Jens T. Stevens

Long-term climate and competition explain forest mortality patterns under extreme drought

Rising temperatures are amplifying drought-induced stress and mortality in forests globally. It remains uncertain, however, whether tree mortality across drought-stricken landscapes will be concentrated in particular climatic and competitive environments. We investigated the effects of long-term average climate [i.e. 35-year mean annual climatic water deficit (CWD)] and competition (i.e. tree basal area) on tree mortality patterns, using extensive aerial mortality surveys conducted throughout the forests of California during a 4-year statewide extreme drought lasting from 2012 to 2015. During this period, tree mortality increased by an order of magnitude, typically from tens to hundreds of dead trees per km2 , rising dramatically during the fourth year of drought. Mortality rates increased independently with average CWD and with basal area, and they increased disproportionately in areas that were both dry and dense. These results can assist forest managers and policy-makers in identifying the most drought-vulnerable forests across broad geographic areas.

Plant communities in harsh sites are less invaded: a summary of observations and proposed explanations.

Within the invasion ecology literature, it is often noted that abiotically stressful environments are typically less invaded by non-native plants than nearby less-stressful environments. However, until now no one had collected and summarized examples of this pattern. This paper first compiles evidence that plant communities in many harsh habitats are less invaded, and then synthesizes possible explanations for this pattern. We discuss that harsh sites may be less invaded because, compared to moderate sites, they may receive lower propagule pressure, particularly from well-suited plants, and because their abiotic and biotic characteristics may make them inherently more resistant to invasion.

Average Stand Age from Forest Inventory Plots Does Not Describe Historical Fire Regimes in Ponderosa Pine and Mixed-Conifer Forests of Western North America

Quantifying historical fire regimes provides important information for managing contemporary forests. Historical fire frequency and severity can be estimated using several methods; each method has strengths and weaknesses and presents challenges for interpretation and verification. Recent efforts to quantify the timing of historical high-severity fire events in forests of western North America have assumed that the “stand age” variable from the US Forest Service Forest Inventory and Analysis (FIA) program reflects the timing of historical high-severity (i.e. stand-replacing) fire in ponderosa pine and mixed-conifer forests. To test this assumption, we re-analyze the dataset used in a previous analysis, and compare information from fire history records with information from co-located FIA plots. We demonstrate that 1) the FIA stand age variable does not reflect the large range of individual tree ages in the FIA plots: older trees comprised more than 10% of pre-stand age basal area in 58% of plots analyzed and more than 30% of pre-stand age basal area in 32% of plots, and 2) recruitment events are not necessarily related to high-severity fire occurrence. Because the FIA stand age variable is estimated from a sample of tree ages within the tree size class containing a plurality of canopy trees in the plot, it does not necessarily include the oldest trees, especially in uneven-aged stands. Thus, the FIA stand age variable does not indicate whether the trees in the predominant size class established in response to severe fire, or established during the absence of fire. FIA stand age was not designed to measure the time since a stand-replacing disturbance. Quantification of historical “mixed-severity” fire regimes must be explicit about the spatial scale of high-severity fire effects, which is not possible using FIA stand age data.

Forest disturbance accelerates thermophilization of understory plant communities

Summary 1. Climate change is likely to shift plant communities towards species from warmer regions, a process termed ‘thermophilization’. In forests, canopy disturbances such as fire may hasten this process by increasing temperature and moisture stress in the understory, yet little is known about the mechanisms that might drive such shifts, or the consequences of these processes for plant diversity. 2. We sampled understory vegetation across a gradient of disturbance severity from a large-scale natural experiment created by the factorial combination of forest thinning and wildfire in California. Using information on evolutionary history and functional traits, we tested the hypothesis that disturbance severity should increase community dominance by species with southern-xeric biogeographic affinities. We also analysed how climatic productivity mediates the effect of disturbance severity, and quantified the functional trait response to disturbance, to investigate potential mechanisms behind thermophilization. 3. The proportion of north-temperate flora decreased, while the proportion of southern-xeric flora increased, with greater disturbance severity and less canopy closure. Disturbance caused a greater reduction of north-temperate flora in productive (wetter) forests, while functional trait analyses suggested that species colonizing after severe disturbance may be adapted to increased water stress. Forests with intermediate disturbance severity, where abundances of northern and southern species were most equitable, had the highest stand-scale understory diversity. 4. Synthesis: Canopy disturbance is likely to accelerate plant community shifts towards species from warmer regions, via its effects on understory microclimate at small scales. Understory diversity can be enhanced by intermediate disturbance regimes that promote the coexistence of species with different biogeographic affinities.

Snowpack, fire, and forest disturbance: interactions affect montane invasions by non‐native shrubs

Montane regions worldwide have experienced relatively low plant invasion rates, a trend attributed to increased climatic severity, low rates of disturbance, and reduced propagule pressure relative to lowlands. Manipulative experiments at elevations above the invasive range of non-native species can clarify the relative contributions of these mechanisms to montane invasion resistance, yet such experiments are rare. Furthermore, global climate change and land use changes are expected to cause decreases in snowpack and increases in disturbance by fire and forest thinning in montane forests. We examined the importance of these factors in limiting montane invasions using a field transplant experiment above the invasive range of two non-native lowland shrubs, Scotch broom (Cytisus scoparius) and Spanish broom (Spartium junceum), in the rain-snow transition zone of the Sierra Nevada of California. We tested the effects of canopy closure, prescribed fire, and winter snow depth on demographic transitions of each species. Establishment of both species was most likely at intermediate levels of canopy disturbance, but at this intermediate canopy level, snow depth had negative effects on winter survival of seedlings. We used matrix population models to show that an 86% reduction in winter snowfall would cause a 2.8-fold increase in population growth rates in Scotch broom and a 3.5-fold increase in Spanish broom. Fall prescribed fire increased germination rates, but decreased overall population growth rates by reducing plant survival. However, at longer fire return intervals, population recovery between fires is likely to keep growth rates high, especially under low snowpack conditions. Many treatment combinations had positive growth rates despite being above the current invasive range, indicating that propagule pressure, disturbance, and climate can all strongly affect plant invasions in montane regions. We conclude that projected reductions in winter snowpack and increases in forest disturbance are likely to increase the risk of invasion from lower elevations.

Alternative characterization of forest fire regimes: incorporating spatial patterns

ContextThe proportion of fire area that experienced stand-replacing fire effects is an important attribute of individual fires and fire regimes in forests, and this metric has been used to group forest types into characteristic fire regimes. However, relying on proportion alone ignores important spatial characteristics of stand-replacing patches, which can have a strong influence on post-fire vegetation dynamics.ObjectivesWe propose a new more ecologically relevant approach for characterizing spatial patterns of stand-replacing patches to account for potential limitation of conifer seed dispersal.MethodsWe applied a simple modified logistic function to describe the relationship between the proportion of total stand-replacing patch area and an interior buffer distance on stand-replacing patches.ResultsThis approach robustly distinguishes among different spatial configurations of stand-replacing area in both theoretical and actual fires, and does so uniquely from commonly used descriptors of spatial configuration.ConclusionsOur function can be calculated for multiple fires over a given area, allowing for meaningful ecological comparisons of stand-replacing effects among different fires and regions.

Scale‐dependent effects of post‐fire canopy cover on snowpack depth in montane coniferous forests

Winter snowpack in dry montane regions provides a valuable ecosystem service by storing water into the growing season. Wildfire in coniferous montane forests has the potential to indirectly affect snowpack accumulation and ablation (mass loss) rates by reducing canopy cover, which reduces canopy interception of snow but also increases solar radiation and wind speed. These counteracting effects create uncertainty regarding the canopy conditions that maximize post-fire snowpack duration, which is of concern as montane regions across the western United States experience increasingly warm, dry winters with below-average snowpack. The net effect of wildfire on snowpack depth and duration across the landscape is uncertain, and likely scale dependent. In this study, I tested whether intermediate levels of wildfire severity maximize snowpack depth by increasing accumulation while slowing ablation, using gridded, repeated snow depth measurements from three fires in the Sierra Nevada of California. Increasing fire severity had a strong negative effect on snowpack depth, suggesting that increased ablation after fire, rather than increased accumulation, was the dominant control over snowpack duration. Contrary to expectations, the unburned forest condition had the highest overall snowpack depth, and mean snow depth among all site visits was reduced by 78% from unburned forest to high-severity fire. However, at the individual tree scale, snowpack depth was greater under canopy openings than underneath canopy, controlling for effects of fire severity and aspect. This apparent paradox in snowpack response to fire at the stand vs. individual tree scales is likely due to greater variation in canopy cover within unburned and very low severity areas, which creates smaller areas for snow accumulation while reducing ablation via shading. Management efforts to maximize snowpack duration in montane forests should focus on retaining fine-scale heterogeneity in forest structure.

Extreme mid‐winter drought weakens tree hydraulic–carbohydrate systems and slows growth

Rising temperatures and extended periods of drought compromise tree hydraulic and carbohydrate systems, threatening forest health globally. Despite winters biological significance to many forests, the effects of warmer and dryer winters on tree hydraulic and carbohydrate status have largely been overlooked. Here we report a sharp and previously unknown decline in stem water content of three conifer species during Californias anomalous 2015 mid-winter drought that was followed by dampened spring starch accumulation. Recent precipitation and seasonal vapor pressure deficit (VPD) anomaly, not absolute VPD, best predicted the hydraulic patterns observed. By linking relative water content and hydraulic conductivity (Kh ), we estimated that stand-level Kh declined by 52% during Californias 2015 mid-winter drought, followed by a 50% reduction in spring starch accumulation. Further examination of tree increment records indicated a concurrent decline of growth with rising mid-winter, but not summer, VPD anomaly. Thus, our findings suggest a seasonality to tree hydraulic and carbohydrate declines, with consequences for annual growth rates, raising novel physiological and ecological questions about how rising winter temperatures will affect forest vitality as climate changes.