Temperature memory and non-structural carbohydrates mediate legacies of a hot drought in trees across the southwestern USA.

Trees are long-lived organisms that integrate climate conditions across years or decades to produce secondary growth. This integration process is sometimes referred to as 'climatic memory.' While widely perceived, the physiological processes underlying this temporal integration, such as the storage and remobilization of non-structural carbohydrates (NSC), are rarely explicitly studied. This is perhaps most apparent when considering drought legacies (perturbed post-drought growth responses to climate), and the physiological mechanisms underlying these lagged responses to climatic extremes. Yet, drought legacies are likely to become more common if warming climate brings more frequent drought. To quantify the linkages between drought legacies, climate memory and NSC, we measured tree growth (via tree ring widths) and NSC concentrations in three dominant species across the southwestern USA. We analyzed these data with a hierarchical mixed effects model to evaluate the time-scales of influence of past climate (memory) on tree growth. We then evaluated the role of climate memory and the degree to which variation in NSC concentrations were related to forward-predicted growth during the hot 2011-2012 drought and subsequent 4-year recovery period. Populus tremuloides exhibited longer climatic memory compared to either Pinus edulis or Juniperus osteosperma, but following the 2011-2012 drought, P. tremuloides trees with relatively longer memory of temperature conditions showed larger (more negative) drought legacies. Conversely, Pinus edulis trees with longer temperature memory had smaller (less negative) drought legacies. For both species, higher NSC concentrations followed more negative (larger) drought legacies, though the relevant NSC fraction differed between P. tremuloides and P. edulis. Our results suggest that differences in tree NSC are also imprinted upon tree growth responses to climate across long time scales, which also underlie tree resilience to increasingly frequent drought events under climate change.

Adapting western North American forests to climate change and wildfires: 10 common questions.

We review science-based adaptation strategies for western North American (wNA) forests that include restoring active fire regimes and fostering resilient structure and composition of forested landscapes. As part of the review, we address common questions associated with climate adaptation and realignment treatments that run counter to a broad consensus in the literature. These include the following: (1) Are the effects of fire exclusion overstated? If so, are treatments unwarranted and even counterproductive? (2) Is forest thinning alone sufficient to mitigate wildfire hazard? (3) Can forest thinning and prescribed burning solve the problem? (4) Should active forest management, including forest thinning, be concentrated in the wildland urban interface (WUI)? (5) Can wildfires on their own do the work of fuel treatments? (6) Is the primary objective of fuel reduction treatments to assist in future firefighting response and containment? (7) Do fuel treatments work under extreme fire weather? (8) Is the scale of the problem too great? Can we ever catch up? (9) Will planting more trees mitigate climate change in wNA forests? And (10) is post-fire management needed or even ecologically justified? Based on our review of the scientific evidence, a range of proactive management actions are justified and necessary to keep pace with changing climatic and wildfire regimes and declining forest heterogeneity after severe wildfires. Science-based adaptation options include the use of managed wildfire, prescribed burning, and coupled mechanical thinning and prescribed burning as is consistent with land management allocations and forest conditions. Although some current models of fire management in wNA are averse to short-term risks and uncertainties, the long-term environmental, social, and cultural consequences of wildfire management primarily grounded in fire suppression are well documented, highlighting an urgency to invest in intentional forest management and restoration of active fire regimes.

Low-density aspen seedling establishment is widespread following recent wildfires in the western United States.

Sexual regeneration is increasingly recognized as an important regeneration pathway for aspen in the western United States, a region previously thought to be too dry for seedling establishment except for during unusually wet periods. Because of this historical assumption, information on aspen seedling establishment and factors influencing its occurrence is limited and frequently anecdotal. We conducted a systematic field survey of 15 recent fire footprints that burned in 2018 in the western United States to quantify how common aspen seedling establishment is following fire and to identify factors associated with establishment. We found aspen seedling establishment in 12 of 15 (80%) of fire footprints surveyed, although densities were mostly low. Establishment probability was positively associated with mean annual precipitation and negatively associated with seed-source distance and the density of asexual aspen regeneration. Our results suggest that aspen seedling establishment may be a widespread, if often low-density, feature in postdisturbance areas. Even in low numbers, aspen seedlings may play a disproportionately large role in aspen regeneration ecology, providing adaptive capacity and facilitating local range expansion.

Temporal controls on crown nonstructural carbohydrates in southwestern US tree species.

In trees, large uncertainties remain in how nonstructural carbohydrates (NSCs) respond to variation in water availability in natural, intact ecosystems. Variation in NSC pools reflects temporal fluctuations in supply and demand, as well as physiological coordination across tree organs in ways that differ across species and NSC fractions (e.g., soluble sugars vs starch). Using landscape-scale crown (leaves and twigs) NSC concentration measurements in three foundation tree species (Populus tremuloides, Pinus edulis, Juniperus osteosperma), we evaluated in situ, seasonal variation in NSC responses to moisture stress on three timescales: short-term (via predawn water potential), seasonal (via leaf δ13C) and annual (via current year's ring width index). Crown NSC responses to moisture stress appeared to depend on hydraulic strategy, where J. osteosperma appears to regulate osmotic potentials (via higher sugar concentrations), P. edulis NSC responses suggest respiratory depletion and P. tremuloides responses were consistent with direct sink limitations. We also show that overly simplistic models can mask seasonal and tissue variation in NSC responses, as well as strong interactions among moisture stress at different timescales. In general, our results suggest large seasonal variation in crown NSC concentrations reflecting the multiple cofunctions of NSCs in plant tissues, including storage, growth and osmotic regulation of hydraulically vulnerable leaves. We emphasize that crown NSC pool size cannot be viewed as a simple physiological metric of stress; in situ NSC dynamics are complex, varying temporally, across species, among NSC fractions and among tissue types.

Testing a pyroclimatic hypothesis on the Mexico-United States border.

The "pyroclimatic hypothesis" proposed by F. Biondi and colleagues provides a basis for testable expectations about climatic and other controls of fire regimes. This hypothesis asserts an a priori relationship between the occurrence of widespread fire and values of a relevant climatic index. Such a hypothesis provides the basis for predicting spatial and temporal patterns of fire occurrence based on climatic control. Forests near the Mexico-United States border offer a place to test the relative influence of climatic and other controls in mountain ranges that are ecologically similar and subject to broadly similar top-down climatic influence, but with differing cultural influences. We tested the pyroclimatic hypothesis by comparing fire history information from the Mesa de las Guacamayas, a mountain range in northwestern Chihuahua, with previously published fire data from the Chiricahua Mountains, in southeastern Arizona, approximately 150 km away. We developed a priori hypothetical models of fire occurrence and compared their performance to empirical climate-based models. Fires were frequent at all Mesa de las Guacamayas study sites through the mid-20th century and continued uninterrupted to the present at one site, in contrast to nearly complete fire exclusion after 1892 at sites in the Chiricahua Mountains. The empirical regression models explained a higher proportion of the variability in fire regime associated with climate than did the a priori models. Actual climate-fire relationships diverged in each country after 1892. The a priori models predicted continuing fires at the same rate per century as prior to 1892; fires did in fact continue in Mexico, albeit with some alteration of fire regimes, but ceased in the United States, most likely due to changes in land use. The cross-border comparison confirms that a frequent-fire regime could cease without a climatic cause, supporting previous arguments that bottom-up factors such as livestock grazing can rapidly and drastically alter surface fire regimes. Understanding the historical patterns of climate controls on fire could inform the use of historical data as ecological reference conditions and for future sustainability.

El Niño-southern oscillation effect on a fire regime in northeastern Mexico has changed over time.

The El Niño Southern Oscillation (ENSO) is a climate-forcing mechanism that has been shown to affect precipitation and the occurrence of wildfires in many parts of the world. In the southern United States and northern Mexico, warm events (El Niño) are associated with moist winter conditions and fewer fires, while cool events (La Niñia) tend to favor dry winters and more fires. We tested this relationship in a region of northeastern Mexico by characterizing the historical fire regime and climatic influences: Fire regimes were reconstructed from fire-scar samples collected from 100 trees in three high-elevation sites on Peña Nevada in southern Nuevo Le6n. The sites were approximately 25 ha each, and the site centers were approximately 1 km apart. The earliest recorded fire occurred in 1521 and the time period we used for analysis was 1645-1929. The sites were characterized by frequent surface fires before the 1920s. In the three sites, mean fire intervals ranged from 8.6 to 9.6 years (all fires) and 11.9 to 18.6 years (fires that scarred > or = 25% of recording trees). The per-tree mean fire return interval was 17 years, and all three sites burned in the same year seven times between 1774 and 1929. After 1929, fires were nearly eliminated in all sites, likely due to human causes. We found a temporal change in the association between ENSO events and fires; before the 1830s La Niña events were significantly associated with fire years, while after the 1830s this association was not significant. In 1998, when the most severe El Niño event of the past century occurred, the three sites experienced severe, stand-replacing fires that killed many trees that had survived multiple surface fires in the past. Prior to the 1830s, fires tended to occur during dry La Niña years, but since then both La Niña and El Niño have been associated with dry years in this region, especially during the last three decades. This result suggests that ENSO effects have changed over time in this location and that phases of ENSO are not consistent indicators of precipitation, fire occurrence, or fire behavior in this area of northeastern Mexico.