Sugar Pine Research Articles

Testing source elevation versus genotype as predictors of sugar pine performance in a post‐fire restoration planting

AbstractClimate change is motivating a reassessment of how seeds are selected for reforestation, as rapid environmental change can lead to local maladaptation in trees. Genetic association studies and past seed source climate both have the potential to help identify appropriate planting stock, but these techniques have not been compared and tested as part of an operational planting program. In this study, we combined an analysis of single nucleotide polymorphisms (SNPs) associated with environmental gradients in sugar pine (Pinus lambertiana) with an analysis of post‐fire seedling survival and growth in a restoration experiment. Our genotype–environment association (GEA) tests of 92 individuals from varying climates within CA revealed 829 SNPs (out of 300,604) with significant association with climate gradients, especially April snowpack. Of these, 323 either had annotations that suggested potential functional importance or were identified by two different methods. We then built Bayesian models of survival and growth for all seedlings in a separate post‐fire planting experiment, to test the relative predictive ability of source elevation (a common proxy for source climate) versus the proportion of seedling alleles expected to be locally advantageous based on GEA. Across three sites within the King Fire scar in Eldorado National Forest in 2017, 2018, and 2019, 1774 seedlings were planted. Of these, 206 had enough green needles in 2020 to allow sample collection, and 161 were successfully genotyped. We found that source elevation was generally better at predicting seedling performance than genotype indices, perhaps because of the limited scope of the association analysis. Seed sources from 500 to 1800 ft (152.4–548.6 m) lower in elevation, and one seed zone further south generally performed as well or better than local seed sources. This result, and those of similar previous studies, suggest that “climate matching” using past climate information for existing seed sourcing units is a reasonable starting point for finding seedlings suited to already‐altered planting site climate conditions. However, further tests with more extensive genomic and performance data may improve the utility of genotype information for seed selection.

Non-composted and blended agro-industrial by-products an as alternative soilless substrate affect potted zonal geranium (Pelargonium x hortorum) and murva (Murraya paniculata) production

Due to their hiking prices and import restrictions, peat moss and coco coir are not readily available in many developing countries including Pakistan. Therefore, it is necessary to explore alternative soilless substrates having low cost and comparable physico-chemical properties with peat moss and coco coir for local and global market. For this purpose, indigenous agro-industrial by-products, viz., sesame straw, date palm coir, sugarcane pressmud, sugar beet waste and pine bark were collected, cleaned, crushed (where needed) and blended for physico-chemical analyses. Zonal geranium ‘Pinto White’ and murva (Murraya paniculata Kaneh.) were used to evaluate the efficacy of various combinations of substrates on plant growth and production. Three experiments were conducted with six treatments of each experiment replicated three times. Blending of sesame straw with sugarcane pressmud and pine bark (4:4:2 by volume) yielded best results and marketable plants were produced with tallest height (35.8 cm), canopy diameter (20.8 cm), leaf area (29.2 cm2), leaf total chlorophyll contents (62.1 SPAD) and flowers per plant (5.0 No.) along with shortest production time (103 days). Same results were recorded when murva plants were grown in 4:4:2 blending of sesame straw, sugarcane pressmud and pine bark. Zonal geranium and murva plants did not perform well when sesame straw was blended with date palm coir and sugar beet waste. Blending of sesame straw with sugarcane pressmud and pine bark had low pH (7.0), electrical conductivity (1.37 mS cm-1), bulk density (0.29 g cm-3) while high water retention (34.7 %) as compared to other blended substrates. However, shrinkage of substrates in containers was highest (7.7 %) when sesame straw was blended with sugarcane pressmud and pine bark in 5:3:2 followed by 4:4:2. pH value of all blended substrates decreased at termination except when sesame straw, sugarcane pressmud and pine bark were blended in 4:4:2 and 5:2:3, which was increased from 7.0 to 7.1. Electrical conductivity of all blended substrates was decreased at termination of the experiments. It can be concluded that blending of sesame straw, sugarcane pressmud and pine bark in 4:4:2 (v/v/v) significantly improved physico-chemical characteristics of substrates as well as plant growth and production attributes of zonal geranium and murva. Therefore, this indigenous soilless substrate, which is cheaper and readily available to nurserymen can be used as alternative of peat moss and coco coir for containerized plant production.

Stand diversity increases pine resistance and resilience to compound disturbance

BackgroundDrought, fire, and insects are increasing mortality of pine species throughout the northern temperate zone as climate change progresses. Tree survival may be enhanced by forest diversity, with growth rates often higher in mixed stands, but whether tree defenses are likewise aided remains in question. We tested how forest diversity-productivity patterns relate to growth and defense over three centuries of climate change, competition, wildfire, and bark beetle attack. We used detailed census data from a fully mapped 25.6-ha forest dynamics plot in California, USA to conduct a spatially explicit, dendroecological assessment of large-diameter Pinus lambertiana survival following fire reintroduction. Our structural equation models investigated direct and indirect pathways by which growth, defense, and forest composition together mediated pine resistance and resilience.ResultsIn the historical era of frequent, mixed-severity fire (pre-1900), trees that were ultimately resistant or susceptible to the post-fire bark beetle epidemic all showed similar growth and defenses, as measured by axial resin duct traits. During the era of fire exclusion (1901–2012), however, susceptible trees had slower growth. Following fire re-entry in 2013, both growth and defense declined precipitously for susceptible trees, resulting in fatal bark beetle attack. Spatial analysis showed that monodominant crowding by shade-tolerant competitors contributed to the long-term stress that prevented susceptible trees from recuperating defenses quickly following fire re-entry. For beetle-resistant trees, however, we found positive feedbacks between diversity, growth, and survival: trees in species-rich communities had higher growth rates pre-fire, which promoted a rapid recuperation of defenses following fire that helped trees resist bark beetle attack. Overall, this associational resistance outweighed associational susceptibility (+8.6% vs. −6.4% change in individual tree survival odds), suggesting a relaxation effect that ultimately allowed 58% of large pines to survive.ConclusionsThough climate change threatens forest biodiversity, biodiversity is key to forest climate adaptation in return. Our findings demonstrate centennial-scale feedbacks by which forest diversity increases pine resistance and resilience to climate-amplified disturbances. The spatially explicit, dendroecological framework provides new insights into diversity-productivity theory, while also informing climate-adaptive forest management by identifying thresholds of tree density and richness that maximize large pine survival.

Placing 21st Century Warming in Southern California, USA in a Multi-Century Historical Context

Warming in southern California during the 21st century is unprecedented in the instrumental record. To place this warming in a multi-century historical context, we analyzed tree ring data sampled from Jeffrey pine (Pinus jeffreyi) and sugar pine (Pinus lambertiana) collected from minimally disturbed, old-growth high-elevation forests within Mt. San Jacinto State Park California, USA. Based on a calibration/verification period of 1960–2020 between earlywood radial growth and California Climate Division 6 climate data, we reconstructed annual (November–October) minimum temperature (Tmin) from 1658 to 2020. During the 61-year calibration/verification period, instrumental Tmin increased (r = 0.69, p < 0.01) and was positively associated with annual radial growth (r = 0.71, p < 0.01). Using regime shift analysis, we found that the 363-year reconstruction revealed Tmin stability until 1958 and then decreased until 1980, followed by the two warmest regimes (1981–2007, 2008–2020) on record. The last 13-year period was 0.77 °C warmer than the multi-century average with nine of the ten warmest years in the reconstruction recorded. These results suggest that 21st century warming in southern California is unique in the context of the past four centuries, indicating the rarity of exceptional warmth captured in the tree ring record.

Threats to the persistence of sugar pine (Pinus lambertiana) in the western USA

This study assesses how numerous stressors shape the vital rates (survival, growth, and fecundity) of sugar pine across the vast majority of its range. Sugar pine (Pinus lambertiana Dougl.) is the largest Pinus species, an important timber species, and a component of several dry conifer forest types of western North America, in particular the extensive Sierra Nevada mixed conifer forest. The species faces several challenges in the Anthropocene, including a disrupted fire regime, an invasive pathogen, forest structure changes, and drought with ensuing bark beetle epidemics. Managers are concerned about the conservation outlook for sugar pine, but it is unclear where and how to best invest conservation resources. Using data from the US Forest Service’s Forest Inventory and Analysis program, we estimate the parameters for the vital rate functions and use these to construct an integral projection model which predicts the effects of various stressors on the asymptotic population growth rate. The asymptotic population growth rate is near or slightly below one even under reference conditions, and the actual abundance (in terms of both stem density and basal area) slightly declined over the duration of the study (2001–2019). The analysis reveals that wildfire, white pine blister rust, and forest density are key drivers of the demographic rates of sugar pine across its range. Drought and site dryness had lesser, but still meaningful, effects. Fire has strong negative effects on survival, resulting in a strongly negative population trajectory on burned sites. Conversely, lower than average forest density (plot-level basal area) results in a positive population growth rate via beneficial effects on individual growth. These results highlight the value of fire hazard mitigation, particularly where it also reduces forest density, in the conservation of this important species.

Predicting snag fall in an old-growth forest after fire

BackgroundSnags, standing dead trees, are becoming more abundant in forests as tree mortality rates continue to increase due to fire, drought, and bark beetles. Snags provide habitat for birds and small mammals, and when they fall to the ground, the resulting logs provide additional wildlife habitat and affect nutrient cycling, fuel loads, and fire behavior. Predicting how long snags will remain standing after fire is essential for managing habitat, understanding chemical cycling in forests, and modeling forest succession and fuels. Few studies, however, have quantified how fire changes snag fall dynamics.ResultsWe compared post-fire fall rates of snags that existed pre-fire (n = 2013) and snags created during or after the fire (n = 8222), using 3 years of pre-fire and 5 years of post-fire data from an annually monitored, 25.6-ha spatially explicit plot in an old-growth Abies concolor–Pinus lambertiana forest in the Sierra Nevada, CA, USA. The plot burned at low to moderate severity in the Rim Fire of 2013. We used random forest models to (1) identify predictors of post-fire snag fall for pre-existing and new snags and (2) assess the influence of spatial neighborhood and local fire severity on snag fall after fire. Fall rates of pre-existing snags increased 3 years after fire. Five years after fire, pre-existing snags were twice as likely to fall as new snags. Pre-existing snags were most likely to persist 5 years after fire if they were > 50 cm in diameter, > 20 m tall, and charred on the bole to heights above 3.7 m. New snags were also more likely to persist 5 years after fire if they were > 20 m tall. Spatial neighborhood (e.g., tree density) and local fire severity (e.g., fire-caused crown injury) within 15 m of each snag barely improved predictions of snag fall after fire.ConclusionsLand managers should expect fall rates of pre-existing snags to exceed fall rates of new snags within 5 years after fire, an important habitat consideration because pre-existing snags represent a wider range of size and decay classes.

Thinning and prescribed burning increase shade-tolerant conifer regeneration in a fire excluded mixed-conifer forest

Fire exclusion and past management have altered the composition, structure, and function of frequent-fire forests throughout western North America. In mixed-conifer forests of the California Sierra Nevada, fire exclusion has exacerbated the effects of drought and endemic bark beetles, resulting in extensive mortality of fire-adapted pine species. Thinning and prescribed fire are widely used in these forests to reduce fuels, moderate fire behavior, and restore ecosystems. Tree regeneration influences future forest composition and structure, and therefore future resilience to disturbances, but long-term effects of thinning and prescribed burning on tree regeneration after prolonged fire exclusion are poorly understood. We measured tree regeneration one year prior to, and periodically for 16 years following thinning and prescribed burning in a mixed-conifer forest in the Sierra Nevada, California, USA. We asked three questions. How did the composition and density of tree regeneration change after thinning and prescribed burning? Did pretreatment vegetation types influence conifer regeneration density after treatments? Did planting after overstory thinning increase regeneration density of native pine species?Sixteen years after treatments, combined natural regeneration of shade-tolerant white fir (Abies concolor) and incense-cedar (Calocedrus decurrens) averaged 2,032 trees per hectare (tph) after understory thinning, and 7,745 tph after understory thinning combined with prescribed burning, increases of 37 % and 146 % from pretreatment densities. In contrast, combined natural regeneration of white fir and incense-cedar averaged 497 tph after overstory thinning, 780 tph after overstory thinning with prescribed burning, 113 tph after prescribed burning alone, and 807 tph in untreated controls, all of which were declines from pretreatment densities. Natural regeneration of white fir and incense-cedar was consistently an order of magnitude greater than Jeffrey pine (Pinus jeffreyi) and sugar pine (Pinus lambertiana), whose combined densities 16 years after treatments averaged 37 tph across treatments and did not significantly respond to thinning and/or prescribed burning. Natural conifer regeneration after treatments varied by pre-treatment vegetation type (closed canopy, Ceanothus cordulatus shrub dominated, and open sparse), with large increases of natural regeneration after understory thinning in closed canopy and Ceanothus shrub vegetation types. Planting increased sugar pine regeneration density after overstory thinning, marginally increased Jeffrey pine regeneration after overstory thinning combined with prescribed burning, and increased white fir regeneration after overstory thinning with and without burning. No treatments reduced white fir and incense-cedar natural regeneration while simultaneously increasing natural pine regeneration, suggesting new thinning, burning, and planting approaches may be required to meet regeneration restoration objectives.

Disease resistance in whitebark pine and potential for restoration of a threatened species

Societal impact statementForests world‐wide are being negatively affected by non‐native, invasive pathogens and pests, and some tree species face uncertain futures. To retain these species as components of future forests, the rare genetic resistance that exists needs to be identified and harnessed. The applied tree improvement program for whitebark pine (Pinus albicaulis), a threatened (in the United States) and endangered (in Canada) keystone species in many forests in western North America, provides an example of what can be accomplished in a relatively short timeframe. The level and frequency of resistance vary by location, and this information will be used to implement the national restoration plan.Summary Forest trees face serious threats from non‐native diseases and pests, often causing high mortality of both the existing trees and regeneration. Developing populations with genetic resistance can help restore forests and retain affected species. Resistance programs have historically focused on species of high economic importance; however, the threats to species of little direct economic value that provide other important ecosystem services are also great. We examined the frequency, level, and geographic variation in genetic resistance to white pine blister rust in the threatened Pinus albicaulis (whitebark pine), a keystone species in high‐elevation ecosystems in western North America. In the two trials reported here, 2‐year‐old seedling progeny of 225 whitebark pine parent trees were inoculated with two geographic sources of the fungal pathogen Cronartium ribicola and evaluated over 5 years for an array of resistance traits. The trials focused primarily on parent trees from the Oregon and Washington populations. We found unexpectedly high levels of quantitative resistance in some seedling families and populations, in stark contrast to levels observed in similar resistance programs with other North American white pine species such as Pinus monticola and Pinus lambertiana. The level of resistance found in some whitebark pine populations provides optimism about potential recovery efforts for this species. Restoration efforts are underway by government agencies, tribes, and non‐government organizations in both the United States and Canada. These efforts may help boost support for applied genetic resistance programs in other forest tree species severely affected by non‐native pathogens or pests.

Differences in regeneration niche mediate how disturbance severity and microclimate affect forest species composition

Climate change is altering forest composition through species-specific responses to fire and drought. Future forest composition will depend on how the different regeneration niches of co-occurring species align with current environmental conditions, especially after fire, which can promote germination by exposing mineral soil. Few studies, however, have examined the effects of disturbance severity and microclimate on post-fire regeneration to define and compare the regeneration niches of co-occurring tree species. We used seven years of annual demography and microenvironment data from a 25.6-ha fully censused, stem-mapped forest dynamics plot in California, USA, to examine how disturbance severity, snow duration, and temperature extremes affect the survival of Abies concolor and Pinus lambertiana seedlings that germinated naturally after a low- to moderate-severity fire. We defined disturbance severity at the microsite level, based on characteristics of the substrate, and at the neighborhood level, based on tree mortality. Both disturbance severity and snow duration had species-specific effects on seedling survival, but these differed by life stage. During the germination year, later snow disappearance was associated with a 0.5 increase in survival probability for A. concolor but hardly affected P. lambertiana; in contrast, higher neighborhood disturbance severity increased survival of both species. After the germination year, higher substrate burn severity was associated with a 0.8 increase in survival probability for A. concolor but hardly affected P. lambertiana; higher neighborhood disturbance severity and later snow disappearance increased annual survival of both species, but maximum summer temperature had minimal effect. Overall, available seed, higher substrate burn severity, higher neighborhood disturbance severity, and later snow disappearance promoted natural regeneration. However, lower substrate burn severity and earlier snow disappearance in the germination year disadvantaged A. concolor seedlings, increasing the relative abundance of P. lambertiana seedlings compared to the local tree population. Our results indicate that natural post-fire compositional shifts toward drought-tolerant Pinus species–and away from less drought-tolerant Abies species–are possible in the Sierra Nevada, with potential benefits for forest persistence under climate change. Broadly, we show that species differences in regeneration niches shape how disturbance severity and microclimate affect forest species composition.

A comprehensive resilience assessment of Mexican tree species and their relationship with drought events over the last century.

The resilience of forests to drought events has become a major natural resource sustainability concern, especially in response to climate change. Yet, little is known about the legacy effects of repeated droughts, and tree species ability to respond across environmental gradients. In this study, we used a tree-ring database (121 sites) to evaluate the overall resilience of tree species to drought events in the last century. We investigated how climate and geography affected the response at the species level. We evaluated temporal trends of resilience using a predictive mixed linear modeling approach. We found that pointer years (e.g., tree growth reduction) occurred during 11.3% of the 20th century, with an average decrease in tree growth of 66% compared to the previous period. The occurrence of pointer years was associated with negative values of the Standardized Precipitation Index (SPI, 81.6%) and Palmer Drought Severity Index (PDSI, 77.3%). Tree species differed in their resilience capacity, however, species inhabiting xeric conditions were less resistant but with higher recovery rates (e.g., Abies concolor, Pinus lambertiana, and Pinus jeffreyi). On average, tree species needed 2.7 years to recover from drought events, with extreme cases requiring more than a decade to reach pre-drought tree growth rates. The main abiotic factor related to resilience was precipitation, confirming that some tree species are better adapted to resist the effects of droughts. We found a temporal variation for all tree resilience indices (scaled to 100), with a decreasing resistance (-0.56 by decade) and resilience (-0.22 by decade), but with a higher recovery (+1.72 by decade) and relative resilience rate (+0.33 by decade). Our results emphasize the importance of time series of forest resilience, particularly by distinguishing the species-level response in the context of legacy of droughts, which are likely to become more frequent and intense under a changing climate.

Degradation and restoration of Indigenous California black oak (Quercus kelloggii) stands in the northern Sierra Nevada

BackgroundThe cultural connections of the Maidu to the lands they inhabit are profound with burning being a major component of their culture. California black oak plays an important role in the lifeways of many Indigenous tribes and Native peoples tend black oaks. We used multiple lines of evidence to reconstruct Indigenous fire use in a mixed conifer forest in the northern Sierra Nevada. This includes summarizing oral traditions by the Mountain Maidu, quantifying current and historical forest structure, and characterizing fire effects in an area impacted by the 2021 Dixie Fire.ResultsThe promotion of fire, both lighting- and human-ignited, has always been part of Mountain Maidu land stewardship. The mixed conifer forests in our focal study site had very high tree densities that were dominated by small, shade-tolerant species and the historic mean fire return interval was 8.6 years. Twenty-three fire intervals of 10 years or less and 11 fire intervals of 5 years or less were recorded. Based on forest inventory data collected in 1924, a majority of the forests in the broader region surrounding our focal site (82%) did not experience high levels of conifer competition. This is a stark contrast to contemporary forest conditions which showed that 73% of forests either were experiencing full site occupancy or were within the zone of imminent mortality. Contemporary forests not experiencing high levels of conifer competition had the highest density of large California black oak while forests in the zone of imminent mortality had the lowest density of large California black oak.ConclusionsFire exclusion and suppression led to a slow, albeit catastrophic, shift for California black oaks. The mixed conifer forests sampled in the early 1920s had very low tree densities and were dominated by large ponderosa and sugar pine trees which would have allowed California black oak to thrive. The number of very short fire intervals recorded is difficult to explain by lightning alone, pointing to the management of this area for Indigenous purposes. The 2021 wildfire resulted in 100% mortality of all conifer trees in the area but 61% of the California black oaks resprouted. This provides an opportunity to remove the conifer snags and allow the California black oaks to regain dominance. It would take active stewardship to produce the oak dominance again but it is possible and could be co-led by the Maidu and federal managers.

Dendroclimatic Analysis of Sierra Nevada Mixed Conifer Species between Different Diameter Size Classes

Climate change is expected to lead to higher temperatures in the Mediterranean region of northern California in the Sierra Nevada. Dendroclimatic studies typically focus on large, old trees, but there relatively limited understanding on how climatic sensitivity can vary with trees of different size classes. We collected tree increment cores and compared radial growth responses of small (20.32 to ≤40.64 cm), medium (40.64 to ≤60.96 cm), and large (>60.96 cm) diameter mixed conifer species in the Sierra Nevada to different climate variables (temperature, precipitation, and climate moisture index (CMI)). The most common tree species encountered were white fir (Abies concolor (Gord. & Glend.) Lindl.), followed by sugar pine (Pinus lambertiana Dougl.), ponderosa pine (Pinus ponderosa Dougl. Ex P. & C. Laws), and incense cedar (Calocedrus decurrens (Torr.) Florin). One of the most coherent responses from all diameter groups and across all species was the positive response to increasing minimum winter temperatures. All diameter groups and species also responded positively to precipitation and CMI at some point in the analysis period, which is the seasonal window of April of the prior year to October of the current year of ring formation. Perhaps the most notable difference when comparing the three diameter groups to climate was the higher occurrence of negative responses to the temperature of the previous year from the largest diameter group, as well as the higher number of negative responses to temperature in general. These results suggest that larger trees may be more sensitive to future climate projections compared with smaller trees and they may carry those effects into the next year. The use of dendroclimatology to assess how mixed conifer species in the Sierra Nevada responded to past climate is a key resource that can be used to infer how trees may respond to a future changing climate.

Fire-caused mortality within tree neighborhoods increases growth of Pinus lambertiana more than growth of Abies concolor

Global increases in large-tree mortality and multiple disturbances such as fire, drought, and pestilence increase the importance of understanding how large-tree growth responds to changes in competitive neighborhoods. We used 34,175 spatially mapped trees within 25.6 ha that burned at low to moderate severity in 2013 to investigate how changes to tree neighborhoods affected growth of 3,652 surviving trees of two species, Abies concolor (white fir) and Pinus lambertiana (sugar pine). Using diameter measurements taken in 2014 and 2019, we estimated post-fire radial growth of each surviving tree. We modeled annual basal area increment as a function of diameter and extracted residuals to yield relative growth rate (RGR). We used a multi-model approach and AIC to compare the species-specific effects on RGR of all post-fire tree mortality within a neighborhood radius versus mortality broken down by cause. To represent density reduction around each surviving tree, we computed crowding metrics for neighborhood radii up to 20 m for all tree mortality combined and for each mortality cause. The best-fit model included terms for density reduction within 10 m of the surviving tree due to 1) fire or mechanical damage and 2) bark beetle mortality, as well as their interactions with species and diameter. Density reduction due to fire or mechanical damage had a positive effect on RGR that was nearly twice as strong for P. lambertiana as A. concolor. Tree diameter interacted with density reduction for just P. lambertiana. Small-diameter (10-cm) P. lambertiana had a fourfold larger increase in basal area increment than large-diameter (60-cm) P. lambertiana as density reduction due to fire or mechanical damage ranged from 0 to 100 %. The strong growth response of medium-diameter P. lambertiana to density reductions could eventually increase the abundance of large-diameter (≥60 cm) P. lambertiana. In conjunction with mortality rates, which were lower for large-diameter P. lambertiana (0.97 %) than A. concolor (3.31 %), this could increase the relative abundance of large-diameter P. lambertiana from 51.2 % in 2020 to 62.6 % by 2050. Changes to competitive neighborhoods can have species-specific effects on large-tree growth, an important consideration for long-term management of fire-prone forests.

Repeated burns fail to restore pine regeneration to the natural range of variability in a Sierra Nevada mixed‐conifer forest, U.S.A.

Fire‐exclusion has acted as a major perturbation on dry conifer forests in the western United States, increasing tree density and, in mixed‐conifer forests, the dominance of shade‐tolerant species. Restoration efforts aim to reverse these effects by reducing stand density, restoring relative proportions of tree species, and reintroducing recurrent fire, but there are limited long‐term data on the effects of repeated burning on tree regeneration. We analyzed two decades of seedling and overstory data from the Teakettle Experimental Forest in the southern Sierra Nevada, California, United States to determine how thinning and repeated burning affect seedling establishment and overstory recruitment. Across treatments, pine seedling densities remained much lower than shade‐tolerant seedling densities. We found repeated burns led to modest increases in sugar pine (Pinus lambertiana) and substantial increases in incense‐cedar (Calocedrus decurrens) seedling densities 4 years postburn. No significant differences in seedling densities among repeated burning treatments were detected for Jeffrey pine (P. jeffreyi) or white fir (Abies concolor). Estimates of natural midstory recruitment were much higher among white fir and incense‐cedar than pines, even following treatments. However, postharvest planting increased rates of pine midstory recruitment in overstory thinned treatments. Our results suggest that fire‐exclusion may have shifted the ecosystem out of its initial domain of attraction, creating a forest dominated by shade‐tolerant species that exhibits hysteresis by resisting a return to a natural range of variability even after restoring structure and process. Planting pine species may be effective at overcoming this resistance to restore the forest to a pine‐dominated state.

Tree resistance to drought and bark beetle-associated mortality following thinning and prescribed fire treatments

Long-term trends show increased tree mortality over the last several decades, coinciding with above-average temperatures, high climatic water deficits, and bark beetle outbreaks. California’s recent unprecedented drought (2012–2016) highlights the need to evaluate whether thinning and prescribed fire can improve individual tree drought resistance and reduce bark beetle-associated mortality. Using a thinning and prescribed fire study on the Stanislaus-Tuolumne Experimental Forest in the central Sierra Nevada implemented prior to the drought (2011–2013), we used dendrochronological methods to estimate metrics of tree vigor (i.e., growth and resin ducts) of sugar pine (Pinus lambertiana Douglas) and white fir (Abies lowiana [Gordon & Glend.] A. Murray bis) among treatments, as well as between trees that died from bark beetle-associated mortality and their paired counterparts that survived. We used tree vigor to estimate drought resistance as the ratio between growth during drought (2012–2016) and pre-drought (2007–2011) for both species. For sugar pine, we also created analogous ratios for multiple resin duct characteristics to evaluate defense during drought. Our findings indicate that lower competition increased growth resistance of white fir, while prescribed fire had negligible impacts on growth. This translated to lower mortality, with live white fir showing higher growth resistance than those that died. While competition did not strongly affect sugar pine growth, greater growth resistance was noted for trees that lived than trees that died. However, reduced competition and prescribed fire increased defense resistance and resin duct density and relative resin duct area were negatively associated with sugar pine mortality. Live sugar pine showed greater defense resistance than dead counterparts particularly under higher levels of competition. These findings suggest thinning can promote or maintain growth during severe drought conditions and prescribed fire can be applied with negligible costs to tree growth while also producing the additional benefit of stimulating defense systems in sugar pine, which may enable them to better survive bark beetle outbreaks. Therefore, susceptibility to bark beetle-associated mortality may be ameliorated through increasing tree vigor with a combination of forest thinning and prescribed fire.

Predictive accuracy of post-fire conifer death declines over time in models based on crown and bole injury.

A key uncertainty of empirical models of post-fire tree mortality is understanding the drivers of elevated post-fire mortality several years following fire, known as delayed mortality. Delayed mortality can represent a substantial fraction of mortality, particularly for large trees that are a conservation focus in western US coniferous forests. Current post-fire tree mortality models have undergone limited evaluation of how injury level and time since fire interact to influence model accuracy and predictor variable importance. Less severe injuries potentially serve as an indicator for vulnerability to additional stressors such as bark beetle attack or moisture stress. We used a collection of 164,293 individual tree records to examine post-fire tree mortality in eight western USA conifers: Abies concolor, Abies grandis, Calocedrus decurrens, Larix occidentalis, Pinus contorta, Pinus lambertiana, Pinus ponderosa, and Pseudotsuga menziesii. We evaluated the importance of fire injury predictors on discriminating between surviving trees versus immediate and delayed post-fire mortality. We fit balanced random forest models for each species using cumulative tree mortality from 1 to 5-years post-fire. We compared these results to multi-class random forest models using first-year mortality, 2-5-year mortality, and survival 5-years post-fire as a response variable. Crown volume scorched, diameter at breast height, and relative bark char height, were used as predictor variables. The cumulative mortality models all predicted trees that died within 1-year of fire with high accuracy but failed to predict 2-5-year mortality. The multi-class models were an improvement but had lower accuracy for predicting 2-5-year mortality. Multi-class model accuracies ranged from 85% to 95% across all species for predicting 1-year post-fire mortality, 42%-71% for predicting 2-5-year mortality, and 64%-85% for predicting trees that lived past 5-years. Our study highlights the differences in tree species tolerance to fire injury and suggests that including second-order predictors such as beetle attack or climatic water stress before and after fire will be critical to improve accuracy and better understand the mechanisms and patterns of fire-caused tree death. Random forest models have potential for management applications such as post-fire harvesting and simulating future stand dynamics.

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

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

The effects of a half century of warming and fire exclusion on montane forests of the Klamath Mountains, California, USA

AbstractClimate warming and altered disturbance regimes are changing forest composition and structure worldwide. Given that species often exhibit individualistic responses to change, making predictions about the cumulative effects of multiple stressors across environmental gradients is challenging, especially in diverse communities. For example, warming temperatures are predicted to drive species upslope, whereas fire exclusion promotes the expansion of species at lower elevations where fire was historically frequent. We resampled 148 vegetation plots to assess 46 years (1969–2015) of species and community‐level response to warming and fire exclusion in a topographically complex landscape in the Klamath Mountains, California, USA, a diverse region that served as a climate refugia throughout the Holocene. We compared cover and assessed change in the elevational distributions of 12 conifer species at different life stages (i.e., seedlings, saplings, canopy). We observed consistent but non‐significant shifts upward in elevation for eight species, and a significant shift upward for one species, all of which were far less than expectations based on recent warming. Six species declined in total cover and another five declined in at least one life stage, whereas the drought‐ and fire‐intolerant Abies concolor increased by 30.7%. The largest declines were at lower elevations in drought‐tolerant, early‐seral species (Pinus lambertiana and Pinus ponderosa) and at higher elevations for the shade‐tolerant Abies magnifica var. shastensis and the regionally rare Abies lasiocarpa. Regionally rare (Picea engelmannii) and endemic (Picea breweriana) species had reductions in early life stages, portending future declines. Multivariate analyses revealed a high degree of inertia with a minor, but significant, shift in composition and a slight decrease in species turnover along the elevation gradient driven by the expansion of A. concolor. Our results indicate that most species are declining, especially at lower and mid‐elevations where fire exclusion has increased the cover of shade‐tolerant species and reduced the recruitment for fire‐adapted species. Collectively, declines in most species, insufficient upward movement to track warming, reductions in drought‐ and fire‐tolerant early‐seral species, and an increase in a single, shade‐tolerant species will leave these communities maladapted to projected climate scenarios and questions the potential for future climate refugia in this region.

Merging prescribed fires and timber harvests in the Sierra Nevada: Burn season and pruning influences in young mixed conifer stands

In dry, productive forests where historically infrequent high-severity fires are now common, new silvicultural systems will be needed to better align management activity with the ecosystem's dependent disturbance regime of frequent low and moderate-severity fires. Merging timber harvests with prescribed fire programs can be advantageous because each disturbance provides benefits that the other cannot provide alone. We conducted a study aimed at providing information to managers interested in merging gap-based silviculture with frequent prescribed fire. We studied the influence of burn season (spring versus fall) on canopy mortality and damage by conducting prescribed burns in 13-14 year old stands that had been regenerated with gap-based silviculture. We also pruned sugar pine (Pinus lambertiana) and incense-cedar (Calocedrus decurrens) prior to burns to evaluate the influence of pruning on fire related mortality and damage. Fall (21%) and spring (19%) burns resulted in similar amounts of mortality two years following burns, but the fall burns consumed considerably more fuel compared to spring burns. Percent volume crown scorch was greater in spring burns and greater when crown bases were low to the ground. Fall burns were generally favorable assuming a management context where fuel consumption and survival is desirable. However, either burn season may be acceptable using fire as a thinning mechanism is desired to encourage the development of low-density, mature stands. As a pre-fire treatment, pruning did not clearly reduce fire-related mortality or crown damage. Considering that pruning itself is a form of crown damage, it could be considered counterproductive as a pre-fire treatment because of increased heat entering pruned crowns as a result of increased surface fuel and the loss of heat-buffering lower branches. Merging gap-based silviculture with prescribed fires in perpetuity may be initially complex operationally in the Sierra Nevada, but it offers managers a disturbance regime-guided method for sustaining heterogeneity at fine and coarse scales while maintaining low surface fuels. The timing of introducing fires into young stands as well as traditional timber management tools such as rotation ages and harvest intervals can be altered depending on exact objectives.

Radial thinning ineffective at increasing large sugar pine survival

White pines (Pinus, subgenus Strobus) in North America are highly vulnerable to the native mountain pine beetle (Dendroctonus ponderosae), invasive white pine blister rust (Cronartium ribicola), and consequences of altered disturbance regimes, with sugar pine (Pinus lambertiana) being among the most in peril. Radial thinning treatments reflect an effort to balance the needs of late seral, high forest cover for some wildlife species with the desire to also promote habitat for species requiring higher light and lower competition, as well as to create heterogenous forest structure that can be more resilient to wildfire and drought. We examined 16-year effects of three radial thinning treatments that cleared approximately 0.017 ha to 0.05 ha around focal large sugar pines on reducing tree mortality and increasing sugar pine recruitment relative to untreated, control trees. Thinning treatments by order of increasing intensity included a compressed radius treatment which removed all trees and shrubs to 3 m from the crown edge, an extended radius with large tree retention treatment which removed trees <64 cm DBH and shrubs to 7.6 m from the crown edge, and an extended radius treatment which removed all trees and shrubs to 7.6 m outside of the crown edge. None of the three radial thinning treatments tested reduced mortality relative to the control, and mountain pine beetle caused the most mortality across the four treatments. However, mortality trended lower in the extended radius with retention treatment (19% compared to 30–37%) and was the only treatment where no mortality from wind or insects other than mountain pine beetle occurred. For trees that survived through the 16-year monitoring period, only trees in the extended radius treatment experienced increased growth above pre-treatment values and higher presence of sugar pine seedlings. Our results show no clear benefit was derived from the radial thinning treatments tested. While radial thinning may be a useful tool when used in conjunction with other silvicultural prescriptions that create heterogeneous stands and landscapes with a variety of forest structures, it seems unlikely that employing only radial thinning will meet restoration efforts that aim to decrease large tree mortality. This study highlights the value of long-term monitoring in the continual effort to develop new treatments that are effective at increasing forest resilience.