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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.

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]

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]

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.

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]

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]

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.

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]

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.)

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]

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.

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.

Large trees have disproportionally large competitive advantage in access to light, which has been proposed to increase growth dominance of large trees (e.g., a larger proportion of volume growth than standing volume is in large trees in a stand). Tree growth may also be limited by the availability of other resources besides light. Nutrient deficiency, especially of nitrogen, is considered common among temperate forests including mixedconifer forests of the Inland Northwest of the United States. Data from a long-term forest nutrition study across four Inland Northwest states were analyzed to evaluate nitrogen ( potassium fertilization's effects on growth dominance over an eight-year period following treatment. Our results show that growth dominance varied substantially across plots in each vegetation series and fertilization treatment, and its mean values generally were at minuscule magnitudes, negative, and not significantly different from 0. We propose that this lack of a clear pattern in growth dominance was the result of the mixed-species composition where shade-tolerant species remained in lower crown positions, yet their relative growth kept pace with large trees. Limited moisture availability at dry sites may have hampered the development of growth dominance. Growth dominance also was lowered by mortality observed among relatively large trees. The largely negative growth dominance across fertilization treatments indicated that small trees seem to have maintained higher relative growth rates than large trees, even if absolute growth was higher among large trees. In the case fertilization does improve stand growth, a significant part of this improved growth will be lost in density-dependent mortality over time if not captured through thinning/harvesting. [ABSTRACT FROM AUTHOR]

In the Douglas-fir (Pseudotsuga menziesii Mirb. Franco) and redwood (Sequoia sempervirens (D. Don) Endl.) forests of the central California coast, Sudden Oak Death (SOD) has led to landscape-scale mortality of tanoak (Notholithocarpus densiflorus (Hook. and Arn.) Manos, Cannon and S.H. Oh). As tanoak mortality progresses, fuel loads and potential fire behavior in these forests are changing. We documented increases in fuel loads over time in long-term monitoring plots in infested forests at Point Reyes National Seashore. Throughout the study, we observed a significant positive relationship between dead tanoak basal area and surface fuels. We used the fire behavior modeling program BehavePlus to compare potential fire behavior between diseased and healthy stands. Model outputs indicated the potential for longer flame lengths, higher rates of spread and more intense surface fire in diseased stands. The potential for increased fire intensity in diseased redwood and Douglas-fir forests may create additional challenges for fire and natural resources managers and may affect the ecology of these forests into the future.

In the Douglas-fir (Pseudotsuga menziesii Mirb. Franco) and redwood (Sequoia sempervirens (D. Don) Endl.) forests of the central California coast, Sudden Oak Death (SOD) has led to landscape-scale mortality of tanoak (Notholithocarpus densiflorus (Hook. and Arn.) Manos, Cannon and S.H. Oh). As tanoak mortality progresses, fuel loads and potential fire behavior in these forests are changing. We documented increases in fuel loads over time in long-term monitoring plots in infested forests at Point Reyes National Seashore. Throughout the study, we observed a significant positive relationship between dead tanoak basal area and surface fuels. We used the fire behavior modeling program BehavePlus to compare potential fire behavior between diseased and healthy stands. Model outputs indicated the potential for longer flame lengths, higher rates of spread and more intense surface fire in diseased stands. The potential for increased fire intensity in diseased redwood and Douglas-fir forests may create additional challenges for fire and natural resources managers and may affect the ecology of these forests into the future.

One mechanism of invasive species success is the production of allelopathic chemicals that negatively affect native competitors. A highly invasive shrub, Cytisus scoparius, impedes Douglas-fir tree establishment in clearcuts, even years after its removal. This impediment may be from the allelopathic alkaloids of C. scoparius that could indirectly hinder Douglas-fir by inhibiting their mutualistic ectomycorrhizal fungi (EMF). I extracted and quantified alkaloids from C. scoparius tissue for use in a laboratory bioassay. I then tested if and how these alkaloids affected EMF growth. In a second assay, I tested the effects of three concentrations of pure sparteine, the primary alkaloid in C. scoparius , on fungal growth. Sparteine was the only alkaloid recovered from the extraction which yielded 0.32 mg sparteine/g fresh weight, a lower concentration than previously reported values. Both the crude extract and pure sparteine significantly affected fungal growth, but only sparteine produced a species-specific response. Growth was inhibited by increasing sparteine concentrations, and most species were inhibited at 1.4 mM, the concentration found in C. scoparius. One common EMF, Wilcoxina mikolae, was unaffected by sparteine while others, like Suillus caerulescens and Cenococcum geophilum, were more sensitive and stopped growing entirely at 5 to 10 mM. These results suggest that the alkaloids of C. scoparius may seriously hinder EMF, and indirectly Douglas-fir, contributing to the competitive dominance of the invasive shrub.

Many fungi in the Ophiostomatales are vectored by bark beetles that introduce these fungi directly into their tree hosts. Most of these fungal associates have little effect on their hosts, but some can cause serious diseases. One such fungus, Leptographium wageneri, causes an economically and ecologically important tree disease known as black stain root disease (BSRD). For this study, 159 full genome sequences of L. wageneri were analyzed using a population genomics approach to investigate the epidemiology, dispersal capabilities, and reproductive biology of this fungus. Analyses were performed with SNP haplotypes from 155 isolates of L. wageneri var. pseudotsugae collected in 16 Douglas-fir stands in Oregon and 4 isolates of L. wageneri var. wageneri collected in pinyon pine stands in southern California. These two host-specific varieties appear to be evolutionarily divergent, likely due a combination of factors such as host differentiation and geographic isolation. We analyzed gene flow and population structure within and among Douglas-fir plantations in western Oregon to infer the relative importance of local vs. long distance dispersal in structuring populations of L. wageneri var. pseudotsugae. Long-distance gene flow has occurred between Douglas-fir plantations, contributing to diversity and population structure within stands, and likely reflecting the behavior of an important insect vector. Genetic clustering analyses revealed the presence of unique local clusters within stands and plantations in addition to those common among multiple stands or plantations. Although populations of L. wageneri var. pseudotsugae are primarily asexual, two mating types were present in many stands, suggesting that recombination is at least possible and may contribute to genetic diversity.

Abstract Douglas‐fir (Pseudotsuga menziesii) is one of the world's premier lumber species and somatic embryogenesis (SE) is the most promising method for rapid propagation of superior tree genotypes. The development and optimization of SE protocols in conifers is hindered by a lack of knowledge of the molecular basis of embryogenesis and limited sequence data. In Arabidopsis, the LEAFY COTYLEDON1 (AtLEC1) gene is a master regulator of embryogenesis that induces SE when expressed ectopically. We isolated the LEC1 homologue from Douglas‐fir, designated as PmLEC1. PmLEC1 expression in somatic embryos and developing seeds demonstrated a unique, alternating pattern of expression with the highest levels during early stages of embryogenesis. PmLEC1 protein accumulation during seed development correlated with its transcriptional levels during early embryogenesis; however, substantial protein levels persisted until 2 weeks on germination medium. Treatment of mature, stratified seeds with 2,4‐epibrassinolide, sorbitol, mannitol, or NaCl upregulated PmLEC1 expression, which may provide strategies to induce SE from mature tissues. Sequence analysis of the PmLEC1 gene revealed a 5′ UTR intron containing binding sites for transcription factors (TFs), such as ABI3, LEC2, FUS3, and AGL15, which are critical regulators of embryogenesis in angiosperms. Regulatory elements for these and other seed‐specific TFs and biotic and abiotic signals were identified within the PmLEC1 locus. Most importantly, functional analysis of PmLEC1 showed that it rescued the Arabidopsis lec1‐1 null mutant and, in the T2 generation, led to the development of embryo‐like structures, indicating a key role of PmLEC1 in the regulation of embryogenesis.

In the 2017 and 2018, 2.55 million hectares burned across British Columbia, Canada, including unanticipated large and high-severity fires in many dry forests. To transform forest and fire management to achieve resilience to future megafires requires improved understanding historical fire frequency, severity, and spatial patterns. Our dendroecological reconstructions of 35 plots in a 161-hectare study area in a dry Douglas-fir forest revealed historical fires that burned at a wide range of frequencies and severities at both the plot- and study-area scales. The 23 fires between 1619 and 1943 burned at intervals of 10–30 years, primarily at low- to moderate-severity that scarred trees but generated few cohorts. In contrast, current fire-free intervals of 70–180 years exceed historical maximum intervals. Of the six widespread fires from 1790 to 1905, the 1863 fire affected 86% of plots and was moderate in severity with patches of higher severity that generated cohorts at fine scales only. These results indicate the severity of fires varied at fine spatial scales, and offer little support for the common assertion that periodic, high-severity, stand-initiating events were a component of the mixed-severity fire regime in these forest types. Many studies consider fires in the late 1800s relatively severe because they generated new cohorts of trees, and thus, emphasize the importance of high-severity fires in a mixed-severity fire regime. In our study area, the most widespread and severe fire was not a stand-initiating fire. Rather, the post-1863 cohorts persisted due disruption of the fire regime in the twentieth century when land-use shifted from Indigenous fire stewardship and early European settler fires to fire exclusion and suppression. In absence of low- to moderate-severity fires, contemporary forests are dense with closed canopies that are vulnerable to high-severity fire. Future management should reduce forest densities and to restore stand- and landscape-level heterogeneity and increase forest resilience. The timing and size of repeat treatments such as thinning of subcanopy trees and prescribed burning, including Indigenous fire stewardship, can be guided by our refined understanding of the mixed-severity fire regime that was historically dominated by low- to moderate-severity fires in this dry forest ecosystem.

Forests are important for biodiversity, timber production and carbon accumulation, but these ecosystem services may be impacted by climate change. Field data collected from individual forest types occurring across a climatic gradient can contribute to forecasting these consequences. We examined how changes in temperature, precipitation and aridity affect ecosystem services in 23 mature Douglas-fir (Pseudotsuga menziesii) forests in nine climatic regions across a 900 km gradient in British Columbia, Canada. Using Canadian National Forest Inventory methodology, we assessed richness and diversity of plant functional groups, site index, and above- and below-ground carbon stocks. As aridity increased, ecosystem-level tree species richness declined on average from four to one species, Douglas-fir site index declined from 30 to 15 m, and ecosystem carbon storage decreased from 565 to 222 Mg ha–1. Tree species richness was positively and herb species richness negatively correlated with carbon storage. Carbon storage by ecosystem compartment was largest in aboveground live tree biomass, declining in the following order: mineral soils > coarse woody debris and dead standing trees > forest floor > small and fine woody debris > understory plants. Mineral soil carbon at depths of 0-15 cm, 15-35 cm, and 35-55 cm increased with increasing mean annual precipitation and decreasing aridity. Our results indicate that as aridity increases and precipitation decreases, tree species richness, site index and carbon storage in existing Douglas-fir forests declines. However, assisted or natural migration of Douglas-fir into more humid regions could be associated with more diverse, productive, carbon-rich forests. This study informs carbon stock vulnerability and provides empirical data essential for carbon stock forecasts.

Source terms of the number of firebrands released as trees/shrubs torch are essential for helping to more accurately implement detailed physics-based models of wildfire propagation, particularly when torching occurs. A key challenge in estimating these source terms is the difficulty in measuring firebrand generation rates from the source. Typical studies have reported generation characteristics (firebrands/m2), but enumeration of the total number of firebrands released has remained elusive. Recently, a fire-resistant fabric was successfully employed to quantify “hot” firebrands (i.e., can potentially lead to further fire spread) generated during tree-scale burns. In this paper, a total of 71 tree/shrubs were burned to quantify hot firebrand production statistics for Douglas-fir, ponderosa pine, and sagebrush. A network of 65 fire-resistant fabric stations were deployed during the burns to measure the “hot” firebrands that are released. These stations are placed strategically at several radii and angles in the prominent wind direction. Using the number of firebrands determined from the fabric, a first-order extrapolation method was developed and evaluated to estimate the source terms (i.e., total number of hot firebrands released). Variation in the firebrand production with the height of the trees and for three species is reported. The total number of hot firebrands produced typically increased with the height of the tree/shrub burned as expected. The specific hot firebrand production (firebrands produced per kg dry mass burned) is exponentially dependent on the moisture content of the tree and had an inconclusive correlation with the height of the trees. Overall, for trees of comparable moisture content, sagebrush produced higher specific hot firebrands when compared to ponderosa pine and Douglas-fir species.

Douglas‐fir (Pseudotsuga menziesii) is one of the world's premier lumber species and somatic embryogenesis (SE) is the most promising method for rapid propagation of superior tree genotypes. The development and optimization of SE protocols in conifers is hindered by a lack of knowledge of the molecular basis of embryogenesis and limited sequence data. In Arabidopsis, the LEAFY COTYLEDON1 (AtLEC1) gene is a master regulator of embryogenesis that induces SE when expressed ectopically. We isolated the LEC1 homologue from Douglas‐fir, designated as PmLEC1. PmLEC1 expression in somatic embryos and developing seeds demonstrated a unique, alternating pattern of expression with the highest levels during early stages of embryogenesis. PmLEC1 protein accumulation during seed development correlated with its transcriptional levels during early embryogenesis; however, substantial protein levels persisted until 2 weeks on germination medium. Treatment of mature, stratified seeds with 2,4‐epibrassinolide, sorbitol, mannitol, or NaCl upregulated PmLEC1 expression, which may provide strategies to induce SE from mature tissues. Sequence analysis of the PmLEC1 gene revealed a 5′ UTR intron containing binding sites for transcription factors (TFs), such as ABI3, LEC2, FUS3, and AGL15, which are critical regulators of embryogenesis in angiosperms. Regulatory elements for these and other seed‐specific TFs and biotic and abiotic signals were identified within the PmLEC1 locus. Most importantly, functional analysis of PmLEC1 showed that it rescued the Arabidopsis lec1‐1 null mutant and, in the T2 generation, led to the development of embryo‐like structures, indicating a key role of PmLEC1 in the regulation of embryogenesis. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Sustainable forest management strategies include emulating historical disturbance regimes to achieve multiple objectives. Partial-harvesting strategies are used to overcome conflicts between timber production and wildlife habitat conservation; however, the potential impacts on complex disturbance interactions and ecological functions remain largely unknown. In 1984, a controlled experiment was initiated in the dry forests of central British Columbia, Canada, to test partial harvesting intended to enhance mule deer habitat while allowing timber extraction. To determine the short- and long-term impacts on complex disturbance regimes, we quantified changes in forest structure and susceptibility to western spruce budworm, Douglas-fir beetle, and wildfire. We compared structural attributes in 2014 (30 years after the first harvest) and 2015 (1 year after the second harvest) in treated forests, and contrasted them with control forests that were measured in 2015. In the short term (1 year post-harvest), partial harvesting altered forest structure by reducing total canopy cover, subcanopy tree density and basal area, and increasing the abundance of large woody surface fuels. In the long term (30 years post-harvest), the forest canopy attributes did not differ between the treatment and control areas, partly due to increased growth of subcanopy trees. Harvesting had little impact on forest susceptibility to western spruce budworm. Susceptibility to Douglas-fir beetle was lower in the short term due to fewer available mature host trees, but increased to levels similar to the control forest over the long term. Reduced canopy fuels and increased canopy base height decreased the likelihood of crown fire in favor of surface fire. In the long term, canopy fuels and likelihood of crown fire recovered, but woody fuel loads remained low after 30 years. Harvesting to enhance mule deer habitat interacts with biotic and abiotic disturbances in the short and long term. Potential cascading affects depended more on the decision to remove harvesting residuals to mitigate potential Douglas-fir beetle infestations and wildfire than on time since treatment. Provided partial harvesting occurs at intervals ≤ 30 years and residuals are immediately removed, timber extraction and mule deer habitat can be compatible with complex disturbance regimes and sustainable forest management.

Abstract The U.S. Forest Service is reconsidering policies that limit the size of trees that can be removed in the course of restoration treatments in dry forests of eastern Oregon. To evaluate the effects of diameter limits on the ability of managers to meet restoration objectives, we used an existing network of long‐term research plots to summarize historical and contemporary structure and composition of mixed‐conifer forests within a one million‐ha study area in eastern Oregon. Then, we used a novel thinning simulation procedure to quantify the degree to which thinning using different diameter limits restored stands to historical conditions. Contemporary mixed‐conifer forests within the study area are significantly denser, have more basal area, and have a greater proportion of shade‐tolerant species than historical conditions. Our simulations of thinning under current policy that prohibits cutting of trees ≥53 cm show that a quarter of mixed‐conifer stands cannot be restored to within the historical range of basal area or density. Those stands that could be restored to within historical basal area ranges still had a substantially higher component of shade‐tolerant trees than historical stands. Permitting larger shade‐tolerant trees to be removed allowed restoration of all or most of stands to within historical structural and compositional ranges. Forest conditions in the late 1800s may not necessarily provide the best template for management because climate and disturbance projections suggest that eastern Oregon forests will be less well suited to shade‐tolerant species in the future. Adapting stands to future conditions will require robust monitoring of forest structural and compositional response to restoration treatments.

Herbivory by black-tailed deer affects growth form and survival of conifer seedlings in western Oregon and Washington, especially Douglas-fir. Several deterrents have been employed to reduce damage to forest resources yet most are ineffective or cost prohibitive. Use of chemical repellents is socially appealing because they offer a potential non-lethal alternative to reduce plant damage (i.e., browse). Commercial deer repellents may employ one or more mechanisms in their formulation, which affect taste, odor, visual, and/or tactile cues when consumed by deer. I evaluated the commercial SeaDust Wildlife Controllant™ as a tool to repel black-tailed deer from Douglas-fir seedlings during spring bud burst in western Oregon (mid-May through early July), because its ingredients have the potential to employ multiple avoidance mechanisms targeted on tactile, taste, and odor cues. Evidence indicated that deer browse was affected by an interaction of treatment and site; therefore, sites were evaluated separately. Treated seedlings were browsed less by deer than untreated seedlings on 2 of 3 tree farms where percent browse ranged from 0.08%-0.17% in treated plots and 0.15%-0.37% in control plots. Browse was similar at the third tree farm (0.15%). However, number of seedlings browsed in the third tree farm was greater than one site and less than the other, suggesting that statistical significance may not represent biological significance in this study. Future research is needed to incorporate acceptable loss to browsing and cost:benefit analyses.

BackgroundThe development of affordable woody biomass feedstocks represents a significant opportunity in the development of cellulosic biofuels. Primary woodchips produced by forest mills are considered an ideal feedstock, but the prices they command on the market are currently too expensive for biorefineries. In comparison, forestry residues represent a potential low-cost input but are considered a more challenging feedstock for sugar production due to complexities in composition and potential contamination arising from soil that may be present. We compare the sugar yields, changes in composition in Douglas-fir woodchips and forestry residues after pretreatment using ionic liquids and enzymatic saccharification in order to determine if this approach can efficiently liberate fermentable sugars.ResultsThese samples were either mechanically milled through a 2 mm mesh or pretreated as received with the ionic liquid (IL) 1-ethyl-3-methylimidazolium acetate [C2mim][OAc] at 120°C and 160°C. IL pretreatment of Douglas-fir woodchips and forestry residues resulted in approximately 71-92% glucose yields after enzymatic saccharification. X-ray diffraction (XRD) showed that the pretreated cellulose was less crystalline after IL pretreatment as compared to untreated control samples. Two-dimensional nuclear magnetic resonance spectroscopy (2D-NMR) revealed changes in lignin and hemicellulose structure and composition as a function of pretreatment. Mass balances of sugar and lignin streams for both the Douglas-fir woodchips and forestry residues throughout the pretreatment and enzymatic saccharification processes are presented.ConclusionsWhile the highest sugar yields were observed with the Douglas-fir woodchips, reasonably high sugar yields were obtained from forestry residues after ionic liquid pretreatment. Structural changes to lignin, cellulose and hemicellulose in the woodchips and forestry residues of Douglas-fir after [C2mim][OAc] pretreatment are analyzed by XRD and 2D-NMR, and indicate that significant changes occurred. Irrespective of the particle sizes used in this study, ionic liquid pretreatment successfully allowed high glucose yields after enzymatic saccharification. These results indicate that forestry residues may be a more viable feedstock than previously thought for the production of biofuels.

The U.S. Forest Service is reconsidering policies that limit the size of trees that can be removed in the course of restoration treatments in dry forests of eastern Oregon. To evaluate the effects of diameter limits on the ability of managers to meet restoration objectives, we used an existing network of long‐term research plots to summarize historical and contemporary structure and composition of mixed‐conifer forests within a one million‐ha study area in eastern Oregon. Then, we used a novel thinning simulation procedure to quantify the degree to which thinning using different diameter limits restored stands to historical conditions. Contemporary mixed‐conifer forests within the study area are significantly denser, have more basal area, and have a greater proportion of shade‐tolerant species than historical conditions. Our simulations of thinning under current policy that prohibits cutting of trees ≥53 cm show that a quarter of mixed‐conifer stands cannot be restored to within the historical range of basal area or density. Those stands that could be restored to within historical basal area ranges still had a substantially higher component of shade‐tolerant trees than historical stands. Permitting larger shade‐tolerant trees to be removed allowed restoration of all or most of stands to within historical structural and compositional ranges. Forest conditions in the late 1800s may not necessarily provide the best template for management because climate and disturbance projections suggest that eastern Oregon forests will be less well suited to shade‐tolerant species in the future. Adapting stands to future conditions will require robust monitoring of forest structural and compositional response to restoration treatments. [ABSTRACT FROM AUTHOR]

Clearcutting reduces proximity to seed sources and mycorrhizal inoculum potential for regenerating seedlings. Partial retention of legacy trees and protection of refuge plants, as well as preservation of the forest floor, can maintain mycorrhizal networks that colonize germinants and improve nutrient supply. However, little is known of overstory retention levels that best protect mycorrhizal inoculum while also providing sufficient light and soil resources for seedling establishment. To quantify the effect of tree retention on seedling regeneration, refuge plants, and resource availability, we compared five harvesting methods with increasing retention of overstory trees (clearcutting (0% retention), seed tree (10% retention), 30% patch retention, 60% patch retention, and 100% retention in uncut controls) in an interior Douglas-fir-dominated forest in British Columbia. Regeneration increased with proximity to legacy trees in partially cut forests, with increasing densities of interior Douglas-fir, western redcedar, grand fir, and western hemlock seedlings with overstory tree retention. Clearcutting reduced cover of ectomycorrhizal refuge plants (from 80 to 5%) while promoting arbuscular mycorrhizal plants the year after harvest. Richness of shrubs, herbs, and mosses declined with increasing harvesting intensity, but tree richness remained at control levels. The presence of legacy trees in all partially cut treatments mitigated these losses. Light availability declined with increasing overstory cover and proximity to leave trees, but it still exceeded 1,000 W m−2 in the clearcut, seed tree and 30% retention treatments. Increasing harvesting intensity reduced aboveground and belowground C stocks, particularly in live trees and the forest floor, although forest floor losses were also substantial where thinning took place in the 60% retention treatment. The loss of forest floor carbon, along with understory plant richness with intense harvesting was likely associated with a loss of ectomycorrhizal inoculum potential. This study suggests that dispersed retention of overstory trees where seed trees are spaced ~10–20 m apart, and aggregated retention where openings are