Terrestrial laser scanning and low magnetic field digitization yield similar architectural coarse root traits for 32-year-old Pinus ponderosa trees.

BACKGROUND: Understanding how trees develop their root systems is crucial for the comprehension of how wildland and urban forest ecosystems plastically respond to disturbances such as harvest, fire, and climate change. The interplay between the endogenously determined root traits and the response to environmental stimuli results in tree adaptations to biotic and abiotic factors, influencing stability, carbon allocation, and nutrient uptake. Combining the three-dimensional structure of the root system, with root morphological trait information promotes a robust understanding of root function and adaptation plasticity. Low Magnetic Field Digitization coupled with AMAPmod (botAnique et Modelisation de l'Architecture des Plantes) software has been the best-performing method for describing root system architecture and providing reliable measurements of coarse root traits, but the pace and scale of data collection remain difficult. Instrumentation and applications related to Terrestrial Laser Scanning (TLS) have advanced appreciably, and when coupled with Quantitative Structure Models (QSM), have shown some potential toward robust measurements of tree root systems. Here we compare, we believe for the first time, these two methodologies by analyzing the root system of 32-year-old Pinus ponderosa trees. RESULTS: In general, at the total root system level and by root-order class, both methods yielded comparable values for the root traits volume, length, and number. QSM for each root trait was highly sensitive to the root size (i.e., input parameter PatchDiam) and models were optimized when discrete PatchDiam ranges were specified for each trait. When examining roots in the four cardinal direction sectors, we observed differences between methodologies for length and number depending on root order but not volume. CONCLUSIONS: We believe that TLS and QSM could facilitate rapid data collection, perhaps in situ, while providing quantitative accuracy, especially at the total root system level. If more detailed measures of root system architecture are desired, a TLS method would benefit from additional scans at differing perspectives, avoiding gravitational displacement to the extent possible, while subsampling roots by hand to calibrate and validate QSM models. Despite some unresolved logistical challenges, our results suggest that future use of TLS may hold promise for quantifying tree root system architecture in a rapid, replicable manner.

Long-term frequent fire and cattle grazing alter dry forest understory vegetation.

Understanding fire and large herbivore interactions in interior western forests is critical, owing to the extensive and widespread co-occurrence of these two disturbance types and multiple present and future implications for forest resilience, conservation and restoration. However, manipulative studies focused on interactions and outcomes associated with these two disturbances are rare in forested rangelands. We investigated understory vegetation response to 5-year spring and fall prescribed fire and domestic cattle grazing exclusion in ponderosa pine stands and reported long-term responses, almost two decades after the first entry fires. In fall burn areas open to cattle grazing, total understory cover prior to utilization was about 12% lower compared with fall burn areas where cattle were experimentally excluded. This response was not strongly driven by a particular palatable or unpalatable plant functional group. Fire and grazing are likely interacting in a numerically mediated process, as we found little evidence to support a functionally moderated pathway. Post-fire green-up may equalize forage to a certain extent and concentrate herbivores in the smaller burned areas within pastures, constraining a positive understory response to burning. Fall fire and grazing also increased annual forbs and resprouting shrubs. The effects of spring burning were relatively minor, and we found no interaction with grazing. The nonnative annual grass Bromus tectorum (cheatgrass) remains a problematic invader linked to fall burning but not grazing in stands that had higher propagule pressure when the experiment was initiated. At these sites, exotic grass was a major component of the vegetation by 2015, and invasion was also increasing in spring burn and unburned areas. Information from our study suggests that frequent fall fires and cattle grazing combined may reduce understory resilience in similar dry ponderosa pine forests. Consideration of longer fire return intervals, resting areas after fire, virtual fencing, or burning entire pastures may help to mitigate the effects noted in this study.

Nonstructural carbohydrates explain post-fire tree mortality and recovery patterns.

Trees use nonstructural carbohydrates (NSCs) to support many functions, including recovery from disturbances. However, NSC's importance for recovery following fire and whether NSC depletion contributes to post-fire delayed mortality are largely unknown. We investigated how fire affects NSCs based on fire-caused injury from a prescribed fire in a young Pinus ponderosa (Lawson & C. Lawson) stand. We assessed crown injury (needle scorch and bud kill) and measured NSCs of needles and inner bark (i.e., secondary phloem) of branches and main stems of trees subject to fire and at an adjacent unburned site. We measured NSCs pre-fire and at six timesteps post-fire (4 days-16 months). While all trees initially survived the fire, NSC concentrations declined quickly in burned trees relative to unburned controls over the same post-fire period. This decline was strongest for trees that eventually died, but those that survived recovered to unburned levels within 14 months post-fire. Two months post-fire, the relationship between crown scorch and NSCs of the main stem inner bark was strongly negative (Adj-R2 = 0.83). Our results support the importance of NSCs for tree survival and recovery post-fire and suggest that post-fire NSC depletion is in part related to reduced photosynthetic leaf area that subsequently limits carbohydrate availability for maintaining tree function. Crown scorch is a commonly measured metric of tree-level fire severity and is often linked to post-fire tree outcome (i.e., recovery or mortality). Thus, our finding that NSC depletion may be the mechanistic link between the fire-caused injury and tree outcome will help improve models of post-fire tree mortality and forest recovery.

Increased aridity is associated with stronger tradeoffs in ponderosa pine vital functions.

Trees must allocate resources to core functions like growth, defense, and reproduction. These allocation patterns have profound effects on forest health, yet little is known about how core functions trade off over time, and even less is known about how a changing climate will impact tradeoffs. We conducted a 21-year survey of growth, defense, and reproduction in 80 ponderosa pine individuals spanning eight populations across environmental gradients along the Colorado Front Range, USA. We used linear mixed models to describe tradeoffs among these functions and to characterize variability among and within individuals over time. Growth and defense were lower in years of high cone production, and local drought conditions amplified year-to-year tradeoffs between reproduction and growth, where trees located at sites with hotter and drier climates showed stronger tradeoffs between reproduction and growth. Our results support the environmental stress hypothesis of masting, which predicts that greater interannual variation in tree functions will be associated with more marginal environments, such as those that are prone to drought. With warming temperatures and increased exposure to drought stress, trees will be faced with stronger interannual tradeoffs, which could lead to further decreases in growth and defensive efforts, ultimately increasing risks of mortality.

Postfire futures in southwestern forests: Climate and landscape influences on trajectories of recovery and conversion.

Southwestern ponderosa pine forests are vulnerable to fire-driven conversion in a warming and drying climate, yet little is known about what kinds of ecological communities may replace them. To characterize postfire vegetation trajectories and their environmental determinants, plant assemblages (361 sample plots including 229 vascular plant species, surveyed in 2017) were sampled within eight burns that occurred between 2000 and 2003. I used nonmetric multidimensional scaling, k-means clustering, principal component analysis, and random forest models to assess relationships between vegetation pattern, topographic and landscape factors, and gridded climate data. I describe seven postfire community types, including regenerating forests of ponderosa pine, aspen, and mixed conifers, shrub-dominated communities of Gambel oak and mixed species, and herb-dominated communities of native bunchgrasses and mixtures of ruderal, native, and nonnative species. Forest recovery was generally associated with cooler, mesic sites in proximity to forested refugia; shifts toward scrub and grassland types were most common in warmer, dryer locations distant from forested refugia. Under future climate scenarios, models project decreases in postfire forest recovery and increases in nonforest vegetation. However, forest to nonforest conversion was partially offset under a scenario of reduced burn severity and increased retention of forested refugia, highlighting important management opportunities. Burning trends in the southwestern United States suggest that postfire vegetation will occupy a growing landscape fraction, compelling renewed management focus on these areas and paradigm shifts that accommodate ecological change. I illustrate how management decisions around resisting, accepting, or directing change could be informed by an understanding of processes and patterns of postfire community variation and likely future trajectories.

Essential Oil Compositions of Pinus Species (P. contorta Subsp. contorta, P. ponderosa var. ponderosa, and P. flexilis); Enantiomeric Distribution of Terpenoids in Pinus Species.

Pinus species are important in traditional medicine throughout their ranges, and pine essential oils are of interest in aromatherapy and as topical treatments. In this work, the leaf (needle) essential oils of Pinus ponderosa var. ponderosa and Pinus contorta subsp. contorta from Oregon and Pinus flexilis growing in Idaho, have been obtained by hydrodistillation and analyzed by gas chromatographic techniques. The leaf essential oil of P. ponderosa was dominated by ß-pinene (21.5-55.3%), methyl chavicol (8.5-41.5%), α-pinene (3.6-9.6%), δ-3-carene (3.6-6.2%), and α-terpineol (1.4-5.3%). The major components of P. contorta essential oil were ß-phellandrene (23.8%), terpinen-4-ol (11.0%). The essential oil of P. flexilis was dominated by α-pinene (37.1%), ß-pinene (21.9%), bornyl acetate (12.8%), and camphene (8.5%). Chiral gas chromatography revealed the enantiomeric ratios of α-pinene and limonene to be variable, but (-)-ß-pinene predominated in Pinus essential oils.

Restoration and fuel hazard reduction result in equivalent reductions in crown fire behavior in dry conifer forests.

Over the past several decades, the management of historically frequent-fire forests in the western United States has received significant attention due to the linked ecological and social risks posed by the increased occurrence of large, contiguous patches of high-severity fire. As a result, efforts are underway to simultaneously reduce potential fire and fuel hazards and restore characteristics indicative of historical forest structures and ecological processes that enhance the diversity and quality of wildlife habitat across landscapes. Despite widespread agreement on the need for action, there is a perceived tension among scientists concerning silvicultural treatments that modify stands to optimally reduce potential fire behavior (fuel hazard reduction) versus those that aim to emulate historical forest structures and create structurally complex stands (restoration). In this work, we evaluated thinning treatments in the Black Hills National Forest that exemplify the extremes of a treatment continuum that ranges from fuel hazard reduction to restoration. The goal of this work was to understand how the differing three-dimensional stand structures created by these treatment approaches altered potential fire behavior. Our results indicate that restoration treatments created higher levels of vertical and horizontal structural complexity than the fuel hazard reduction treatments but resulted in similar reductions to potential crown fire behavior. There were some trade-offs identified as the restoration treatments created larger openings, which generated faster mean rates of fire spread; however, these increased spread rates did not translate to higher levels of canopy consumption. Overall, our results suggest that treatments can create vertical and horizontal complexity desired for restoration and wildlife habitat management while reducing fire hazard and that they can be used in concert with traditional fuel hazard reduction treatments to reduce landscape scale fire risk. We also provide some suggestions to land managers seeking to design and implement prescriptions that emulate historical structures and enhance forest complexity.

Heterogeneous isotope effects decouple conifer leaf and branch sugar δ18O and δ13C.

Isotope ratios of tree-ring cellulose are a prominent tool to reconstruct paleoclimate and plant responses to environmental variation. Current models for cellulose isotope ratios assume a transfer of the environmental signals recorded in bulk leaf water to carbohydrates and ultimately into stem cellulose. However, the isotopic signal of carbohydrates exported from leaf to branch may deviate from mean leaf values if spatial heterogeneity in isotope ratios exists in the leaf. We tested whether the isotopic heterogeneity previously observed along the length of a ponderosa pine (Pinus ponderosa) leaf water was preserved in photosynthetic products. We observed an increase in both sugar and bulk tissue δ18O values along the needle, but the increase in carbohydrate δ18O values was dampened relative to the trend observed in leaf water. In contrast, δ13C values of both sugar and bulk organic matter were invariant along the needle. Phloem-exported sugar measured in the branch below the needles did not match whole-needle values of δ18O or δ13C. Instead, there was a near-constant offset observed between the branch and needle sugar δ13C values, while branch δ18O values were most similar to δ18O values observed for sugar at the base of the needle. The observed offset between the branch and needle sugar δ18O values likely arises from partial isotope oxygen exchange between sugars and water during phloem loading and transport. An improved understanding of the conditions producing differential δ13C and δ18O isotope effects between branch phloem and needle sugars could improve tree-ring-based climate reconstructions.

Simulated treatment effects on bird communities inform landscape-scale dry conifer forest management.

Human land use and climate change have increased forest density and wildfire risk in dry conifer forests of western North America, threatening various ecosystem services, including habitat for wildlife. Government policy supports active management to restore historical structure and ecological function. Information on potential contributions of restoration to wildlife habitat can allow assessment of tradeoffs with other ecological benefits when prioritizing treatments. We predicted avian responses to simulated treatments representing alternative scenarios to inform landscape-scale forest management planning along the Colorado Front Range. We used data from the Integrated Monitoring in Bird Conservation Regions program to inform a hierarchical multispecies occupancy model relating species occupancy and richness with canopy cover at two spatial scales. We then simulated changes in canopy cover (remotely sensed in 2018) under three alternative scenarios, (1) a "fuels reduction" scenario representing landscape-wide 30% reduction in canopy cover, (2) a "restoration" scenario representing more nuanced, spatially variable treatments targeting historical conditions, and (3) a reference, no-change scenario. Model predictions showed areas of potential gains and losses for species richness, richness of ponderosa pine forest habitat specialists, and the ratio of specialists to generalists at two (1 km2 and 250 m2 ) spatial scales. Under both fuels reduction and restoration scenarios, we projected greater gains than losses for species richness. Surprisingly, despite restoration more explicitly targeting ecologically relevant historical conditions, fuels reduction benefited bird species richness over a greater spatial extent than restoration, particularly in the lower montane life zone. These benefits reflected generally positive species associations with moderate canopy cover promoted more consistently under the fuels reduction scenario. In practice, contemporary forest management is likely to lie somewhere between the fuels reduction and restoration scenarios represented here. Therefore, our results inform where and how active forest management can best support avian diversity. Although our study raises questions regarding the value of including landscape-scale heterogeneity as a management objective, we do not question the value of targeting finer scale heterogeneity (i.e., stand and treatment level). Rather, our results combined with those from previous work clarify the scale at which targeting structural heterogeneity and historical reference conditions can promote particular ecosystem services.

Ecological forecasting of tree growth: Regional fusion of tree-ring and forest inventory data to quantify drivers and characterize uncertainty.

Robust ecological forecasting of tree growth under future climate conditions is critical to anticipate future forest carbon storage and flux. Here, we apply three ingredients of ecological forecasting that are key to improving forecast skill: data fusion, confronting model predictions with new data, and partitioning forecast uncertainty. Specifically, we present the first fusion of tree-ring and forest inventory data within a Bayesian state-space model at a multi-site, regional scale, focusing on Pinus ponderosa var. brachyptera in the southwestern US. Leveraging the complementarity of these two data sources, we parsed the ecological complexity of tree growth into the effects of climate, tree size, stand density, site quality, and their interactions, and quantified uncertainties associated with these effects. New measurements of trees, an ongoing process in forest inventories, were used to confront forecasts of tree diameter with observations, and evaluate alternative tree growth models. We forecasted tree diameter and increment in response to an ensemble of climate change projections, and separated forecast uncertainty into four different causes: initial conditions, parameters, climate drivers, and process error. We found a strong negative effect of fall-spring maximum temperature, and a positive effect of water-year precipitation on tree growth. Furthermore, tree vulnerability to climate stress increases with greater competition, with tree size, and at poor sites. Under future climate scenarios, we forecast increment declines of 22%-117%, while the combined effect of climate and size-related trends results in a 56%-91% decline. Partitioning of forecast uncertainty showed that diameter forecast uncertainty is primarily caused by parameter and initial conditions uncertainty, but increment forecast uncertainty is mostly caused by process error and climate driver uncertainty. This fusion of tree-ring and forest inventory data lays the foundation for robust ecological forecasting of aboveground biomass and carbon accounting at tree, plot, and regional scales, including iterative improvement of model skill.

Changes in tree drought sensitivity provided early warning signals to the California drought and forest mortality event.

Climate warming in recent decades has negatively impacted forest health in the western United States. Here, we report on potential early warning signals (EWS) for drought-related mortality derived from measurements of tree-ring growth (ring width index; RWI) and carbon isotope discrimination (∆13 C), primarily focused on ponderosa pine (Pinus ponderosa). Sampling was conducted in the southern Sierra Nevada Mountains, near the epicenter of drought severity and mortality associated with the 2012-2015 California drought and concurrent outbreak of western pine beetle (Dendroctonus brevicomis). At this site, we found that widespread mortality was presaged by five decades of increasing sensitivity (i.e., increased explained variation) of both tree growth and ∆13 C to Palmer Drought Severity Index (PDSI). We hypothesized that increasing sensitivity of tree growth and ∆13 C to hydroclimate constitute EWS that indicate an increased likelihood of widespread forest mortality caused by direct and indirect effects of drought. We then tested these EWS in additional ponderosa pine-dominated forests that experienced varying mortality rates associated with the same California drought event. In general, drier sites showed increasing sensitivity of RWI to PDSI over the last century, as well as higher mortality following the California drought event compared to wetter sites. Two sites displayed evidence that thinning or fire events that reduced stand basal area effectively reversed the trend of increasing hydroclimate sensitivity. These comparisons indicate that reducing competition for soil water and/or decreasing bark beetle host tree density via forest management-particularly in drier regions-may buffer these forests against drought stress and associated mortality risk. EWS such as these could provide land managers more time to mitigate the extent or severity of forest mortality in advance of droughts. Substantial efforts at deploying additional dendrochronological research in concert with remote sensing and forest modeling will aid in forecasting of forest responses to continued climate warming.

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

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

Bimodal cambial activity and false-ring formation in conifers under a monsoon climate.

Tracking wood formation in semiarid regions during the seasonal march of precipitation extremes has two important applications. It can provide (i) insight into the adaptive capacities of trees to drought and (ii) a basis for a richer interpretation of tree-ring data, assisting in a deeper understanding of past and current climate. In the southwestern USA, the anatomical signature of seasonally bimodal precipitation is the 'false ring'-a band of latewood-like cells in the earlywood. These occur when a particularly deep drought during the early growing season ends abruptly with timely, mid-growing season monsoonal rains. Such conditions presented in southern Arizona in 2014, enabling us to explore false-ring formation in ponderosa pine (Pinus ponderosa Lawson and C. Lawson) and Douglas-fir (Pseudotsuga menziesii Mirb. Franco) in mixed-conifer forest at 2573 m above sea level. We ask: what were the cell-by-cell timings and durations in the phases of wood cell development in 2014? How do these seasonal patterns relate to strongly fluctuating environmental conditions during the growing season? We took weekly microcores from March through November from six ponderosa pine and seven Douglas-fir trees at a well-instrumented flux tower site. Thin sections were prepared, and we counted cells in cambial, expansion, cell wall thickening and mature phases. For ponderosa pine trees forming a false ring, the first impact of intensifying seasonal drought was seen in the enlarging phase and then, almost a month later, in cambial activity. In this species, recovery from drought was associated with recovery first in cambial activity, followed by cell enlargement. This timing raised the possibility that cell division may be affected by atmospheric moisture increases before soil recharge. In both species, the last false-ring cells matured during the summer rainy season. Bimodal cambial activity coincident with moisture availability was observed in both species, whether or not they formed a false ring. This deeper knowledge of the precise timing of both developmental and environmental events should help define mechanistic connections among these factors in creating bimodal growth patterns.

Native American fire management at an ancient wildland-urban interface in the Southwest United States.

The intersection of expanding human development and wildland landscapes-the "wildland-urban interface" or WUI-is one of the most vexing contexts for fire management because it involves complex interacting systems of people and nature. Here, we document the dynamism and stability of an ancient WUI that was apparently sustainable for more than 500 y. We combine ethnography, archaeology, paleoecology, and ecological modeling to infer intensive wood and fire use by Native American ancestors of Jemez Pueblo and the consequences on fire size, fire-climate relationships, and fire intensity. Initial settlement of northern New Mexico by Jemez farmers increased fire activity within an already dynamic landscape that experienced frequent fires. Wood harvesting for domestic fuel and architectural uses and abundant, small, patchy fires created a landscape that burned often but only rarely burned extensively. Depopulation of the forested landscape due to Spanish colonial impacts resulted in a rebound of fuels accompanied by the return of widely spreading, frequent surface fires. The sequence of more than 500 y of perennial small fires and wood collecting followed by frequent "free-range" wildland surface fires made the landscape resistant to extreme fire behavior, even when climate was conducive and surface fires were large. The ancient Jemez WUI offers an alternative model for fire management in modern WUI in the western United States, and possibly other settings where local management of woody fuels through use (domestic wood collecting) coupled with small prescribed fires may make these communities both self-reliant and more resilient to wildfire hazards.

Forest restoration treatments in a ponderosa pine forest enhance physiological activity and growth under climatic stress.

As the climate warms, drought will increasingly occur under elevated temperatures, placing forest ecosystems at growing risk of extensive dieback and mortality. In some cases, increases in tree density following early 20th-century fire suppression may exacerbate this risk. Treatments designed to restore historical stand structure and enhance resistance to high-severity fire might also alleviate drought stress by reducing competition, but the duration of these effects and the underlying mechanisms remain poorly understood. To elucidate these mechanisms, we evaluate tree growth, mortality, and tree-ring stable-carbon isotope responses to stand-density reduction treatments with and without prescribed fire in a ponderosa pine forest of western Montana. Moderate and heavier cutting experiments (basal area reductions of 35% and 56%, respectively) were initiated in 1992, followed by prescribed burning in a subset of the thinned units. All treatments led to a growth release that persisted to the time of resampling. The treatments had little effect on climate-growth relationships, but they markedly altered seasonal carbon isotope signals and their relationship to climate. In burned and unburned treatments, carbon isotope discrimination (Δ13 C) increased in the earlywood (EW) and decreased in the latewood (LW) relative to the control. The sensitivity of LW Δ13 C to late-summer climate also increased in all treatments, but not in the control. Such increased sensitivity indicates that the reduction in competition enabled trees to continue to fix carbon for new stem growth, even when the climate became sufficiently stressful to stop new assimilation in slower-growing trees in untreated units. These findings would have been masked had we not separated EW and LW. The importance of faster growth and enhanced carbon assimilation under late-summer climatic stress became evident in the second decade post-treatment, when mountain pine beetle activity increased locally, and tree mortality rates in the controls of both experiments increased to more than twice those in their respective treatments. These findings highlight that, when thinning is used to restore historical forest structure or increase resistance to high-severity fire, there will likely be additional benefits of enhanced growth and physiological activity under climatic stress, and the effects may persist for more than two decades.

Dry conifer forest restoration benefits Colorado Front Range avian communities.

Fire suppression has increased stand density and risk of severe, stand-replacing wildfire in lower elevation dry conifer forests of western North America, threatening ecological function. The U.S. Forest Service's Collaborative Forest Landscape Restoration Program (CFLRP) aims to mitigate impacts to ecological function, while mandating effectiveness monitoring to verify restoration success. Expected benefits include improved conditions for biodiversity, but relatively few empirical studies evaluate restoration effects on biodiversity. We applied the Integrated Monitoring in Bird Conservation Regions program to survey birds in relation to CFLRP treatments along the Colorado Front Range in 2015-2017. We employed hierarchical models to analyze species occupancy and richness at 1972 points nested within 141 1-km2 grid cells. Our objectives were to investigate (1) species occupancy relationships with treatments at local (point) and landscape (grid) spatial scales, (2) potential mechanisms for treatment relationships considering species and treatment relationships with forest structure and composition (i.e., habitat relationships), and (3) treatment and habitat relationships with species richness. The data supported positive and negative point-level treatment relationships, suggesting uneven species distributions between treated and untreated points. At the grid scale, however, we only found positive species relationships with percent area treated, and accordingly, grid-level species richness increased with treatment extent. Potential mechanisms for treatment relationships included treatments generating foraging opportunities for aerial insectivores by opening the canopy, improving conditions for ground-associated species by increasing herbaceous growth, and limiting opportunities for shrub-nesting species by reducing shrub cover. Landscape-scale patterns suggest CFLRP treatments can benefit avian communities by generating habitat for open-forest species without necessarily eliminating habitat for closed-forest species. Our results provide evidence for a commonly expected but rarely verified pattern of increased species richness with forest heterogeneity. We suggest restoration treatments will most benefit forest bird diversity by reducing canopy cover, encouraging herbaceous ground cover, limiting ladder fuel species, and encouraging shrub diversity in canopy openings, while maintaining some dense forest stands on the landscape.

Limitations to recovery following wildfire in dry forests of southern Colorado and northern New Mexico, USA.

Climate warming is contributing to increases in wildfire activity throughout the western United States, leading to potentially long-lasting shifts in vegetation. The response of forest ecosystems to wildfire is thus a crucial indicator of future vegetation trajectories, and these responses are contingent upon factors such as seed availability, interannual climate variability, average climate, and other components of the physical environment. To better understand variation in resilience to wildfire across vulnerable dry forests, we surveyed conifer seedling densities in 15 recent (1988-2010) wildfires and characterized temporal variation in seed cone production and seedling establishment. We then predicted postfire seedling densities at a 30-m resolution within each fire perimeter using downscaled climate data, monthly water balance models, and maps of surviving forest cover. Widespread ponderosa pine (Pinus ponderosa) seed cone production occurred at least twice following each fire surveyed, and pulses of conifer seedling establishment coincided with years of above-average moisture availability. Ponderosa pine and Douglas-fir (Pseudotsuga menziesii) seedling densities were higher on more mesic sites and adjacent to surviving trees, though there were also important interspecific differences, likely attributable to drought and shade tolerance. We estimated that postfire seedling densities in 42% (for ponderosa pine) and 69% (for Douglas-fir) of the total burned area were below the lowest reported historical tree densities in these forests. Spatial models demonstrated that an absence of mature conifers (particularly in the interior of large, high-severity patches) limited seedling densities in many areas, but 30-yr average actual evapotranspiration and climatic water deficit limited densities on marginal sites. A better understanding of the limitations to postfire forest recovery will refine models of vegetation dynamics and will help to improve strategies of adaptation to a warming climate and shifting fire activity.

Large-scale forest restoration stabilizes carbon under climate change in Southwest United States.

Higher tree density, more fuels, and a warmer, drier climate have caused an increase in the frequency, size, and severity of wildfires in western U.S. forests. There is an urgent need to restore forests across the western United States. To address this need, the U.S. Forest Service began the Four Forest Restoration Initiative (4FRI) to restore four national forests in Arizona. The objective of this study was to evaluate how restoration of ~400,000 ha under the 4FRI program and projected climate change would influence carbon dynamics and wildfire severity from 2010 to 2099. Specifically, we estimated forest carbon fluxes, carbon pools and wildfire severity under a moderate and fast 4FRI implementation schedule and compared those to status quo and no-harvest scenarios using the LANDIS-II simulation model and climate change projections. We found that the fast-4FRI scenario showed early decreases in ecosystem carbon due to initial thinning/prescribed fire treatments, but total ecosystem carbon increased by 9-18% over no harvest by the end of the simulation. This increased carbon storage by 6.3-12.7 million metric tons, depending on the climate model, equating to removal of carbon emissions from 55,000 to 110,000 passenger vehicles per year until the end of the century. Nearly half of the additional carbon was stored in more stable soil pools. However, climate models with the largest predicted temperature increases showed declines by late century in ecosystem carbon despite restoration. Our study uses data from a real-world, large-scale restoration project and indicates that restoration is likely to stabilize carbon and the benefits are greater when the pace of restoration is faster.

Wildfires and climate change push low-elevation forests across a critical climate threshold for tree regeneration.

Climate change is increasing fire activity in the western United States, which has the potential to accelerate climate-induced shifts in vegetation communities. Wildfire can catalyze vegetation change by killing adult trees that could otherwise persist in climate conditions no longer suitable for seedling establishment and survival. Recently documented declines in postfire conifer recruitment in the western United States may be an example of this phenomenon. However, the role of annual climate variation and its interaction with long-term climate trends in driving these changes is poorly resolved. Here we examine the relationship between annual climate and postfire tree regeneration of two dominant, low-elevation conifers (ponderosa pine and Douglas-fir) using annually resolved establishment dates from 2,935 destructively sampled trees from 33 wildfires across four regions in the western United States. We show that regeneration had a nonlinear response to annual climate conditions, with distinct thresholds for recruitment based on vapor pressure deficit, soil moisture, and maximum surface temperature. At dry sites across our study region, seasonal to annual climate conditions over the past 20 years have crossed these thresholds, such that conditions have become increasingly unsuitable for regeneration. High fire severity and low seed availability further reduced the probability of postfire regeneration. Together, our results demonstrate that climate change combined with high severity fire is leading to increasingly fewer opportunities for seedlings to establish after wildfires and may lead to ecosystem transitions in low-elevation ponderosa pine and Douglas-fir forests across the western United States.

Fire legacies in eastern ponderosa pine forests.

Disturbance legacies structure communities and ecological memory, but due to increasing changes in disturbance regimes, it is becoming more difficult to characterize disturbance legacies or determine how long they persist. We sought to quantify the characteristics and persistence of material legacies (e.g., biotic residuals of disturbance) that arise from variation in fire severity in an eastern ponderosa pine forest in North America. We compared forest stand structure and understory woody plant and bird community composition and species richness across unburned, low-, moderate-, and high-severity burn patches in a 27-year-old mixed-severity wildfire that had received minimal post-fire management. We identified distinct tree densities (high: 14.3 ± 7.4 trees per ha, moderate: 22.3 ± 12.6, low: 135.3 ± 57.1, unburned: 907.9 ± 246.2) and coarse woody debris cover (high: 8.5 ± 1.6% cover per 30 m transect, moderate: 4.3 ± 0.7, low: 2.3 ± 0.6, unburned: 1.0 ± 0.4) among burn severities. Understory woody plant communities differed between high-severity patches, moderate- and low-severity patches, and unburned patches (all p < 0.05). Bird communities differed between high- and moderate-severity patches, low-severity patches, and unburned patches (all p < 0.05). Bird species richness varied across burn severities: low-severity patches had the highest (5.29 ± 1.44) and high-severity patches had the lowest (2.87 ± 0.72). Understory woody plant richness was highest in unburned (5.93 ± 1.10) and high-severity (5.07 ± 1.17) patches, and it was lower in moderate- (3.43 ± 1.17) and low-severity (3.43 ± 1.06) patches. We show material fire legacies persisted decades after the mixed-severity wildfire in eastern ponderosa forest, fostering distinct structures, communities, and species in burned versus unburned patches and across fire severities. At a patch scale, eastern and western ponderosa system responses to mixed-severity fires were consistent.