Western Ecological Research Center

Fire is a worldwide phenomenon that appears in the geological record soon after the appearance of terrestrial plants. Fire influences global ecosystem patterns and processes, including vegetation distribution and structure, the carbon cycle, and climate. Although humans and fire have always coexisted, our capacity to manage fire remains imperfect and may become more difficult in the future as climate change alters fire regimes. This risk is difficult to assess, however, because fires are still poorly represented in global models. Here, we discuss some of the most important issues involved in developing a better understanding of the role of fire in the Earth system.

Several recent papers have suggested replacing the terminology of fire intensity and fire severity. Part of the problem with fire intensity is that it is sometimes used incorrectly to describe fire effects, when in fact it is justifiably restricted to measures of energy output. Increasingly, the term has created confusion because some authors have restricted its usage to a single measure of energy output referred to as fireline intensity. This metric is most useful in understanding fire behavior in forests, but is too narrow to fully capture the multitude of ways fire energy affects ecosystems. Fire intensity represents the energy released during various phases of a fire, and different metrics such as reaction intensity, fireline intensity, temperature, heating duration and radiant energy are useful for different purposes. Fire severity, and the related term burn severity, have created considerable confusion because of recent changes in their usage. Some authors have justified this by contending that fire severity is defined broadly as ecosystem impacts from fire and thus is open to individual interpretation. However, empirical studies have defined fire severity operationally as the loss of or change in organic matter aboveground and belowground, although the precise metric varies with management needs. Confusion arises because fire or burn severity is sometimes defined so that it also includes ecosystem responses. Ecosystem responses include soil erosion, vegetation regeneration, restoration of community structure, faunal recolonization, and a plethora of related response variables. Although some ecosystem responses are correlated with measures of fire or burn severity, many important ecosystem processes have either not been demonstrated to be predicted by severity indices or have been shown in some vegetation types to be unrelated to severity. This is a critical issue because fire or burn severity are readily measurable parameters, both on the ground and with remote sensing, yet ecosystem responses are of most interest to resource managers.

Persistent changes in tree mortality rates can alter forest structure, composition, and ecosystem services such as carbon sequestration. Our analyses of longitudinal data from unmanaged old forests in the western United States showed that background (noncatastrophic) mortality rates have increased rapidly in recent decades, with doubling periods ranging from 17 to 29 years among regions. Increases were also pervasive across elevations, tree sizes, dominant genera, and past fire histories. Forest density and basal area declined slightly, which suggests that increasing mortality was not caused by endogenous increases in competition. Because mortality increased in small trees, the overall increase in mortality rates cannot be attributed solely to aging of large trees. Regional warming and consequent increases in water deficits are likely contributors to the increases in tree mortality rates.

Plant invasions are widely recognized as significant threats to biodiversity conservation worldwide. One way invasions can affect native ecosystems is by changing fuel properties, which can in turn affect fire behavior and, ultimately, alter fire regime characteristics such as frequency, intensity, extent, type, and seasonality of fire. If the regime changes subsequently promote the dominance of the invaders, then an invasive plant-fire regime cycle can be established. As more ecosystem components and interactions are altered, restoration of preinvasion conditions becomes more difficult. Restoration may require managing fuel conditions, fire regimes, native plant communities, and other ecosystem properties in addition to the invaders that caused the changes in the first place. We present a multiphase model describing the interrelationships between plant invaders and fire regimes, provide a system for evaluating the relative effects of invaders and prioritizing them for control, and recommend ways to restore preinvasion fire regime properties.

Community assembly theory suggests that two processes affect the distribution of trait values within communities: competition and habitat filtering. Within a local community, competition leads to ecological differentiation of coexisting species, while habitat filtering reduces the spread of trait values, reflecting shared ecological tolerances. Many statistical tests for the effects of competition exist in the literature, but measures of habitat filtering are less well-developed. Here, we present convex hull volume, a construct from computational geometry, which provides an n-dimensional measure of the volume of trait space occupied by species in a community. Combined with ecological null models, this measure offers a useful test for habitat filtering. We use convex hull volume and a null model to analyze California woody-plant trait and community data. Our results show that observed plant communities occupy less trait space than expected from random assembly, a result consistent with habitat filtering.

Humans and their ancestors are unique in being a fire-making species, but 'natural' (i.e. independent of humans) fires have an ancient, geological history on Earth. Natural fires have influenced biological evolution and global biogeochemical cycles, making fire integral to the functioning of some biomes. Globally, debate rages about the impact on ecosystems of prehistoric human-set fires, with views ranging from catastrophic to negligible. Understanding of the diversity of human fire regimes on Earth in the past, present and future remains rudimentary. It remains uncertain how humans have caused a departure from 'natural' background levels that vary with climate change. Available evidence shows that modern humans can increase or decrease background levels of natural fire activity by clearing forests, promoting grazing, dispersing plants, altering ignition patterns and actively suppressing fires, thereby causing substantial ecosystem changes and loss of biodiversity. Some of these contemporary fire regimes cause substantial economic disruptions owing to the destruction of infrastructure, degradation of ecosystem services, loss of life, and smoke-related health effects. These episodic disasters help frame negative public attitudes towards landscape fires, despite the need for burning to sustain some ecosystems. Greenhouse gas-induced warming and changes in the hydrological cycle may increase the occurrence of large, severe fires, with potentially significant feedbacks to the Earth system. Improved understanding of human fire regimes demands: (1) better data on past and current human influences on fire regimes to enable global comparative analyses, (2) a greater understanding of different cultural traditions of landscape burning and their positive and negative social, economic and ecological effects, and (3) more realistic representations of anthropogenic fire in global vegetation and climate change models. We provide an historical framework to promote understanding of the development and diversification of fire regimes, covering the pre-human period, human domestication of fire, and the subsequent transition from subsistence agriculture to industrial economies. All of these phases still occur on Earth, providing opportunities for comparative research.

Although disturbances such as fire and native insects can contribute to natural dynamics of forest health, exceptional droughts, directly and in combination with other disturbance factors, are pushing some temperate forests beyond thresholds of sustainability. Interactions from increasing temperatures, drought, native insects and pathogens, and uncharacteristically severe wildfire are resulting in forest mortality beyond the levels of 20th-century experience. Additional anthropogenic stressors, such as atmospheric pollution and invasive species, further weaken trees in some regions. Although continuing climate change will likely drive many areas of temperate forest toward large-scale transformations, management actions can help ease transitions and minimize losses of socially valued ecosystem services.

Climate change is expected to drive increased tree mortality through drought, heat stress, and insect attacks, with manifold impacts on forest ecosystems. Yet, climate-induced tree mortality and biotic disturbance agents are largely absent from process-based ecosystem models. Using data sets from the western USA and associated studies, we present a framework for determining the relative contribution of drought stress, insect attack, and their interactions, which is critical for modeling mortality in future climates. We outline a simple approach that identifies the mechanisms associated with two guilds of insects - bark beetles and defoliators - which are responsible for substantial tree mortality. We then discuss cross-biome patterns of insect-driven tree mortality and draw upon available evidence contrasting the prevalence of insect outbreaks in temperate and tropical regions. We conclude with an overview of tools and promising avenues to address major challenges. Ultimately, a multitrophic approach that captures tree physiology, insect populations, and tree-insect interactions will better inform projections of forest ecosystem responses to climate change.

Abstract: Terrain ruggedness is often an important variable in wildlife habitat models. Most methods used to quantify ruggedness are indices derived from measures of slope and, as a result, are strongly correlated with slope. Using a Geographic Information System, we developed a vector ruggedness measure (VRM) of terrain based on a geomorphological method for measuring vector dispersion that is less correlated with slope. We examined the relationship of VRM to slope and to 2 commonly used indices of ruggedness in 3 physiographically different mountain ranges within the Mojave Desert of the southwestern United States. We used VRM, slope, distance to water, and springtime bighorn sheep ( Ovis canadensis nelsoni ) adult female locations to model sheep habitat in the 3 ranges. Using logistic regression, we determined that the importance of ruggedness in habitat selection remained consistent across mountain ranges, whereas the relative importance of slope varied according to the characteristic physiography of each range. Our results indicate that the VRM quantifies local variation in terrain more independently of slope than other methods tested, and that VRM and slope distinguish 2 different components of bighorn sheep habitat.

Summary Atmospheric carbon dioxide concentration ([CO 2 ]) is increasing, which increases leaf‐scale photosynthesis and intrinsic water‐use efficiency. These direct responses have the potential to increase plant growth, vegetation biomass, and soil organic matter; transferring carbon from the atmosphere into terrestrial ecosystems (a carbon sink). A substantial global terrestrial carbon sink would slow the rate of [CO 2 ] increase and thus climate change. However, ecosystem CO 2 responses are complex or confounded by concurrent changes in multiple agents of global change and evidence for a [CO 2 ]‐driven terrestrial carbon sink can appear contradictory. Here we synthesize theory and broad, multidisciplinary evidence for the effects of increasing [CO 2 ] (iCO 2 ) on the global terrestrial carbon sink. Evidence suggests a substantial increase in global photosynthesis since pre‐industrial times. Established theory, supported by experiments, indicates that iCO 2 is likely responsible for about half of the increase. Global carbon budgeting, atmospheric data, and forest inventories indicate a historical carbon sink, and these apparent iCO 2 responses are high in comparison to experiments and predictions from theory. Plant mortality and soil carbon iCO 2 responses are highly uncertain. In conclusion, a range of evidence supports a positive terrestrial carbon sink in response to iCO 2 , albeit with uncertain magnitude and strong suggestion of a role for additional agents of global change.

Tree mortality is a key factor influencing forest functions and dynamics, but our understanding of the mechanisms leading to mortality and the associated changes in tree growth rates are still limited. We compiled a new pan-continental tree-ring width database from sites where both dead and living trees were sampled (2970 dead and 4224 living trees from 190 sites, including 36 species), and compared early and recent growth rates between trees that died and those that survived a given mortality event. We observed a decrease in radial growth before death in ca. 84% of the mortality events. The extent and duration of these reductions were highly variable (1-100 years in 96% of events) due to the complex interactions among study species and the source(s) of mortality. Strong and long-lasting declines were found for gymnosperms, shade- and drought-tolerant species, and trees that died from competition. Angiosperms and trees that died due to biotic attacks (especially bark-beetles) typically showed relatively small and short-term growth reductions. Our analysis did not highlight any universal trade-off between early growth and tree longevity within a species, although this result may also reflect high variability in sampling design among sites. The intersite and interspecific variability in growth patterns before mortality provides valuable information on the nature of the mortality process, which is consistent with our understanding of the physiological mechanisms leading to mortality. Abrupt changes in growth immediately before death can be associated with generalized hydraulic failure and/or bark-beetle attack, while long-term decrease in growth may be associated with a gradual decline in hydraulic performance coupled with depletion in carbon reserves. Our results imply that growth-based mortality algorithms may be a powerful tool for predicting gymnosperm mortality induced by chronic stress, but not necessarily so for angiosperms and in case of intense drought or bark-beetle outbreaks.

Recent observations of elevated tree mortality following climate extremes, like heat and drought, raise concerns about climate change risks to global forest health. We currently lack both sufficient data and understanding to identify whether these observations represent a global trend toward increasing tree mortality. Here, we document events of sudden and unexpected elevated tree mortality following heat and drought events in ecosystems that previously were considered tolerant or not at risk of exposure. These events underscore the fact that climate change may affect forests with unexpected force in the future. We use the events as examples to highlight current difficulties and challenges for realistically predicting such tree mortality events and the uncertainties about future forest condition. Advances in remote sensing technology and greater availably of high-resolution data, from both field assessments and satellites, are needed to improve both understanding and prediction of forest responses to future climate change.

Top predators often have powerful direct effects on prey populations, but whether these direct effects propagate to the base of terrestrial food webs is debated. There are few examples of trophic cascades strong enough to alter the abundance and composition of entire plant communities. We show that the introduction of arctic foxes (Alopex lagopus) to the Aleutian archipelago induced strong shifts in plant productivity and community structure via a previously unknown pathway. By preying on seabirds, foxes reduced nutrient transport from ocean to land, affecting soil fertility and transforming grasslands to dwarf shrub/forb-dominated ecosystems.

Wildfires are often perceived as destructive disturbances, but we propose that when integrating evolutionary and socioecological factors, fires in most ecosystems can be understood as natural processes that provide a variety of benefits to humankind. Wildfires generate open habitats that enable the evolution of a diversity of shade‐intolerant plants and animals that have long benefited humans. There are many provisioning, regulating, and cultural services that people obtain from wildfires, and prescribed fires and wildfire management are tools for mimicking the ancestral role of wildfires in an increasingly populated world.

Dietary diversity often varies inversely with prey resource abundance. This pattern, although typically measured at the population level, is usually assumed to also characterize the behavior of individual animals within the population. However, the pattern might also be produced by changes in the degree of variation among individuals. Here we report on dietary and associated behavioral changes that occurred with the experimental translocation of sea otters from a food-poor to a food-rich environment. Although the diets of all individuals were broadly similar in the food-rich environment, a behaviorally based dietary polymorphism existed in the food-poor environment. Higher dietary diversity under low resource abundance was largely driven by greater variation among individuals. We further show that the dietary polymorphism in the food-poor environment included a broad suite of correlated behavioral variables and that the individuals that comprised specific behavioral clusters benefited from improved foraging efficiency on their individually preferred prey. Our findings add to the growing list of examples of extreme individuality in behavior and prey choice within populations and suggest that this phenomenon can emerge as a behavioral manifestation of increased population density. Individuality in foraging behavior adds complexity to both the fitness consequences of prey selection and food web dynamics, and it may figure prominently as a diversifying process over evolutionary timescales.

Forest structure and species composition in many western U.S. coniferous forests have been altered through fire exclusion, past and ongoing harvesting practices, and livestock grazing over the 20th century. The effects of these activities have been most pronounced in seasonally dry, low and mid-elevation coniferous forests that once experienced frequent, low to moderate intensity, fire regimes. In this paper, we report the effects of Fire and Fire Surrogate (FFS) forest stand treatments on fuel load profiles, potential fire behavior, and fire severity under three weather scenarios from six western U.S. FFS sites. This replicated, multisite experiment provides a framework for drawing broad generalizations about the effectiveness of prescribed fire and mechanical treatments on surface fuel loads, forest structure, and potential fire severity. Mechanical treatments without fire resulted in combined 1-, 10-, and 100-hour surface fuel loads that were significantly greater than controls at three of five FFS sites. Canopy cover was significantly lower than controls at three of five FFS sites with mechanical-only treatments and at all five FFS sites with the mechanical plus burning treatment; fire-only treatments reduced canopy cover at only one site. For the combined treatment of mechanical plus fire, all five FFS sites with this treatment had a substantially lower likelihood of passive crown fire as indicated by the very high torching indices. FFS sites that experienced significant increases in 1-, 10-, and 100-hour combined surface fuel loads utilized harvest systems that left all activity fuels within experimental units. When mechanical treatments were followed by prescribed burning or pile burning, they were the most effective treatment for reducing crown fire potential and predicted tree mortality because of low surface fuel loads and increased vertical and horizontal canopy separation. Results indicate that mechanical plus fire, fire-only, and mechanical-only treatments using whole-tree harvest systems were all effective at reducing potential fire severity under severe fire weather conditions. Retaining the largest trees within stands also increased fire resistance.

Mega‐fires are often defined according to their size and intensity but are more accurately described by their socioeconomic impacts. Three factors – climate change, fire exclusion, and antecedent disturbance, collectively referred to as the “mega‐fire triangle” – likely contribute to today's mega‐fires. Some characteristics of mega‐fires may emulate historical fire regimes and can therefore sustain healthy fire‐prone ecosystems, but other attributes decrease ecosystem resiliency. A good example of a program that seeks to mitigate mega‐fires is located in Western Australia, where prescribed burning reduces wildfire intensity while conserving ecosystems. Crown‐fire‐adapted ecosystems are likely at higher risk of frequent mega‐fires as a result of climate change, as compared with other ecosystems once subject to frequent less severe fires. Fire and forest managers should recognize that mega‐fires will be a part of future wildland fire regimes and should develop strategies to reduce their undesired impacts.

We provide a first detailed analysis of long-term, annual-resolution demographic trends in a temperate forest. After tracking the fates of 21,338 trees in a network of old-growth forest plots in the Sierra Nevada of California, we found that mortality rate, but not the recruitment rate, increased significantly over the 22 years of measurement (1983-2004). Mortality rates increased in both of two dominant taxonomic groups (Abies and Pinus) and in different forest types (different elevational zones). The increase in overall mortality rate resulted from an increase in tree deaths attributed to stress and biotic causes, and coincided with a temperature-driven increase in an index of drought. Our findings suggest that these forests (and by implication, other water-limited forests) may be sensitive to temperature-driven drought stress, and may be poised for die-back if future climates continue to feature rising temperatures without compensating increases in precipitation.

Summary 1. Deserts are one of the least invaded ecosystems by plants, possibly due to naturally low levels of soil nitrogen. Increased levels of soil nitrogen caused by atmospheric nitrogen deposition may increase the dominance of invasive alien plants and decrease the diversity of plant communities in desert regions, as it has in other ecosystems. Deserts should be particularly susceptible to even small increases in soil nitrogen levels because the ratio of increased nitrogen to plant biomass is higher compared with most other ecosystems. 2. The hypothesis that increased soil nitrogen will lead to increased dominance by alien plants and decreased plant species diversity was tested in field experiments using nitrogen additions at three sites in the in the Mojave Desert of western North America. 3. Responses of alien and native annual plants to soil nitrogen additions were measured in terms of density, biomass and species richness. Effects of nitrogen additions were evaluated during 2 years of contrasting rainfall and annual plant productivity. The rate of nitrogen addition was similar to published rates of atmospheric nitrogen deposition in urban areas adjacent to the Mojave Desert (3·2 g N m −2 year −1 ). The dominant alien species included the grasses Bromus madritensis ssp. rubens and Schismus spp. ( S. arabicus and S. barbatus ) and the forb Erodium cicutarium . 4. Soil nitrogen addition increased the density and biomass of alien annual plants during both years, but decreased density, biomass and species richness of native species only during the year of highest annual plant productivity. The negative response of natives may have been due to increased competitive stress for soil water and other nutrients caused by the increased productivity of aliens. 5. The effects of nitrogen additions were significant at both ends of a natural nutrient gradient, beneath creosote bush Larrea tridentata canopies and in the interspaces between them, although responses varied among individual alien species. The positive effects of nitrogen addition were highest in the beneath‐canopy for B. rubens and in interspaces for Schismus spp. and E. cicutarium . 6. The results indicated that increased levels of soil nitrogen from atmospheric nitrogen deposition or from other sources could increase the dominance of alien annual plants and possibly promote the invasion of new species in desert regions. Increased dominance by alien annuals may decrease the diversity of native annual plants, and increased biomass of alien annual grasses may also increase the frequency of fire. 7. Although nitrogen deposition cannot be controlled by local land managers, the managers need to understand its potential effects on plant communities and ecosystem properties, in particular how these effects may interact with land‐use activities that can be managed at the local scale. These interactions are currently unknown, and hinder the ability of managers to make appropriate land‐use decisions related to nitrogen deposition in desert ecosystems. 8. Synthesis and applications. The effects of nitrogen deposition on invasive alien plants should be considered when deciding where to locate new conservation areas, and in evaluating the full scope of ecological effects of new projects that would increase nitrogen deposition rates.

Abstract C 4 photosynthesis had a mid‐Tertiary origin that was tied to declining atmospheric CO 2 , but C 4 ‐dominated grasslands did not appear until late Tertiary. According to the ‘CO 2 ‐threshold’ model, these C 4 grasslands owe their origin to a further late Miocene decline in CO 2 that gave C 4 grasses a photosynthetic advantage. This model is most appropriate for explaining replacement of C 3 grasslands by C 4 grasslands, however, fossil evidence shows C 4 grasslands replaced woodlands. An additional weakness in the threshold model is that recent estimates do not support a late Miocene drop in p CO 2 . We hypothesize that late Miocene climate changes created a fire climate capable of replacing woodlands with C 4 grasslands. Critical elements were seasonality that sustained high biomass production part of year, followed by a dry season that greatly reduced fuel moisture, coupled with a monsoon climate that generated abundant lightning‐igniting fires. As woodlands became more open from burning, the high light conditions favoured C 4 grasses over C 3 grasses, and in a feedback process, the elevated productivity of C 4 grasses increased highly combustible fuel loads that further increased fire activity. This hypothesis is supported by paleosol data that indicate the late Miocene expansion of C 4 grasslands was the result of grassland expansion into more mesic environments and by charcoal sediment profiles that parallel the late Miocene expansion of C 4 grasslands. Many contemporary C 4 grasslands are fire dependent and are invaded by woodlands upon cessation of burning. Thus, we maintain that the factors driving the late Miocene expansion of C 4 were the same as those responsible for maintenance of C 4 grasslands today.