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Jon E. Keeley

Publications and source records attributed to Jon E. Keeley.

At least 91 records · Page 5Linked to original sources

Fire-adaptive trait evolution

Until relatively recently the importance of fire and the origin of fire-adaptive traits have received minimal attention from paleoecologists, and appreciation of this importance has varied across the different mediterranean-type climate (MTC) ecosystems. For example, Axelrod (1973) and Raven & Axelrod (1978) wrote extensive treatises on the origins of the California flora, and yet gave little or no mention to the issue of fire in the evolution of these taxa. Hopper (2009) suggests that fire has only been an incidental factor in the evolution of the Western Australian flora. These investigators have weighed climate and soils far above fire as an important evolutionary driver in these plant assemblages and have downplayed this component of community assembly (see Fig. 1.4). Axelrod (1989) even went so far as to suggest fire was irrelevant to the evolution of California chaparral. Although he acknowledged that fire could have played a role in the spread of chaparral-like vegetation during the late Tertiary (2–10 Ma), he insisted that fire had played no significant role in the origin of “adaptive types.” In his view, “Several lines of evidence suggest that the modern fire-adapted taxa may not reflect an evolutionary response to fire. The diverse adaptations to fire probably represent features that originated without the stimulus of fire. . .” Contrary to this belief, we suggest there is sufficient reason to accept a fire origin for many fire-adaptive traits in mediterranean-type vegetation (MTV), and that fire has been a potential ecosystem process on landscapes far longer than the late Tertiary (Bowman et al . 2009; Pausas & Keeley 2009).

Book chapter

Fire and the origins of Mediterranean-type vegetation

The mediterranean-type climate (MTC) is widely agreed to have been in place in all five MTC regions since at least the late Pliocene (see Fig. 9.1), ~2 Ma, with much of the contemporary mediterranean-type vegetation (MTV) present and contributing to a highly fire-prone environment. There is far less agreement on: (1) the timing of the origin of the MTC, (2) the timing of and factors responsible for the origins of MTV, and (3) the paleohistory of fire and extent to which it has played a role in the origins of MTV. Ample evidence exists to suggest a much earlier origin of MTC and MTV. A widely held paradigm is that many of the woody sclerophylls that comprise MTV are much older than the Pliocene and thus have not adapted to contemporary fire-prone MTC conditions (Axelrod 1989; Herrera 1992; Verdú et al . 2003; Ackerly 2004a). Most of these have origins in the Tertiary Period of the early Cenozoic and are viewed as relictual taxa that represent evolutionary inertia and are present today merely by chance avoidance of random extinctions.

Book chapter

Alien species and fire

A large diversity of alien plants is found in most mediterranean-type climate (MTC) regions and fire is sometimes closely linked to their ability to invade natural ecosystems. This is a concern because aliens often upset natural ecosystem processes, and thus are a major management concern. These five regions not only differ in their contributions of non-native plant species to other regions, but also vary in their susceptibility to invasion by alien species, something often referred to as a community's invasibility. Fire is a key factor behind plant invasions into natural plant communities and particularly critical is the timing of propagule availability and characteristics of the fire regime. Fire also interacts with geology in dictating functional types that become pernicious invasive problems. For example, on coarse-textured low-fertility soils in two of the southern hemisphere MTC regions, shrubs and trees are among the most aggressive invasives, and are capable of invading seemingly undisturbed intact shrublands. However, on more fertile soils such as in California and Chile, grasses and other herbaceous species are bigger threats, but invasion typically requires disturbance and under some circumstances fire can effect type conversion from woody vegetation to alien-dominated grasslands.

Book chapter

Fire management of Mediterranean landscapes

The hazardous mediterranean climate, highly flammable vegetation, and rugged terrain, all important elements of fire behavior, become problems only in the presence of people. People recreate and build homes in the mediterranean wildlands because of the delightful climate and will continue to do so as long as space is available. People start most fires, and their mere presence tends to warp fire suppression strategies because fire agencies must protect lives and property threatened by fires rather than “back off” and build fire lines around fire perimeters . Carl C. Wilson (1979a), Chief of Division of Forest and Fire Research, USFS/Pacific Southwest Forest and Range Experiment Station Human presence in mediterranean-type climate (MTC) regions has differed markedly in the length of human occupation; however, there are remarkable similarities in how early inhabitants altered fire regimes and how modern societies deal with the fire hazard. Here we draw on the history of human impacts outlined in the regional reviews (see Chapters 4–8), the problems created by nineteenth and twentieth century management practices, and conclude with twenty-first century problems and future options. As discussed throughout this book, MTC ecosystems are highly fire adapted but, as illustrated here, contemporary societies have not fully adapted to balancing fire hazard risk and resource needs on these landscapes. Early Human Fire Use and Impacts Fire has been a widely utilized management tool throughout the history of humankind (Pyne 1995). Early hunter–gatherers utilized fire to manage for plant and animal resources. Fire also played an important role in early domestication of crops as clearing off woody or other perennial vegetation would have required fire on many landscapes. With domestication of livestock it was an important tool for increasing forage.

Book chapter

Climate, Fire and Geology in the Convergence of Mediterranean-type Climate Ecosystems

Integrating Climate, Fire and Geology in a Fire-prone World Fire challenges the long-standing hegemony of ecology, biogeography and paleoecology that climate and soils are sufficient to explain the origin and distribution of plant species. In a world where half of the land surface is fire-prone (Krawchuk et al . 2009), understanding the past and predicting the future requires a close integration of climate, fire and geology. The dogma that fire is an anthropogenic phenomenon of little use in understanding paleoecology (Axelrod 1980, 1989), or merely incidental to vegetation development (Hopper 2009), is rapidly being replaced with a better understanding of paleofire's impact on land plant evolution (Scott 2000; Pausas & Keeley 2009). Attempts to model future global vegetation patterns have been demonstrated to be inadequate without including both natural and anthropogenic fire regimes (Bond et al . 2005). Bond and Keeley (2005) outlined the conundrum posed by alternative explanations for the present distribution of vegetation and assembly of communities. Classical explanations have invoked resource-based mechanisms that are driven by climate and soils. There are ecosystems where resource-based mechanisms may be sufficient, but on many seasonally dry landscapes ecosystem processes such as fire play a major role in the organization and evolution of vegetation.

Book chapter

Factors affecting fuel break effectiveness in the control of large fires on the Los Padres National Forest, California

As wildfires have increased in frequency and extent, so have the number of homes developed in the wildland-urban interface. In California, the predominant approach to mitigating fire risk is construction of fuel breaks, but there has been little empirical study of their role in controlling large fires.We constructed a spatial database of fuel breaks on the Los Padres National Forest in southern California to better understand characteristics of fuel breaks that affect the behaviour of large fires and to map where fires and fuel breaks most commonly intersect. We evaluated whether fires stopped or crossed over fuel breaks over a 28-year period and compared the outcomes with physical characteristics of the sites, weather and firefighting activities during the fire event. Many fuel breaks never intersected fires, but others intersected several, primarily in historically fire-prone areas. Fires stopped at fuel breaks 46% of the time, almost invariably owing to fire suppression activities. Firefighter access to treatments, smaller fires and longer fuel breaks were significant direct influences, and younger vegetation and fuel break maintenance indirectly improved the outcome by facilitating firefighter access. This study illustrates the importance of strategic location of fuel breaks because they have been most effective where they provided access for firefighting activities.

California

Estimating aboveground biomass for broadleaf woody plants and young conifers in Sierra Nevada, California forests

Quantification of biomass is fundamental to a wide range of research and natural resource management goals. An accurate estimation of plant biomass is essential to predict potential fire behavior, calculate carbon sequestration for global climate change research, assess critical wildlife habitat, and so forth. Reliable allometric equations from simple field measurements are necessary for efficient evaluation of plant biomass. However, allometric equations are not available for many common woody plant taxa in the Sierra Nevada. In this report, we present more than 200 regression equations for the Sierra Nevada western slope that relate crown diameter, plant height, crown volume, stem diameter, and both crown diameter and height to the dry weight of foliage, branches, and entire aboveground biomass. Destructive sampling methods resulted in regression equations that accurately predict biomass from one or two simple, nondestructive field measurements. The tables presented here will allow researchers and natural resource managers to easily choose the best equations to fit their biomass assessment needs.

California

The Origins of C4 Grasslands: Integrating Evolutionary and Ecosystem Science

The evolution of the C 4 photosynthetic pathway from the ancestral C 3 pathway in grasses led to the establishment of grasslands in warm climates during the Late Miocene (8 to 3 million years ago). This was a major event in plant evolutionary history, and their high rates of foliage production sustained high levels of herbivore consumption. The past decade has seen significant advances in understanding C 4 grassland ecosystem ecology, and now a wealth of data on the geological history of these ecosystems has accumulated and the phylogeny of grasses is much better known. Edwards et al. (p. 587) review this multidisciplinary research area and attempt to synthesize emerging knowledge about the evolution of grass species within the context of plant and ecosystem ecology.

Science

Stem demography and post-fire recruitment of a resprouting serotinous conifer

The contribution of resprouts and seedling recruitment to post-fire regeneration of the South African fynbos conifer Widdringtonia nodiflora was compared eight months after wildfires in 1990. Stems on all trees were killed by fire but resprouting success was > 90 % at all but one site. A demographic study of burned skeletons revealed that prior to these fires, nearly all plants were multi-stemmed (4–9 stems/plant) and multi-aged, indicating continuous sprout production between fires. All stems were killed by these 1990 fires and at most sites > 90 % of the stems were burned to ground level. All diameter stems were susceptible to such incineration as, at most sites, there was no difference in average diameter of stems burned to ground level and those left standing. Individual genets usually had all ramets incinerated to ground level or all ramets charred, but intact, suggesting certain micro-sites burned hotter, whereas other sites were somewhat protected. Although not true of the 1990 fires, there was evidence that occasionally Widdring-tonia stems may survive fire. At one site, four of the 16 plants sampled had a burned stem twice as old as the oldest burned stem on the other 12 plants at the site, suggesting some stems had survived the previous fire (ca. 1970) and this conclusion was supported by fire-scars on these four stems that dated to ca. 1970. Based on the highly significant correlation between stem diameter and cone density left standing after the 1990 fires, we calculated that for most sites > 80 % of the initial cone crop was incinerated by fire. This is important because we observed a strong relationship between size of the canopy cone crop surviving fire and post-fire seedling recruitment. Under these conditions we hypothesize that sprouting confers a selective advantage to genets when fires cause heavy losses of seed. The infrequent occurrence of sprouting in the Cupressaceae suggests the hypothesis that resprouting is an apomorphic or derived trait in Widdringtonia. Data from this study suggests resprouting provides a selective advantage under severe fynbos fires, which are not only 'stand-replacing fires,’but also are intense enough to incinerate cone-bearing stems.

Journal of Vegetation Science

Fire in the Earth system

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.

Science

A burning story: The role of fire in the history of life

Ecologists, biogeographers, and paleobotanists have long thought that climate and soils controlled the distribution of ecosystems, with the role of fire getting only limited appreciation. Here we review evidence from different disciplines demonstrating that wildfire appeared concomitant with the origin of terrestrial plants and played an important role throughout the history of life. The importance of fire has waxed and waned in association with changes in climate and paleoatmospheric conditions. Well before the emergence of humans on Earth, fire played a key role in the origins of plant adaptations as well as in the distribution of ecosystems. Humans initiated a new stage in ecosystem fire, using it to make the Earth more suited to their lifestyle. However, as human populations have expanded their use of fire, their actions have come to dominate some ecosystems and change natural processes in ways that threaten the sustainability of some landscapes. ?? 2009 by American Institute of Biological Sciences.

BioScience

The national Fire and Fire Surrogate study: Effects of fuel reduction methods on forest vegetation structure and fuels

Changes in vegetation and fuels were evaluated from measurements taken before and after fuel reduction treatments (prescribed fire, mechanical treatments, and the combination of the two) at 12 Fire and Fire Surrogate (FFS) sites located in forests with a surface fire regime across the conterminous United States. To test the relative effectiveness of fuel reduction treatments and their effect on ecological parameters we used an informationtheoretic approach on a suite of 12 variables representing the overstory (basal area and live tree, sapling, and snag density), the understory (seedling density, shrub cover, and native and alien herbaceous species richness), and the most relevant fuel parameters for wildfire damage (height to live crown, total fuel bed mass, forest floor mass, and woody fuel mass). In the short term (one year after treatment), mechanical treatments were more effective at reducing overstory tree density and basal area and at increasing quadratic mean tree diameter. Prescribed fire treatments were more effective at creating snags, killing seedlings, elevating height to live crown, and reducing surface woody fuels. Overall, the response to fuel reduction treatments of the ecological variables presented in this paper was generally maximized by the combined mechanical plus burning treatment. If the management goal is to quickly produce stands with fewer and larger diameter trees, less surface fuel mass, and greater herbaceous species richness, the combined treatment gave the most desirable results. However, because mechanical plus burning treatments also favored alien species invasion at some sites, monitoring and control need to be part of the prescription when using this treatment. ?? 2009 by the Ecological Society of America.

Ecological Applications

Fire treatment effects on vegetation structure, fuels, and potential fire severity in western U.S. forests

Abstract. 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. ?? 2009 by the Ecological Society of America.

Ecological Applications

Large, high-intensity fire events in Southern California shrublands: Debunking the fine-grain age patch model

We evaluate the fine-grain age patch model of fire regimes in southern California shrublands. Proponents contend that the historical condition was characterized by frequent small to moderate size, slow-moving smoldering fires, and that this regime has been disrupted by fire suppression activities that have caused unnatural fuel accumulation and anomalously large and catastrophic wildfires. A review of more than 100 19th-century newspaper reports reveals that large, high-intensity wildfires predate modern fire suppression policy, and extensive newspaper coverage plus first-hand accounts support the conclusion that the 1889 Santiago Canyon Fire was the largest fire in California history. Proponents of the fine-grain age patch model contend that even the very earliest 20th-century fires were the result of fire suppression disrupting natural fuel structure. We tested that hypothesis and found that, within the fire perimeters of two of the largest early fire events in 1919 and 1932, prior fire suppression activities were insufficient to have altered the natural fuel structure. Over the last 130 years there has been no significant change in the incidence of large fires greater than 10000 ha, consistent with the conclusion that fire suppression activities are not the cause of these fire events. Eight megafires (???50 000 ha) are recorded for the region, and half have occurred in the last five years. These burned through a mosaic of age classes, which raises doubts that accumulation of old age classes explains these events. Extreme drought is a plausible explanation for this recent rash of such events, and it is hypothesized that these are due to droughts that led to increased dead fine fuels that promoted the incidence of firebrands and spot fires. A major shortcoming of the fine-grain age patch model is that it requires age-dependent flammability of shrubland fuels, but seral stage chaparral is dominated by short-lived species that create a dense surface layer of fine fuels. Results from the Behave Plus fire model with a custom fuel module for young chaparral shows that there is sufficient dead fuel to spread fire even under relatively little winds. Empirical studies of fuel ages burned in recent fires illustrate that young fuels often comprise a major portion of burned vegetation, and there is no difference between evergreen chaparral and semi-deciduous sage scrub. It has also been argued that the present-day fire size distribution in northern Baja California is a model of the historical patterns that were present on southern California landscapes. Applying this model with historical fire frequencies shows that the Baja model is inadequate to maintain these fire-prone ecosystems and further demonstrates that fire managers in southern California are not likely to learn much from studying modern Baja California fire regimes. Further supporting this conclusion are theoretical cellular automata models of fire spread, which show that, even in systems with age dependent flammability, landscapes evolve toward a complex age mosaic with a plausible age structure only when there is a severe stopping rule that constrains fire size, and only if ignitions are saturating. ?? 2009 by the Ecological Society of America.

Ecological Applications

The 2007 southern California wildfires: Lessons in complexity

The 2007 wildfire season in southern California burned over 1,000,000 ac (∼400,000 ha) and included several megafires. We use the 2007 fires as a case study to draw three major lessons about wildfires and wildfire complexity in southern California. First, the great majority of large fires in southern California occur in the autumn under the influence of Santa Ana windstorms. These fires also cost the most to contain and cause the most damage to life and property, and the October 2007 fires were no exception because thousands of homes were lost and seven people were killed. Being pushed by wind gusts over 100 kph, young fuels presented little barrier to their spread as the 2007 fires reburned considerable portions of the area burned in the historic 2003 fire season. Adding to the size of these fires was the historic 2006‐2007 drought that contributed to high dead fuel loads and long distance spotting. As in 2003, young chaparral stands and fuel treatments were not reliable barriers to fire in October 2007. Second, the Zaca Fire in July and August 2007 showed that other factors besides high winds can sometimes combine to create conditions for large fires in southern California. Spring and summer fires in southern California chaparral are usually easily contained because of higher fuel moisture and the general lack of high winds. However, the Zaca Fire burned in a remote wilderness area of rugged terrain that made access difficult. In addition, because of its remoteness, anthropogenic ignitions have been low and stand age and fuel loads were high. Coupled with this was severe drought that year that generated fuel moisture levels considerably below normal for early summer. A third lesson comes from 2007 conifer forest fires in the southern California mountains. In contrast to lower elevation chaparral, fire suppression has led to major increases in conifer forest fuels that can lead to unnaturally severe fires when ignitions escape control. The Slide and Grass Valley Fires of October 2007 occurred in forests that had been subject to extensive fuel treatment, but fire control was complicated by a patchwork of untreated private properties and mountain homes built of highly flammable materials. In a fashion reminiscent of other recent destructive conifer fires in California, burning homes themselves were a major source of fire spread. These lessons suggest that the most important advances in fire safety in this region are to come from advances in fire prevention, fire preparedness, and land-use planning that includes fire hazard patterns.

California

Biogeochemical legacy of prescribed fire in a giant sequoia - Mixed conifer forest: A 16-year record of watershed balances

The effects of prescription burning on watershed balances of major ions in mixed conifer forest were examined in a 16-year paired catchment study in Sequoia National Park, California. The objective was to determine whether fire-related changes in watershed balances persist as long as estimated low-end natural fire-return intervals (???10 years), and whether cumulative net export caused by fire could deplete nutrient stocks between successive fires. Inputs (wet + dry deposition) and outputs (stream export) of N, S, Cl-, HCO3-, Ca2+, Mg2+, Na+, K+, H+, and SiO2 were measured for 7 years preceding, and 9 years following, a prescribed burn of one of the catchments. After fire, runoff coefficients increased by 7% (in dry years) to 35% (in wet years). Inorganic N was elevated in stream water for 3 years after fire. Increased export of water, SO42-, Cl-,SiO2, and base cations continued through the end of the study. Pools and processes attributed to fire led to the cumulative loss, per hectare, of 1.2 kg N, 16 kg S, 25 kg Cl-, 130 kg Ca2+, 19 kg Mg2+, 71 kg Na+, 29 kg K+ and 192 kg Si, above that predicted by prefire regression equations relating export in the paired catchments. This additional export equaled <1% of the N, up to one-third of the Ca and Mg, and up to three-fourths of the K, contained in the forest floor prior to combustion. Changes in watershed balances indicated that low-end natural fire-return intervals may prevent complete reaccumulation of several elements between fires. Copyright 2008 by the American Geophysical Union.

Journal of Geophysical Research G: Biogeosciences

Fire severity and ecosytem responses following crown fires in California shrublands

Chaparral shrublands burn in large high-intensity crown fires. Managers interested in how these wildfires affect ecosystem processes generally rely on surrogate measures of fire intensity known as fire severity metrics. In shrublands burned in the autumn of 2003, a study of 250 sites investigated factors determining fire severity and ecosystem responses. Using structural equation modeling we show that stand age, prefire shrub density, and the shortest interval of the prior fire history had significant direct effects on fire severity, explaining >50% of the variation in severity. Fire severity per se is of interest to resource managers primarily because it is presumed to be an indicator of important ecosystem processes such as vegetative regeneration, community recovery, and erosion. Fire severity contributed relatively little to explaining patterns of regeneration after fire. Two generalizations can be drawn: fire severity effects are mostly short-lived, i.e., by the second year they are greatly diminished, and fire severity may have opposite effects on different functional types. Species richness exhibited a negative relationship to fire severity in the first year, but fire severity impacts were substantially less in the second postfire year and varied by functional type. Much of this relationship was due to alien plants that are sensitive to high fire severity; at all scales from 1 to 1000 m 2 , the percentage of alien species in the postfire flora declined with increased fire severity. Other aspects of disturbance history are also important determinants of alien cover and richness as both increased with the number of times the site had burned and decreased with time since last fire. A substantial number of studies have shown that remote-sensing indices are correlated with field measurements of fire severity. Across our sites, absolute differenced normalized burn ratio (dNBR) was strongly correlated with field measures of fire severity and with fire history at a site but relative dNBR was not. Despite being correlated with fire severity, absolute dNBR showed little or no relationship with important ecosystem responses to wildfire such as shrub resprouting or total vegetative regeneration. These findings point to a critical need for further research on interpreting remote sensing indices as applied to postfire management of these shrublands.

Ecological Applications