziesii (Mirbel) Franco) in North America. For over 70,000 plot-level presence-absences, spatial differences in the survival response to climatic conditions are identified. Results The spatially-variable coefficient model fits the data substantially better than a stationary, i.e. constant-effect analysis (as measured by AIC to account for differences in model complexity). Also, clustering the model terms identifies several potential ecotypes that could not be derived from clustering climatic conditions itself. Comparing these six identified ecotypes to known genetically diverging regions shows some congruence, as well as some mismatches. However, comparing ecotypes among each other, we find clear differences in their climate niches. Conclusion While our approach is data-demanding and computationally expensive, with the increasing availability of data on species distributions this may be a useful first screening step during the search for climate-change adapted varieties. With our unsupervised learning approach being explorative, finely resolved genotypic data would be helpful to improve its quantitative validation.

Summary: 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. [ABSTRACT FROM AUTHOR]

Downed woody debris is important for biodiversity, forest regeneration, and carbon, nutrient, and water cycling, and past studies have examined how the coarse fraction is affected by climate or harvesting. In a field study in Douglas-fir dominated forests, we expand existing knowledge by investigating the interacting effects of climate and harvesting on downed woody debris of all sizes. Across a 900-km long latitudinal gradient in British Columbia, we found that coarse woody debris (CWD, >7.5 cm diameter) in humid climates contained 700% greater carbon stocks, had 500% greater volume, and was more diverse than in arid climates. Pre- and post-harvest, small and fine woody debris comprised a higher proportion of total woody debris carbon stocks in arid than moist climates, especially after clearcutting and seed tree treatments. Harvesting generally decreased total CWD volume, but it was not depleted on any site. Harvesting substantially reduced the volume of large, highly decomposed CWD except at the two most arid sites, and losses of large CWD increased with increasing tree removal. These losses were accompanied by a pulse of fresh, small diameter CWD and SWD which are short-term organic nutrient sources but have less habitat value than larger pieces and contribute to fuel loads. Because CWD was less abundant in arid than humid mature forests, care must be taken on arid sites to avoid its depletion during harvesting, especially clearcutting, where future woody debris inputs will not occur for decades. [ABSTRACT FROM AUTHOR]

In the face of climate change and associated increases in disturbances, some areas, known as refugia, will remain or become newly habitable for species, while others will be lost. Planning and managing for refugia can support biodiversity and conservation. However, without explicit consideration of justice, planning and management for refugia risks unnecessarily limiting information about local conditions and traditional practices that may be contained in Indigenous knowledges, and causing maladaptive consequences such as exclusion of Indigenous communities from decision-making and from protected areas, with loss of use of traditional plants and animals. The article proposes a new concept, Indigenous refugia, that incorporates three types of justice into existing theories of refugia for conservation in the face of climate change: recognition justice as understanding and respect for Indigenous values, experiences, and knowledges; procedural justice in collaboration and decision-making; and distributional justice as access to species and lands that sustain cultural and social processes. It presents a potential example of Indigenous refugia for the planning and management for climate-vulnerable Douglas-fir in New Mexico in collaboration with Pueblo, Diné (Navajo), Nde (Apache), and other Indigenous peoples with ancestral lands in the area. [ABSTRACT FROM AUTHOR]

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Summary: Tree growth performance can be partly explained by genetics, while a large proportion of growth variation is thought to be controlled by environmental factors. However, to what extent DNA methylation, a stable epigenetic modification, contributes to phenotypic plasticity in the growth performance of long‐lived trees remains unclear. In this study, a comparative analysis of targeted DNA genotyping, DNA methylation and mRNAseq profiling for needles of 44‐year‐old Douglas‐fir trees (Pseudotsuga menziesii (Mirb.) Franco) having contrasting growth characteristics was performed. In total, we identified 195 differentially expressed genes (DEGs) and 115 differentially methylated loci (DML) that are associated with genes involved in fitness‐related processes such as growth, stress management, plant development and energy resources. Interestingly, all four intronic DML were identified in mega‐sized (between 100 and 180 kbp in length) and highly expressed genes, suggesting specialized regulation mechanisms of these long intron genes in gymnosperms. DNA repetitive sequences mainly comprising long‐terminal repeats of retroelements are involved in growth‐associated DNA methylation regulation (both hyper‐ and hypomethylation) of 99 DML (86.1% of total DML). Furthermore, nearly 14% of the DML was not tagged by single nucleotide polymorphisms, suggesting a unique contribution of the epigenetic variation in tree growth. [ABSTRACT FROM AUTHOR]

Summary: Tree seedlings from populations native to drier regions are often assumed to be more drought tolerant than those from wetter provenances. However, intraspecific variation in drought tolerance has not been well‐characterized despite being critical for developing climate change mitigation and adaptation strategies, and for predicting the effects of drought on forests.We used a large‐scale common garden drought‐to‐death experiment to assess range‐wide variation in drought tolerance, measured by decline of photosynthetic efficiency, growth, and plastic responses to extreme summer drought in seedlings of 73 natural populations of the two main varieties of Douglas‐fir (Pseudotsuga menziesii var. menziesii and var. glauca).Local adaptation to drought was weak in var. glauca and nearly absent in menziesii. Var. glauca showed higher tolerance to drought but slower growth than var. menziesii. Clinal variation in drought tolerance and growth species‐wide was mainly associated with temperature rather than precipitation. A higher degree of plasticity for growth was observed in var. menziesii in response to extreme drought.Genetic variation for drought tolerance in seedlings within varieties is maintained primarily within populations. Selective breeding within populations may facilitate adaptation to drought more than assisted gene flow. [ABSTRACT FROM AUTHOR]

Fire injury stresses Douglas-fir trees (Pseudotsuga menziesii) that survive a wildfire event, allowing subsequent Douglas-fir beetle (Dendroctonus pseudotsugae) infection to kill trees that may have otherwise survived. This study aimed to determine how fire injury, stand, and tree characteristics drive Douglas-fir beetle host tree selection five years post-fire. We paired 28 adjacent beetle-infected and uninfected stands (infected N = 14) and 140 Douglas-fir trees (infected N = 70) within the 416 Fire burn area in Southwest Colorado. We found no statistically significant differences between infected and uninfected stand characteristics. Individual tree height, DBH, and bark char severity index were significantly higher in infected versus uninfected trees. We created a regression decision tree model to determine the influence of fire injury and tree characteristics on the probability of infection. Trees with a height ≥ 27 m, bark char height < 2.3 m, and DBH < 80 cm had the greatest probability of attack (100%). Trees with a height < 27 m, bark char severity index < 5.5, and DBH < 49 cm had the lowest probability of attack (3.7%). Understanding the influence of fire on Douglas-fir beetle host selection allows land managers to model potential epidemic outbreaks and guide proactive management actions that may reduce beetle outbreak severity or preserve high-value trees not killed by fire. [ABSTRACT FROM AUTHOR]

Downed woody debris is important for biodiversity, forest regeneration, and carbon, nutrient, and water cycling, and past studies have examined how the coarse fraction is affected by climate or harvesting. In a field study in Douglas-fir dominated forests, we expand existing knowledge by investigating the interacting effects of climate and harvesting on downed woody debris of all sizes. Across a 900-km long latitudinal gradient in British Columbia, we found that coarse woody debris (CWD, >7.5 cm diameter) in humid climates contained 700% greater carbon stocks, had 500% greater volume, and was more diverse than in arid climates. Pre- and post-harvest, small and fine woody debris comprised a higher proportion of total woody debris carbon stocks in arid than moist climates, especially after clearcutting and seed tree treatments. Harvesting generally decreased total CWD volume, but it was not depleted on any site. Harvesting substantially reduced the volume of large, highly decomposed CWD except at the two most arid sites, and losses of large CWD increased with increasing tree removal. These losses were accompanied by a pulse of fresh, small diameter CWD and SWD which are short-term organic nutrient sources but have less habitat value than larger pieces and contribute to fuel loads. Because CWD was less abundant in arid than humid mature forests, care must be taken on arid sites to avoid its depletion during harvesting, especially clearcutting, where future woody debris inputs will not occur for decades.

The Elliott State Research Forest comprises 33,700 ha of temperate, Douglas-fir rainforest along North America's Pacific Coast (Oregon, United States). In 2015, naturally regenerated stands at least 92 years old covered 49% of the research area and sawtimber plantations younger than 68 years another 50%. During the winter of 2015–2016, a forest wide inventory sampled both naturally regenerated and plantation stands, recording 97,424 trees on 17,866 plots in 738 stands. The resulting dataset is atypical for the area as plot locations were not restricted to upland, commercially harvestable timber. Multiage stands and riparian areas were therefore documented along with plantations 2–61 years old and trees retained through clearcut harvests. This dataset constitutes the only open access, stand-based forest inventory currently available for a large area within the Oregon Coast Range. The dataset enables development of suites of models as well as many comparisons across stand ages and types, both at stand level and at the level of individual trees. DataSet: https://doi.org/10.7267/xp68kq79g DataSet License: CC-BY-NC-SA [ABSTRACT FROM AUTHOR]

Tree-ring δ15N may depict site-specific, long-term patterns in nitrogen (N) dynamics under N2-fixing species, but field trials with N2-fixing tree species are lacking and the relationship of temporal patterns in tree-ring δ15N to soil N dynamics is controversial. We examined whether the tree-ring δ15N of N2-fixing red alder (Alnus rubra Bong.) would mirror N accretion rates and δ15N of soils and whether the influence of alder-fixed N could be observed in the wood of a neighboring conifer. We sampled a 27-year-old replacement series trial on south-eastern Vancouver Island, with red alder and coastal Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco) planted in five proportions (0/100, 11/89, 25/75, 50/50 and 100/0) at a uniform stem density. An escalation in forest floor N content was evident with an increasing proportion of red alder, equivalent to a difference of ~750 kg N ha−1 between 100% Douglas-fir versus 100% alder. The forest floor horizon also had high δ15N values in treatments with more red alder. Red alder had a consistent quadratic fit in tree-ring δ15N over time, with a net increase of |$\sim$| 1.5‰, on average, from initial values, followed by a plateau or slight decline. Douglas-fir tree-ring δ15N, in contrast, was largely unchanged over time (in three of four plots) but was significantly higher in the 50/50 mix. The minor differences in current leaf litter N content and δ15N between alder and Douglas-fir, coupled with declining growth in red alder, suggests the plateau or declining trend in alder tree-ring δ15N could coincide with lower N2-fixation rates, potentially by loss in alder vigor at canopy closure, or down-regulation via nitrate availability. [ABSTRACT FROM AUTHOR]

Birds are under selective pressure to orient their nests in a direction that reduces the impact of climatic extremes and the risk of the nest being detected by predators. Woodpeckers (Picidae), despite raising young in tree cavities that are safer from predators, should orient the entrances of their nest cavities in a direction that results in a favorable cavity microclimate. Because Picid nest cavity orientation is not well-studied in the Pacific Northwest of North America, we measured nest cavity orientation of White-headed Dryobates albolarvatus, Hairy D. villosus, and Black-backed Picoides arcticus Woodpeckers, Northern Flicker Colaptes auratus, and Williamson's Sapsucker Sphyrapicus thyroideus within burned and unburned, dry conifer forests of the eastern Cascade Range in Washington, USA (N = 684). Each woodpecker species had a mean angle of nest cavity orientation in a northerly or easterly direction and ranged from 12.3° for the Northern Flicker to 117.6° for the Black-backed Woodpecker. Hairy Woodpeckers, Northern Flickers, and White-headed Woodpeckers showed strong evidence for nonrandom nest cavity orientation, and Northern Flickers showed particularly high dispersion with a multimodal distribution of cavity orientations. Although the mean and variance of cavity orientations did not differ substantially between woodpeckers, cavity orientation differed based on nest tree species. Woodpeckers may orient their nest cavities in an easterly direction so that cavities warm quicker in the morning, to avoid hot afternoon temperatures and prevailing west and north westerly winds, and because the majority of cavities were located in burned forest where shading from canopy cover is reduced. [ABSTRACT FROM AUTHOR]

Detailed information about the historical range of variability in wildfire activity informs adaptation to future climate and disturbance regimes. Here, we describe one of the first annually resolved reconstructions of historical (1500–1900 ce) fire occurrence in coast Douglas‐fir dominated forests of the west slope of the Cascade Range in western Oregon. Mean fire return intervals (MFRIs) across 16 sites within our study area ranged from 6 to 165 years. Variability in MFRIs was strongly associated with average maximum summer vapor pressure deficit. Fire occurred infrequently in Douglas‐fir forest stands seral to mountain hemlock or silver fir, but fire frequency was much shorter than predicted by theory in other forest types. MFRIs within Douglas‐fir stands seral to western hemlock or grand fir ranged from 19 to 45 years, and MFRIs in stands seral to Douglas‐fir ranged from 6 to 11 years. There was little synchrony in fire occurrence or tree establishment across 16 sites separated by 4 km. The lack of synchrony in fire suggests that large, wind‐driven fire events that are often considered to be characteristic of coast Douglas‐fir forests were not an important driver of succession in our study area during the last ~400–500 years. Climate was more arid than normal during fire years in most forest types, but historical fire in stands seral to Douglas‐fir was strongly associated with antecedent moisture and less strongly associated with drought. We interpret the extraordinary tempo of fire we observed in stands seral to Douglas‐fir and the unique climate pattern associated with fire in these stands to be indicative of Indigenous fire stewardship. This study provides evidence of far more frequent historical fire in coast Douglas‐fir forests than assumed by managers or scientists—including some of the most frequent fire return intervals documented in the Pacific Northwest. We recommend additional research across the western Cascades to create a comprehensive account of historical fire in highly productive forests with significant cultural, economic, and ecological importance. [ABSTRACT FROM AUTHOR]

Maximum stand density index (SDIMAX) represents the carrying capacity of a forest stand based on the relationship between the number of trees and their size. Plot‐level inventory data provided through a collaborative network of federal, state, and private forest management groups were utilized to develop SDIMAX models for important Pacific Northwest conifers of western Washington and Oregon, USA. The influence of site‐specific climatic and environmental variables was explored within an ensemble learning model. Future climate projections based on global circulation models under different representative CO2 concentration pathways (RCP 4.5 and RCP 8.5) and timeframes (2050s and 2080s) were utilized in a space‐for‐time substitution to understand potential shifts in modeled SDIMAX. A majority of the region showed decreases in carrying capacity under future climate conditions. Modeled mean SDIMAX decreased 5.4% and 11.4% for Douglas‐fir (Pseudotsuga menziesii (Mirb.) Franco) dominated forests and decreased 6.6% and 8.9% for western hemlock (Tsuga heterophylla (Raf.) Sarg.) and Pacific silver fir (Abies amabilis), dominated forests under the RCP 4.5 in the 2050s and RCP 8.5 in the 2080s, respectively. Projected future conditions often fall outside the range of any contemporary climate profile, resulting in what may be referred to as extramural conditions. Within the study region, 45% and 46% of climate variables included in the final model were extramural for the Douglas‐fir and hemlock models, respectively, under RCP 8.5 in the 2080s. Although extrapolating beyond the range of input data is not appropriate and many unknowns remain regarding future climate projections, these results allow for general interpretations of the direction and magnitude of potential shifts in forest carrying capacity. Recommendations for Resource Managers: These results present forest managers and policy‐makers with the ability to make location‐specific interpretations of potential shifts in forest stand carrying capacity under potential future climate projection scenarios, which show forest communities of the Pacific Northwest may be facing a longer, warmer, droughtier growing season.Across the study region, a majority of the landscape is seeing a downward shift in the modeled maximum carrying capacity under future climate projections.The modeled carrying capacity of mixed species stands appear to be more resilient to projected changes in climate.Planting and thinning to lower densities may provide resilience and capture the projected increases in density‐related mortality.Across all projected future climate scenarios fall into extramural conditions, which are a combination of climatic conditions not currently experienced in the study area. [ABSTRACT FROM AUTHOR]

Abstract Detailed information about the historical range of variability in wildfire activity informs adaptation to future climate and disturbance regimes. Here, we describe one of the first annually resolved reconstructions of historical (1500–1900 ce) fire occurrence in coast Douglas‐fir dominated forests of the west slope of the Cascade Range in western Oregon. Mean fire return intervals (MFRIs) across 16 sites within our study area ranged from 6 to 165 years. Variability in MFRIs was strongly associated with average maximum summer vapor pressure deficit. Fire occurred infrequently in Douglas‐fir forest stands seral to mountain hemlock or silver fir, but fire frequency was much shorter than predicted by theory in other forest types. MFRIs within Douglas‐fir stands seral to western hemlock or grand fir ranged from 19 to 45 years, and MFRIs in stands seral to Douglas‐fir ranged from 6 to 11 years. There was little synchrony in fire occurrence or tree establishment across 16 sites separated by 4 km. The lack of synchrony in fire suggests that large, wind‐driven fire events that are often considered to be characteristic of coast Douglas‐fir forests were not an important driver of succession in our study area during the last ~400–500 years. Climate was more arid than normal during fire years in most forest types, but historical fire in stands seral to Douglas‐fir was strongly associated with antecedent moisture and less strongly associated with drought. We interpret the extraordinary tempo of fire we observed in stands seral to Douglas‐fir and the unique climate pattern associated with fire in these stands to be indicative of Indigenous fire stewardship. This study provides evidence of far more frequent historical fire in coast Douglas‐fir forests than assumed by managers or scientists—including some of the most frequent fire return intervals documented in the Pacific Northwest. We recommend additional research across the western Cascades to create a comprehensive account of historical fire in highly productive forests with significant cultural, economic, and ecological importance.

Understanding the genomic and environmental basis of cold adaptation is key to understand how plants survive and adapt to different environmental conditions across their natural range. Univariate and multivariate genome-wide association (GWAS) and genotype-environment association (GEA) analyses were used to test associations among genome-wide SNPs obtained from whole-genome resequencing, measures of growth, phenology, emergence, cold hardiness, and range-wide environmental variation in coastal Douglas-fir (Pseudotsuga menziesii). Results suggest a complex genomic architecture of cold adaptation, in which traits are either highly polygenic or controlled by both large and small effect genes. Newly discovered associations for cold adaptation in Douglas-fir included 130 genes involved in many important biological functions such as primary and secondary metabolism, growth and reproductive development, transcription regulation, stress and signaling, and DNA processes. These genes were related to growth, phenology and cold hardiness and strongly depend on variation in environmental variables such degree days below 0c, precipitation, elevation and distance from the coast. This study is a step forward in our understanding of the complex interconnection between environment and genomics and their role in cold-associated trait variation in boreal tree species, providing a baseline for the species' predictions under climate change.

Climate projections predict more frequent and severe drought in coastal Pseudotsuga menziesii forests of western North America, raising concerns over how to promote drought adaptation. Thinning often increases drought resistance (the ability to maintain growth during a drought) and resilience (the ability to recover growth after a drought), but these effects vary with thinning intensity, shift over time, and may have tradeoffs with fiber production. We collected tree cores from a long-term thinning study with four residual density levels replicated across both uniform thinning and thinning with gaps, and used annual growth data to investigate responses to droughts occurring 8 and 21 years after thinning. For the first drought, resistance and resilience were higher in treatments with lower residual densities. For the second drought, there were no differences in drought response between the lowest and highest residual density treatments, and all treatments had lower drought resistance and resilience than for the first drought. Spatial arrangement had little impact on drought resistance or resilience and residual density level had a significant effect on the periodic annual volume increment—drought resistance tradeoff. Our results suggest that thinning can promote drought adaptation in Pseudotsuga menziesii forests, but these effects dissipate over time. [ABSTRACT FROM AUTHOR]

Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), one of the most commercially important tree species in western North America and one of the most valuable timber trees worldwide, was introduced to Europe in 1827. It became a major species for afforestation in Western Europe after WWII, currently grows in 35 countries on over 0.83 million ha and is one of the most widespread non-native tree species across the continent. A lower sensitivity to drought makes Douglas-fir a potential alternative to the more drought-sensitive Norway spruce so its importance in Europe is expected to increase in the future. It is one of the fastest growing conifer species cultivated in Europe, with the largest reported dimensions of 2.3 m in diameter and 67.5 m in height. Pure stands have high productivity (up to 20 m3 ha−1a−1) and production (over 1000 m3 ha−1). The species is generally regenerated by planting (initial stocking density from less than 1000 seedlings ha−1 to more than 4000 ha−1), using seedlings of European provenance derived from seed orchards or certified seed stands. As the range of end-uses of its wood is very wide, the rotation period of Douglas-fir is highly variable and ranges between 40 and 120 years. When the production of large-sized, knot-free timber is targeted, thinnings are always coupled with pruning up to 6 m. There is an increasing interest in growing Douglas-fir in mixtures and managing stands through close-to-nature silviculture, but the species' intermediate shade tolerance means that it is best managed through group selection or shelterwood systems. [ABSTRACT FROM AUTHOR]

High‐severity wildfires create pulses of snags that serve a variety of functions as they decompose over time. Snag‐related benefits (and hazards) are often linked to specific decomposition stages, but snag decomposition rates and pathways are not well understood in many forest types. We examined temporal patterns of snag decomposition, wildlife cavity creation, and surface woody fuel dynamics in dry coniferous forests of the interior Pacific Northwest region of North America by sampling 159 forest stands within a 39‐year chronosequence of stand‐replacing wildfires in dry coniferous forests dominated by ponderosa pine and Douglas‐fir. We found that most snags broke or fell during the first 15 years after wildfire; small‐diameter snags mostly broke off at or near ground level, while many large‐diameter snags initially broke off above a height of 2 m and then remained standing for an extended period. Ponderosa pine and lodgepole pine snags fell earlier than Douglas‐fir and true fir snags of comparable diameter classes. Wildlife cavities were most common in stands surveyed 8–20 years after fire and in snags with broken tops but were not limited to large diameter snags. Cavity snag diameters ranged from 17 to 98 cm, with 64% of cavity snags having diameters between 30 and 60 cm and 22% of cavity snags having diameters >60 cm. Surface woody fuels increased as snags broke and fell, reaching maximum levels in all size classes 15–20 years after fire. The percentage of large diameter rotten woody fuels increased steadily beginning about 15 years after fire, with implications for fuel management and subsequent wildfire behavior and severity. Our study supports the proposition that fire‐killed snags represent a transient resource pulse in which the relative contribution of snags to different ecological functions varies with snag diameter, species, time since fire, and landscape position. [ABSTRACT FROM AUTHOR]

A series of ISO 5658-2 flame spread experiments on Douglas-fir specimens were carried out to investigate the relationship between flame spread characteristics and thermal thickness of solid combustibles, which is greatly influenced by char formation during combustion. Computations using the fire dynamics simulator (FDS) were performed to recreate these experiments. The measured and calculated flame spread characteristics were then compared and analyzed based on thermal thickness and char residue ratio. The results clearly show that the effects of char formation on the surface of wood during combustion are more noticeable in thermally thick regimes than in thermally thin regimes. The findings provide an important foundational understanding of the influence of char formation on solid combustibles on flame retardant properties, which has not previously been studied. [ABSTRACT FROM AUTHOR]

Cultivation of non‐native tree species is a promising option to adapt managed forests to climate change. However, consequences of non‐native tree species on flora, fauna, and microorganisms that occur in forest ecosystems (forest associates) need to be considered when managing forests. We lack a solid understanding of how cultivating non‐native tree species in pure stands and in mixture with native tree species impacts abundance and species diversity of forest associates. We compared abundance, alpha‐, beta‐ and gamma‐diversity of eight forest‐floor‐associated taxa that are relevant for ecosystem functioning (including fungi, plants, arthropods, and small mammals) between different forest ecosystems. We addressed pure stands of non‐native coniferous Douglas‐fir (Pseudotsuga menziesii) and two native species, broad‐leaved European beech (Fagus sylvatica) and coniferous Norway spruce (Picea abies), as well as mixed stands of European beech with each conifer in two regions with differing site conditions in temperate Northwest Germany. Diversity indices revealed differences among species groups. Gamma‐diversity and alpha‐diversity of forest associates in Douglas‐fir and spruce stands were usually higher than in beech stands. Estimates of species diversity of mixed stands are intermediate between estimates for the respective pure stands. Differences in the diversity between the two study regions were highly taxon specific with no clear support for a general trend toward a higher or lower species diversity of forest associates depending on site quality. Abundance values show a pattern similar to our diversity estimates, but with a higher statistical uncertainty. Non‐native Douglas‐fir stands provided habitats to support associated species communities of equally high or higher diversity than stands of native beech or spruce. Mixed stands of non‐native and native tree species may be a management option to achieve different goals, that is, to provide habitats for species that are strictly associated with beech and to increase resilience to climate change. However, the overall diversity of forest‐floor‐associated biota is not improved by cultivating Douglas‐fir or spruce in mixture with beech. [ABSTRACT FROM AUTHOR]

In forest resource inventory, tree height is often estimated by easily measurable diameter from height-diameter model. In this study, we tried to use random forest (RF), an important machine learning method, to model individual-tree height. Results showed that the optimized RF model had better fitting and prediction accuracy (R2 = 0.8146 and RMSE = 2.2527 m). In terms of relative importance, diameter at breast height (DBH) was the most important factor, followed by neighborhood-related variables and other variables related to environmental conditions. Further, tree height was generally positively affected by DBH, mean diameter of neighbors, DBH dominance, number of neighbors, and mean annual precipitation, but negatively affected by elevation. The results indicated that the RF-based height model was statistically reliable and highly accurate, and it had strong interpretability with ecological significance. Our study will provide a new perspective for the application of machine learning algorithms to forest dynamic modeling. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Abstract High‐severity wildfires create pulses of snags that serve a variety of functions as they decompose over time. Snag‐related benefits (and hazards) are often linked to specific decomposition stages, but snag decomposition rates and pathways are not well understood in many forest types. We examined temporal patterns of snag decomposition, wildlife cavity creation, and surface woody fuel dynamics in dry coniferous forests of the interior Pacific Northwest region of North America by sampling 159 forest stands within a 39‐year chronosequence of stand‐replacing wildfires in dry coniferous forests dominated by ponderosa pine and Douglas‐fir. We found that most snags broke or fell during the first 15 years after wildfire; small‐diameter snags mostly broke off at or near ground level, while many large‐diameter snags initially broke off above a height of 2 m and then remained standing for an extended period. Ponderosa pine and lodgepole pine snags fell earlier than Douglas‐fir and true fir snags of comparable diameter classes. Wildlife cavities were most common in stands surveyed 8–20 years after fire and in snags with broken tops but were not limited to large diameter snags. Cavity snag diameters ranged from 17 to 98 cm, with 64% of cavity snags having diameters between 30 and 60 cm and 22% of cavity snags having diameters >60 cm. Surface woody fuels increased as snags broke and fell, reaching maximum levels in all size classes 15–20 years after fire. The percentage of large diameter rotten woody fuels increased steadily beginning about 15 years after fire, with implications for fuel management and subsequent wildfire behavior and severity. Our study supports the proposition that fire‐killed snags represent a transient resource pulse in which the relative contribution of snags to different ecological functions varies with snag diameter, species, time since fire, and landscape position.

Phenology of diameter growth in trees has been studied for many years but generally using a limited number of sites and genotypes. In this project provenances of Douglas-fir (Pseudotsuga menziesii var. menziesii) planted across a wide range of environments were used to evaluate diameter growth and phenology to an extreme heat event and to seasonal conditions. Sampling was done in nine common gardens in western Oregon and Washington, USA that included a coastal site, a low elevation interior site, and high elevation interior site on three latitude bands. In 2021, three provenances at four sites were monitored for late-season diameter growth following an extreme heat event in June. In 2022, five provenances were evaluated for full-seasonal diameter growth at four growth intervals across all nine sites. Growth after the 2021 extreme heat event varied by planting site, with the greatest growth observed at mild (cooler/wetter) sites and the least growth observed at arid (hotter/drier) sites. Comparisons of diameter growth over the late season period (i.e., July–October) showed that 2022 growth was 1.25-fold to 3.70-fold higher, presumably due to premature growth cessation in 2021. In 2022, diameter growth rates showed significant variation by site and provenance; the interaction between site and provenance was significant in all but the first growth interval. Intra-seasonal growth rates were not consistent across sites; there was substantial diameter growth early in the season on sites with continental climates, little diameter growth by any provenance after early August at three high elevation sites, and most sources continued growth later in the season at coastal sites. One source differed more in phenology across sites than the others, apparently due to its susceptibility to a needle-cast disease. Cambial phenology varied with genetic and site factors that control the timing of growth and intrinsic growth rates. In addition, the interaction between genotype and site resulted in some provenances performing better than others on some sites as a function of disease susceptibility or response to environmental factors. Understanding phenology of seed sources under a range of site conditions should help predict diameter growth under different climate conditions in the future.

Abstract Cultivation of non‐native tree species is a promising option to adapt managed forests to climate change. However, consequences of non‐native tree species on flora, fauna, and microorganisms that occur in forest ecosystems (forest associates) need to be considered when managing forests. We lack a solid understanding of how cultivating non‐native tree species in pure stands and in mixture with native tree species impacts abundance and species diversity of forest associates. We compared abundance, alpha‐, beta‐ and gamma‐diversity of eight forest‐floor‐associated taxa that are relevant for ecosystem functioning (including fungi, plants, arthropods, and small mammals) between different forest ecosystems. We addressed pure stands of non‐native coniferous Douglas‐fir (Pseudotsuga menziesii) and two native species, broad‐leaved European beech (Fagus sylvatica) and coniferous Norway spruce (Picea abies), as well as mixed stands of European beech with each conifer in two regions with differing site conditions in temperate Northwest Germany. Diversity indices revealed differences among species groups. Gamma‐diversity and alpha‐diversity of forest associates in Douglas‐fir and spruce stands were usually higher than in beech stands. Estimates of species diversity of mixed stands are intermediate between estimates for the respective pure stands. Differences in the diversity between the two study regions were highly taxon specific with no clear support for a general trend toward a higher or lower species diversity of forest associates depending on site quality. Abundance values show a pattern similar to our diversity estimates, but with a higher statistical uncertainty. Non‐native Douglas‐fir stands provided habitats to support associated species communities of equally high or higher diversity than stands of native beech or spruce. Mixed stands of non‐native and native tree species may be a management option to achieve different goals, that is, to provide habitats for species that are strictly associated with beech and to increase resilience to climate change. However, the overall diversity of forest‐floor‐associated biota is not improved by cultivating Douglas‐fir or spruce in mixture with beech.

Adult individuals and seeds of two mixed stands of coastal and interior Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) were analysed for genetic differentiation between the two varieties and evidence of intervarietal pollination. Clear genetic discrimination between the two varieties was observed based on multilocus evaluation of nine microsatellite markers using other Douglas-fir stands of known variety composition for comparison. Analysis of pollination distances showed that 80% of pollinations took place within a distance of about 44–55 m. Analysis of stand structure showed clearly separated areas of mainly coastal or interior Douglas-fir within both stands. Together with short pollination distances this led to an apparent dominance of intravarietal pollinations. However, analysis of pollination partners of trees growing near the border of the variety specific areas, does not indicate the existence of reproductive barriers between trees of the two varieties growing in mixed stands. Therefore, commercial seed harvesting in mixed stands should be avoided if the production of seed lots of pure coastal or interior Douglas-fir is intended. [ABSTRACT FROM AUTHOR]

Severe and frequent heat and drought events challenge the survival and development of long-generation trees. In this study, we investigated the genomic basis of heat tolerance, water use efficiency and growth by performing genome-wide association studies in coastal Douglas-fir (Pseudotsuga menziesii) and intervarietal (menziesii × glauca) hybrid seedlings. GWAS results identified 32 candidate genes involved in primary and secondary metabolism, abiotic stress and signaling, among other functions. Water use efficiency (inferred from carbon isotope discrimination), photosynthetic capacity (inferred from %N), height and heat tolerance (inferred from electrolyte leakage in a heat stress experiment) were significantly different among Douglas-fir families and varieties. High-elevation seed sources had increased water use efficiency, which could be a result of higher photosynthetic capacity. Similarly, families with greater heat tolerance also had higher water use efficiency and slower growth, suggesting a conservative growth strategy. Intervarietal hybrids showed increased heat tolerance (lower electrolyte leakage at 50 and 55 °C) and higher water use efficiency compared with coastal families, suggesting that hybridization might be a source of pre-adapted alleles to warming climates and should be considered for large-scale reforestation projects under increasingly arid conditions. [ABSTRACT FROM AUTHOR]

Across the western United States, large, severe wildfires in montane forests are creating treeless patches that can fail to reforest naturally due to a lack of seed sources and a warming climate. Nursery-grown tree seedlings are commonly planted by land managers into these areas to promote forest recovery and resilience, but uncertainty exists about what influences their survival across the landscape. We obtained survival monitoring data that had been collected one growing season after planting for 5656 tree seedlings, which were distributed across nine montane wildfires and four National Forests in Colorado, USA. We used these data to examine how seedling survival varied across a range of factors, including long-term average climatic conditions, post-planting weather conditions, and locations and species of seed lot collection. We found that survival after one growing season averaged 80 % across all plots but ranged from 0 % to 100 %. Survival was greater at cooler, wetter sites and at sites planted in warmer, wetter years. Survival was also greater for ponderosa pine than for Douglas-fir, and when the locally collected seed lot used to produce the seedlings came from a site that was lower in elevation than the planting site. Our results suggest that the location and timing of tree planting, and the plant materials used, play critical roles in planting success in Colorado montane wildfires, and should help land managers optimize post-fire planted seedling survival under both current and future climatic conditions. • Monitoring data were used to examine drivers of planted tree seedling survival. • Data were collected after one growing season in nine montane Colorado wildfires. • Survival was greater for ponderosa pine seedlings than for Douglas-fir seedlings. • Survival was greater at cool, wet sites and at sites planted in warm, wet years. • Survival increased when seed sources were lower in elevation than planting sites. [ABSTRACT FROM AUTHOR]

The Elliott State Research Forest comprises 33,700 ha of temperate, Douglas-fir rainforest along North America’s Pacific Coast (Oregon, United States). In 2015, naturally regenerated stands at least 92 years old covered 49% of the research area and sawtimber plantations younger than 68 years another 50%. During the winter of 2015–2016, a forest wide inventory sampled both naturally regenerated and plantation stands, recording 97,424 trees on 17,866 plots in 738 stands. The resulting dataset is atypical for the area as plot locations were not restricted to upland, commercially harvestable timber. Multiage stands and riparian areas were therefore documented along with plantations 2–61 years old and trees retained through clearcut harvests. This dataset constitutes the only open access, stand-based forest inventory currently available for a large area within the Oregon Coast Range. The dataset enables development of suites of models as well as many comparisons across stand ages and types, both at stand level and at the level of individual trees.

Shear fracture frequently occurs in timber beams and panels subjected to transverse loads. At elevated temperatures, wood will undergo complex physical and chemical processes which significantly affect the shear properties. In this paper, the v-notched Douglas-fir specimens with three different shear planes: (a) Radial-Tangential (RT); (b) Radial-Longitudinal (RL), and (c) Longitudinal-Radial (LR), were fabricated and tested under the elevated temperatures from 20 °C to 180 °C. The digital image correlation (DIC) technique was used to measure the shear strain. It was found that the shear plane had a significant effect on the failure modes, shear strength, and shear modulus. The shear strength and shear modulus generally decreased with the increase of temperature. However, the shear strength was significantly improved when the hardening of the dry lignin occurred between 100 °C and 140 °C. Moreover, the design curve for the shear strength in Eurocode 5 is conservative for all the specimens with different shear planes. [ABSTRACT FROM AUTHOR]

Fire exclusion over the last two centuries has driven a significant fire deficit in the forests of western North America, leading to widespread changes in the composition and structure of these historically fire-adapted ecosystems. Fuel treatments have been increasingly applied over the last few decades to mitigate fire hazard, yet it is unclear whether these fuel-focused treatments restore the fire-adapted conditions and species that will allow forests to persist into the future. A vital prerequisite of restoring fire-adaptedness is ongoing establishment of fire-tolerant tree species, and both the type and reoccurrence of fuel treatments are likely to strongly influence stand trajectories. Here, we leveraged a long-term study of repeated fuel treatments in a Sierra Nevada mixed-conifer forest to examine the regeneration response of six native tree species to the repeated application of common fuel treatments: prescribed fire, mechanical, mechanical plus fire, and untreated controls. Our objectives were to (1) quantify differences in forest structure and composition following the repeated application of alternative fuel treatments that may influence the establishment environment and then (2) identify the stand structure and climate conditions influencing seedling dynamics. We found that both treatment type and intensity are highly influential in shifting forests toward more fire-adapted conditions and determining species-specific regeneration dynamics. Specifically, the conifer species tracked here increased in either colonization or persistence potential following repeated applications of fire, indicating fire may be most effective for restoring regeneration conditions broadly across species. Fire alone, however, was not enough to promote fire-adapted composition, with concurrent mechanical treatments creating more favorable conditions for promoting colonization and increasing abundances of fire-tolerant ponderosa pine. Yet, even with repeated fuel treatment application, establishment of fire-intolerant species far exceeded that of fire-tolerant species over this 20-year study period. Moreover, increasing growing season water stress negatively impacted seedling dynamics across all species regardless of treatment type and intensity, an important consideration for ongoing management under heightened climatic stress. While repeated treatments are waypoints in restoring fire-adapted conditions, more intense treatments via gap-creation or hotter prescribed fires targeting removal of fire-intolerant species will be necessary to sustain recruitment of fire-tolerant species., (© 2024 The Ecological Society of America.)

Purpose: Douglas-fir occupies a growing role in the European timber industries but its effects on soils and the environment are still not well understood. We monitored the biogeochemical cycle of major elements in 11 representative mature stands throughout France over the 2013 to 2020 period to better understand repercussions on soil processes. Methods: The present study is focused on throughfall and soil solution chemistry: concentrations were measured monthly at 15 cm and 60 cm depth and fluxes of major anions and cations were computed. Results: Our results show that nitrate strongly dominated the soil solution chemistry: concentrations were elevated (>20 mg L−1) for 8 out of 11 sites with mean annual NO3− values ranging from 2 to 253 mg L−1. Fluxes of nitrate at 60 cm of depth, ranging from 23 to 309 kg ha−1 yr−1, were much higher than those in throughfall for nine studied sites. Nitrate, which was the dominant anion in soil solution, associated either to Ca2+, Mg2+ and K+, or to Al3+ was lost out of the soil-plant system through leaching. Conclusions: These results suggest a production of NO3− in the soil exceeding tree uptake and microbial immobilization resulting in strong ongoing soil acidification at many sites and potential degradation of water quality. Significant negative correlations were found between nitrate concentration in soil solution and sandy texture, soil C and N, site elevation and, annual rainfall, but further research is necessary to clarify the role of these factors in reducing the negative impacts on terrestrial and aquatic ecosystems. [ABSTRACT FROM AUTHOR]

Patterns in foliar nitrogen (N) stable isotope ratios (δ15N) have been shown to reveal trends in terrestrial N cycles, including the identification of ecosystems where N deficiencies limit forest ecosystem productivity. However, there is a gap in our understanding of within-species variation and species-level response to environmental gradients or forest management. Our objective is to examine the relationship between site index, foliar %N, foliar δ15N and spectral reflectance for managed Douglas-fir (Pseudotsuga menziesii) and loblolly pine (Pinus taeda) plantations across their geographic ranges in the Pacific Northwest and the southeastern United States, respectively. Foliage was measured at 28 sites for reflectance using a handheld spectroradiometer, and further analyzed for δ15N and N concentration. Unlike the prior work for grasslands and shrubland species, our results show that foliar δ15N and foliar %N are not well correlated for these tree species. However, multiple linear regression models suggest a strong predictive ability of spectroscopy data to quantify foliar δ15N, with some models explaining more than 65% of the variance in the δ15N. Additionally, moderate to strong explanations of variance were found between site index and foliar δ15N (R2 = 0.49) and reflectance and site index (R2 = 0.84) in the Douglas-fir data set. The development of relationships between foliar spectral reflectance, δ15N and measures of site productivity provides the first step toward mapping canopy δ15N for these managed forests with remote sensing. [ABSTRACT FROM AUTHOR]

Abstract Conifers often occur along steep gradients of diverse climates throughout their natural ranges, which is expected to result in spatially varying selection to local climate conditions. However, signals of climatic adaptation can often be confounded, because unraveled clines covary with signals caused by neutral evolutionary processes such as gene flow and genetic drift. Consequently, our understanding of how selection and gene flow have shaped phenotypic and genotypic differentiation in trees is still limited. A 40‐year‐old common garden experiment comprising 16 Douglas‐fir (Pseudotsuga menziesii) provenances from a north‐to‐south gradient of approx. 1,000 km was analyzed, and genomic information was obtained from exome capture, which resulted in an initial genomic dataset of >90,000 single nucleotide polymorphisms. We used a restrictive and conservative filtering approach, which permitted us to include only SNPs and individuals in environmental association analysis (EAA) that were free of potentially confounding effects (LD, relatedness among trees, heterozygosity deficiency, and deviations from Hardy–Weinberg proportions). We used four conceptually different genome scan methods based on FST outlier detection and gene–environment association in order to disentangle truly adaptive SNPs from neutral SNPs. We found that a relatively small proportion of the exome showed a truly adaptive signal (0.01%–0.17%) when population substructuring and multiple testing was accounted for. Nevertheless, the unraveled SNP candidates showed significant relationships with climate at provenance origins, which strongly suggests that they have featured adaptation in Douglas‐fir along a climatic gradient. Two SNPs were independently found by three of the employed algorithms, and one of them is in close proximity to an annotated gene involved in circadian clock control and photoperiodism as was similarly found in Populus balsamifera. Synthesis. We conclude that despite neutral evolutionary processes, phenotypic and genomic signals of adaptation to climate are responsible for differentiation, which in particular explain disparity between the well‐known coastal and interior varieties of Douglas‐fir.

Douglas-fir (Pseudotsuga menziesii (Mirbel) Franco), an important North American softwood species, has been cultivated in Europe since 1827 and extended over a large area (831 thousand hectares) due to its (i) high growth and wood production, (ii) tolerance to summer drought, as well as (iii) wood with superior qualities and multiple uses. Since the natural pruning of Douglas-fir is defective/imperfect and slow, even in dense stands, intervention with artificial pruning is required, in stands intended for the production of superior wood assortments (e.g., veneer, solid wood furniture, class A lumber). In this context, the paper presents the results of an artificial pruning experiment in Douglas-fir, which sought to determine the duration of pruning and the costs related to the intervention at heights of a maximum of 2 m (penetration/access pruning) and over 2 m (high pruning). We worked in two stands with Douglasfir (one mixed, the other monospecific), in two teams of two people for each stand. 104 trees were pruned, of which all up to a height of 2 m and 28 up to a maximum height of 4.4 m, of which 75% were pruned to a minimum of 4 m. Average duration per meter of pruning from 0 to 2 m height ranged, in the four teams, between 0'46" and 1'5", a value that increased to 1'36" in the case of intervention at heights over 2 m. Based on these values, it can be considered that the duration of the artificial pruning of Douglas-fir at a height of 6 m reaches values of at least 7'30"-8', similar to the situation in Germany. If we consider the artificial pruning of 360 future crop trees ha-1 (the maximum number proposed in Romania for Douglas-fir stands) up to 6 m high, results a current pruning cost of 1,400 RON ha-1 (284 euro ha-1). This relatively low value economically justifies the application of artificial pruning to Douglas-fir if the production of high-quality wood with superior uses is targeted. [ABSTRACT FROM AUTHOR]

The USDA Forest Service is working closely with collaborative stakeholder groups to accelerate the pace and scale of restoration in fire-prone mixed conifer forests of eastern Oregon. Collaboratively planned restoration projects are typically designed to conserve older trees established before fire exclusion policies and other management practices began to alter forest landscapes beginning in the late 1800s. Tools exist for accurately estimating the age of common species including ponderosa pine, Douglas-fir, and western larch. There are no existing tools available for aging grand/white fir, although an important objective of many restoration projects in mixed conifer stands is to retain older grand/white fir while removing younger individuals that have infilled into stands in the absence of fire to enhance resilience of stands to future climate and disturbance regimes. This article describes the development of tools to age grand fir on the Malheur National Forest by taking simple field measurements of morphological characteristics. Bark fissure depth, height to live foliage, and diameter at breast height were the strongest noncorrelated tree morphological characteristics associated with tree age. Crown class had no predictive power for estimating tree age. A variety of methods are presented that can estimate the age of grand fir with reasonable accuracy and are appropriate for different management objectives. Additional field testing and continued experimentation with different tree aging methods within an adaptive management framework is recommended. Study Implications: Shade tolerant grand fir has expanded dramatically in mixed conifer stands of eastern Oregon in the absence of frequent fire. Collaboratively designed restoration projects in mixed conifer stands usually call for the removal of younger grand fir while maintaining older grand fir that contribute to stand- and landscape-scale biodiversity. It can be difficult to estimate the age of grand fir based on morphological clues because of highly variable growth forms in this species. However, several easy-to-use grand fir aging tools promise to facilitate restoration by making reasonably accurate estimates of tree age. [ABSTRACT FROM AUTHOR]

Outbreaks of western spruce budworm (Choristoneura freemani Freeman) in Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] forests of western Canada have shifted northward in recent decades. Using forest health survey data collected during the previous century (1909 to 2011) to assess the pattern of range shifts, we found that outbreaks have rapidly shifted to higher latitudes and elevations while gradually retracting from the southern and western portions of its range. Given that the survival of western spruce budworm is affected by access to developing buds, we tested the prediction that this shift was associated with climate change-induced alterations in synchrony between budworm larval emergence and Douglas-fir bud development. We simulated the dates of larval emergence and host budburst annually from 1901 to 2011 using existing phenology models and long-term weather data. Over the last century, the timing of larval emergence in relation to budburst has converged toward optimal synchrony at higher latitudes and elevations, while diverging at lower latitudes and elevations. Shifting synchrony arises from differential responses to temperature by the insect and its host tree; both are reliant upon degree day accumulation, but Douglas-fir bud development is putatively dependent upon an additional photoperiodic threshold. Thus, the potential for warming to cause directional shifts in phenological matching between early season folivores and their host trees may be contingent upon the degree to which each depends on different components of their environments for their development.

Key message: Planting sites that are cooler, with more precipitation, early springs, high TD and hot dry summers reduce bole taper in young Douglas-fir trees. Seed-source climates that are drier, with high TD and low snowfall produce seed-sources with lower taper. Analysis of a 10 year reciprocal transplant study was performed to determine the influence of seed-source and local planting site climates on the bole taper of 8995 Douglas-fir (Pseudotsuga menziesii var. menziesii) trees. Trees were planted at 9 sites across Oregon and Washington, with 120 known families taken from 12 seed-source regions across California, Oregon, and Washington. Diameter at breast height (DBH) ranged from 0.1 to 22.9 cm, with tree ages ranging from 2 year-old seedlings to 12 year-old trees. Changes in Gini coefficients (∆G) of diameters along tree boles, as a surrogate for changes in taper, were modeled as a function of age, site climate, and seed-source climate (Wang et al., in PLoS One 11, 2016) using universal response functions (URF) (Wang et al., in Ecol Appl 20:153–163, 2010). Lower Gini coefficients come from less tapered boles, i.e., more cylindrically shaped boles. There was significant influence on taper from five site climate variables: mean annual temperature (MAT), mean annual precipitation (MAP), beginning of frost-free period (bFFP), difference between minimum and maximum monthly temperatures (TD), and summer heat-moisture index (SHM). These results suggest that cooler years with more precipitation, early springs, large TD and hot dry summers reduce tree taper. There was significant influence on taper from three seed-source climate variables: percent precipitation as snow (PAS), Hargrave's climate moisture deficit (CMD), and TD. These results suggest dry regions with large ranges in monthly temperatures and low snowfall will produce seed-sources with lower Gini coefficients. Projections under future climates using an ensemble model show areas where G estimates are currently highest showed increases in G, while areas with the lowest G estimates decreased. Most of the Douglas-fir region shows declines in Gini coefficients under high emissions scenarios. These predicted changes in taper have direct implications for wood volume and carbon mass estimates of trees under future climates. [ABSTRACT FROM AUTHOR]

Bark beetles are casual agents of tree mortality, often affecting landscapes with diverse topography. The Douglas-fir beetle (Dendroctonus pseudotsugae Hopkins) (DFB) uses Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco), as its primary host. Populations often increase following disturbances such as defoliation, fires, or blowdowns and then transition into live healthy trees. Mitigating tree mortality where high mortality levels are incompatible with land manager objectives is of relevance. In this study, we compared mortality levels in eighteen thinned and twelve unthinned pure Douglas-fir stands in the Helena National Forest in Montana. Managed stands were thinned during the summers of 2014 and 2015 and tree mortality evaluated in 2017 and 2018. Douglas-fir beetle populations were at endemic levels in the study area, as represented by low observed tree mortality. Even under endemic populations, 3.6% of Douglas-firs > 12 in diameter at breast height (dbh) were killed in unthinned stands compared to 1.6% in thinned stands. Basal area of trees killed in unthinned stands was 5.4 (± 2.4) ft2/ac compared with 0.5 (± 0.3) ft2/ac in thinned stands. Although insect pressure was low, this study indicates that vegetation management could be a tool to mitigate DFB-caused tree mortality. Additional studies conducted under higher insect populations are needed. Study Implications: Douglas-fir beetle (DFB) can cause extensive mortality of Douglas-fir, particularly in dense stands of large-diameter trees with a high percentage of hosts. Vegetation management through silvicultural thinnings has been proposed as a long-term strategy to mitigate tree mortality from bark beetles. In this study, we compared tree mortality caused by DFB in thinned and unthinned Douglas-fir stands. Douglas-fir beetle populations were at low levels post-thinning and mortality levels were limited. Regardless, we observed increased mortality in denser, unthinned stands compared with thinned stands with lower tree density, suggesting that stand density reductions may mitigate DFB-caused tree mortality. [ABSTRACT FROM AUTHOR]

Douglas‐fir (Pseudotsuga menziesii) is an important component of mixed‐conifer forests of the Western United States. Changes in forest composition from Douglas‐fir encroachment in areas of fire‐suppression could have implications for water balance, supply and stream habitats. Increasing episodes of severe drought linked to climate change threaten forest health. Measurements of tree water inform management and biosphere models. Twenty‐six trees were instrumented with thermal dissipation sapflow probes. Sites were in Petrolia, California, and Maple Creek, California. Monitoring took place from Summer 2015 through Summer 2016. Daily water use was found to decline steeply over each summer dry period. This may limit the influence of Douglas‐fir on soil moisture budgets and streamflow during this time period. Tree size, peak and low flow daily water use averages were statistically related. Little difference was found between drier 2015 and wetter 2016 years. However, larger trees showed greater peak water use in 2015, perhaps from sufficient moisture and greater insolation, and at Petrolia, larger trees showed greater dry period water use in 2016, as greater moisture levels persisted into the summer. Linear mixed effects models of daily integral sap velocity (cm/day) were created with the input variables of solar radiation, vapour pressure deficit (VPD) and soil moisture. The best model using Akaike information criterion scores had fixed effects of solar radiation and VPD. This model was trained on one site and then validated at the second site with goodness of fit tests. The model is provided for estimating Douglas‐fir water use. [ABSTRACT FROM AUTHOR]

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.

A growing body of evidence suggests that climate change is altering the epidemiology of many forest diseases. Nothophaeocryptopus gaeumannii (Rhode) Petrak, an ascomycete native to the Pacific Northwest and the causal agent of the Swiss needle cast (SNC) disease of Douglas-fir [Pseudotsuga menziesii (Mirbel) Franco], is no exception. In the past few decades, changing climatic conditions have coincided with periodic epidemics of SNC in coastal forests and plantations from Southwestern British Columbia (B.C.) to Southwestern Oregon, wherein an increase in the colonization of needles by N. gaeumanii causes carbon starvation, premature needle shedding and a decline in growth. Two major sympatric genetic lineages of N. gaeumannii have been identified in the coastal Pacific Northwest. Past research on these lineages suggests they have different environmental tolerance ranges and may be responsible for some variability in disease severity. In this study, we examined the complex dynamics between biologically pertinent short- and long-term climatic and environmental factors, phylogenetic lineages of N. gaeumannii and the severity patterns of the SNC disease. Firstly, using an ensemble species distribution modeling approach using genetic lineage presences as model inputs, we predicted the probability of occurrence of each lineage throughout the native range of Douglas-fir in the present as well as in 2050 under the “business as usual” (RCP8.5) emissions scenario. Subsequently, we combined these model outputs with short-term climatic and topographic variables and colonization index measurements from monitoring networks across the SNC epidemic area to infer the impacts of climate change on the SNC epidemic. Our results suggest that the current environmental tolerance range of lineage 1 exceeds that of lineage 2, and we expect lineage 1 to expand inland in Washington and Oregon, while we expect lineage 2 will remain relatively constrained to its current range with some slight increases in suitability, particularly in coastal Washington and Oregon. We also found that disease colonization index is associated with the climatic suitability of lineage 1, and that the suitability of the different lineages could impact the vertical patterns of colonization within the crown. We conclude that unabated climate change could cause the SNC epidemic to intensify.

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