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Geology topics

Jon E. Keeley

Publications and source records attributed to Jon E. Keeley.

At least 127 records · Page 7Linked to original sources
USGS2005

Fire and the Miocene expansion of C4 grasslands

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.

Ecology Letters↗
USGS2005

Fuel reduction and woody debris dynamics with early season and late season prescribed fire in a Sierra Nevada mixed conifer forest

Fire exclusion has led to an unnatural accumulation and greater spatial continuity of organic material on the ground in many forests. This material serves both as potential fuel for forest fires and habitat for a large array of forest species. Managers must balance fuel reduction to reduce wildfire hazard with fuel retention targets to maintain other forest functions. This study reports fuel consumption and changes to coarse woody debris attributes with prescribed burns ignited under different fuel moisture conditions. Replicated early season burn, late season burn, and unburned control plots were established in old-growth mixed conifer forest in Sequoia National Park that had not experienced fire for more than 120 years. Early season burns were ignited during June 2002 when fuels were relatively moist, and late season burns were ignited during September/October 2001 when fuels were dry. Fuel loading and coarse woody debris abundance, cover, volume, and mass were evaluated prior to and after the burns. While both types of burns reduced fuel loading, early season burns consumed significantly less of the total dead and down organic matter than late season burns (67% versus 88%). This difference in fuel consumption between burning treatments was significant for most all woody fuel components evaluated, plus the litter and duff layers. Many logs were not entirely consumed – therefore the number of logs was not significantly changed by fire – but burning did reduce log length, cover, volume, and mass. Log cover, volume, and mass were reduced to a lesser extent by early season burns than late season burns, as a result of higher wood moisture levels. Early season burns also spread over less of the ground surface within the burn perimeter (73%) than late season burns (88%), and were significantly patchier. Organic material remaining after a fire can dam sediments and reduce erosion, while unburned patches may help mitigate the impact of fire on fire-sensitive species by creating refugia from which these species can recolonize burned areas. Early season burns may be an effective means of moderating potential ecosystem damage when treating heavy and/or continuous fuels resulting from long periods of fire exclusion, if burning during this season is not detrimental to other forest functions.

California↗
USGS2005

Fire as a global ‘herbivore’: the ecology and evolution of flammable ecosystems

It is difficult to find references to fire in general textbooks on ecology, conservation biology or biogeography, in spite of the fact that large parts of the world burn on a regular basis, and that there is a considerable literature on the ecology of fire and its use for managing ecosystems. Fire has been burning ecosystems for hundreds of millions of years, helping to shape global biome distribution and to maintain the structure and function of fire-prone communities. Fire is also a significant evolutionary force, and is one of the first tools that humans used to re-shape their world. Here, we review the recent literature, drawing parallels between fire and herbivores as alternative consumers of vegetation. We point to the common questions, and some surprisingly different answers, that emerge from viewing fire as a globally significant consumer that is analogous to herbivory.

Trends in Ecology and Evolution↗
USGS2005

Fire history of the San Francisco East Bay region and implications for landscape patterns

The San Francisco East Bay landscape is a rich mosaic of grasslands, shrublands and woodlands that is experiencing losses of grassland due to colonization by shrubs and succession towards woodland associations. The instability of these grasslands is apparently due to their disturbance-dependent nature coupled with 20th century changes in fire and grazing activity. This study uses fire history records to determine the potential for fire in this region and for evidence of changes in the second half of the 20th century that would account for shrubland expansion. This region has a largely anthropogenic fire regime with no lightning-ignited fires in most years. Fire suppression policy has not excluded fire from this region; however, it has been effective at maintaining roughly similar burning levels in the face of increasing anthropogenic fires, and effective at decreasing the size of fires. Fire frequency parallels increasing population growth until the latter part of the 20th century, when it reached a plateau. Fire does not appear to have been a major factor in the shrub colonization of grasslands, and cessation of grazing is a more likely immediate cause. Because grasslands are not under strong edaphic control, rather their distribution appears to be disturbance-dependent, and natural lightning ignitions are rare in the region, I hypothesize that, before the entrance of people into the region, grasslands were of limited extent. Native Americans played a major role in creation of grasslands through repeated burning and these disturbance-dependent grasslands were maintained by early European settlers through overstocking of these range lands with cattle and sheep. Twentieth century reduction in grazing, coupled with a lack of natural fires and effective suppression of anthropogenic fires, have acted in concert to favor shrubland expansion.

International Journal of Wildland Fire↗
USGS2005

Seed germination of Sierra Nevada postfire chaparral species

The California chaparral community has a rich flora of species with different mechanisms for cuing germination to postfire conditions. Here we report further germination experiments that elucidate the response of several widespread shrub species whose germination response was not clear and include other species from the Sierra Nevada, which have not previously been included in germination studies. The shrubs Adenostoma fasciculatum and Eriodictyon crassifolium and the postfire annual Mentzelia dispersa exhibited highly significant germination in response to smoke treatments, with some enhanced germination in response to heating as well. The shrubs Fremontodendron californicum and Malacothamnus fremontii were stimulated only by heat-shock treatments. Seeds buried in the soil for one year exhibited substantially higher germination for controls and most treatments. In the case of two postfire annuals, Mimulus bolanderi and M. gracilipes , germination of fresh seed was significantly greater with smoke or heating but after soil storage, over two-thirds of the control seeds germinated and treatment effects were not significant. These two annuals are generally restricted to postfire conditions and it is suggested that control germination of soil-stored seed may be a light-response (which was not tested here) as previously reported for another chaparral species in that genus.

Madroño↗
USGS2004

Ecological impacts of wheat seeding after a Sierra Nevada wildfire

The Highway Fire burned 1680 ha of mixed ponderosa pine–oak–chaparral in the newly created Giant Sequoia National Monument and the adjacent Sequoia National Forest of Fresno County, California in August 2001. The USDA Forest Service Burned Area Emergency Rehabilitation (BAER) program recommended that portions of the burned forest be seeded with a non-persistent variety of wheat at a density of 157 kg ha –1 (140 lb/ac). The present study compared the vascular plant diversity and cover in seeded and unseeded parts of this burn to evaluate the ecological impact of seeding an alien grass. In the first post-fire growing season, the natural regeneration of unseeded control sites averaged ~55% ground surface covered. Wheat seeding enhanced the ground cover, averaging 95% ground surface cover. Wheat was the dominant species on the seeded sites, comprising 67% of the total cover. Dominance–diversity curves were markedly affected by the seeding and indicated a disruption in the natural ecological structure of these communities. On seeded sites, wheat dominated and all other species were poorly represented whereas, on unseeded control sites, there was a more equitable distribution of species. Correlated with the wheat cover was a significant decrease in species richness at all scales examined. Total species richness was reduced from 152 species across all unseeded sites to 104 species on all seeded sites. Average species richness, at scales from 1 to 1000 m 2 , was 30–40% lower on seeded sites. Species most strongly inhibited were post-fire endemics whose lifecycle is restricted to immediate post-fire environments. Seeded sites had fewer alien species than unseeded sites; however, this may not have any lasting effect since other studies show the primary alien threat is not in the first post-fire year. Seeding was also associated with an order of magnitude drop in Pinus ponderosa seedling recruitment and, coupled with the massive thatch still remaining on the site, it is likely that recruitment will be inhibited in subsequent years.

International Journal of Wildland Fire↗
USGS2004

Impact of antecedent climate on fire regimes in coastal California

Severe fire weather is a major determinant of fire size in coastal California; however, it is unclear to what extent antecedent climate also controls fire activity. This study investigates the relationship between fire activity and climate in central coastal and southern California. Climate variables included the Palmer Drought Severity Index (PDSI), total monthly precipitation, mean monthly maximum temperature and the autumn and winter Southern Oscillation Indices (SOI). For both the central coast and the south coast regions there was no significant relationship between growing season PDSI, precipitation or temperature and number of fires. When examined by season, summer temperatures were positively correlated with number of fires in the central coast and autumn PDSI and precipitation were negatively correlated with fire occurrence in the south coast region. Area burned was not correlated with any current year climate variables in southern California although, in the central coast, drought during spring and autumn were correlated, but explained less than 10% of the variation in the area burned. Although there was a modest relationship between the Southern Oscillation Index (SOI) and local climate parameters, there was only a relatively weak relationship with fire activity. The importance of autumn foehn winds is illustrated by the observation that large fires occur most commonly during the autumn, regardless of PDSI. Antecedent climate, however, does appear to play some role in determining the length of the fire season on these landscape as PDSI is consistently related to the occurrence of large fires that occur before or after the autumn months.

International Journal of Wildland Fire↗
USGS2004

Effects of an introduced pathogen and fire exclusion on the demography of sugar pine

An introduced pathogen, white pine blister rust ( Cronartium ribicola ), has caused declines in five-needled pines throughout North America. Simultaneously, fire exclusion has resulted in dense stands in many forest types, which may create additional stress for these generally shade-intolerant pines. Fire exclusion also allows fuels to accumulate, and it is unclear how affected populations will respond to the reintroduction of fire. Although white pine blister rust and fire exclusion are widely recognized threats, long-term demographic data that document the effects of these stressors are rare. We present population trends from 2168 individuals over 5–15 years for an affected species, sugar pine ( Pinus lambertiana ), at several burned and unburned sites in the Sierra Nevada of California. Size-based matrix models indicate that most unburned populations have negative growth rates (λ range: 0.82–1.04). The growth rate of most populations was, however, indistinguishable from replacement levels (λ = 1.0), implying that, if populations are indeed declining, the progression of any such decline is slow, and longer observations are needed to clearly determine population trends. We found significant differences among population growth rates, primarily due to variation in recruitment rates. Deaths associated with blister rust and stress (i.e., resource competition) were common, suggesting significant roles for both blister rust and fire exclusion in determining population trajectories. Data from 15 prescribed fires showed that the immediate effect of burning was the death of many small trees, with the frequency of mortality returning to pre-fire levels within five years. In spite of a poor prognosis for sugar pine, our results suggest that we have time to apply and refine management strategies to protect this species.

Ecological Applications↗
USGS2004

VTM plots as evidence of historical change: Goldmine or landmine?

VTM (Vegetation Type Map) plots comprise a huge data set on vegetation composition for many parts of California collected mostly between 1929 and 1935. Historical changes in vegetation have been inferred by sampling these areas many decades later and evaluating the changes in plant dominance. VTM plots can not be precisely relocated, and it has been assumed that errors resulting from this problem are inconsequential or can be eliminated by comparison with a composite of multiple contemporary plots. This study examines that assumption for southern California shrubland landscapes by comparing the differences in species composition between closely positioned VTM-sized plots. Comparing shrub species density in 400-m² plots separated by 30 m (center to center), I found that all species exhibited considerable differences in density even over this short distance. This patchiness in shrub distribution could lead to major errors in historical reconstructions from VTM plot data. Two methods are proposed for dealing with this problem. One is to collect multiple samples from the vicinity of the VTM plot and use the observed spatial variation to set bounds on the temporal changes required to represent significant historical change. The other is to look at broad landscape changes reflected in the averages observed in a large sampling of sites.

Madroño↗
USGS2003

Species area relationships in mediterranean-climate plant communities

Aim To determine the best-fit model of species–area relationships for Mediterranean-type plant communities and evaluate how community structure affects these species–area models. Location Data were collected from California shrublands and woodlands and compared with literature reports for other Mediterranean-climate regions. Methods The number of species was recorded from 1, 100 and 1000 m 2 nested plots. Best fit to the power model or exponential model was determined by comparing adjusted r 2 values from the least squares regression, pattern of residuals, homoscedasticity across scales, and semi-log slopes at 1–100 m 2 and 100–1000 m 2 . Dominance–diversity curves were tested for fit to the lognormal model, MacArthur's broken stick model, and the geometric and harmonic series. Results Early successional Western Australia and California shrublands represented the extremes and provide an interesting contrast as the exponential model was the best fit for the former, and the power model for the latter, despite similar total species richness. We hypothesize that structural differences in these communities account for the different species–area curves and are tied to patterns of dominance, equitability and life form distribution. Dominance–diversity relationships for Western Australian heathlands exhibited a close fit to MacArthur's broken stick model, indicating more equitable distribution of species. In contrast, Californian shrublands, both postfire and mature stands, were best fit by the geometric model indicating strong dominance and many minor subordinate species. These regions differ in life form distribution, with annuals being a major component of diversity in early successional Californian shrublands although they are largely lacking in mature stands. Both young and old Australian heathlands are dominated by perennials, and annuals are largely absent. Inherent in all of these ecosystems is cyclical disequilibrium caused by periodic fires. The potential for community reassembly is greater in Californian shrublands where only a quarter of the flora resprout, whereas three quarters resprout in Australian heathlands. Other Californian vegetation types sampled include coniferous forests, oak savannas and desert scrub, and demonstrate that different community structures may lead to a similar species–area relationship. Dominance–diversity relationships for coniferous forests closely follow a geometric model whereas associated oak savannas show a close fit to the lognormal model. However, for both communities, species–area curves fit a power model. The primary driver appears to be the presence of annuals. Desert scrub communities illustrate dramatic changes in both species diversity and dominance–diversity relationships in high and low rainfall years, because of the disappearance of annuals in drought years. Main conclusions Species–area curves for immature shrublands in California and the majority of Mediterranean plant communities fit a power function model. Exceptions that fit the exponential model are not because of sampling error or scaling effects, rather structural differences in these communities provide plausible explanations. The exponential species–area model may arise in more than one way. In the highly diverse Australian heathlands it results from a rapid increase in species richness at small scales. In mature California shrublands it results from very depauperate richness at the community scale. In both instances the exponential model is tied to a preponderance of perennials and paucity of annuals. For communities fit by a power model, coefficients z and log c exhibit a number of significant correlations with other diversity parameters, suggesting that they have some predictive value in ecological communities.

Journal of Biogeography↗
USGS2003

Evolution of CAM and C 4 carbon-concentrating mechanisms

Mechanisms for concentrating carbon around the Rubisco enzyme, which drives the carbon-reducing steps in photosynthesis, are widespread in plants; in vascular plants they are known as crassulacean acid metabolism (CAM) and C 4 photosynthesis. CAM is common in desert succulents, tropical epiphytes, and aquatic plants and is characterized by nighttime fixation of CO2. The proximal selective factor driving the evolution of this CO2-concentrating pathway is low daytime CO2, which results from the unusual reverse stomatal behavior of terrestrial CAM species or from patterns of ambient CO2 availability for aquatic CAM species. In terrestrials the ultimate selective factor is water stress that has selected for increased water use efficiency. In aquatics the ultimate selective factor is diel fluctuations in CO2 availability for palustrine species and extreme oligotrophic conditions for lacustrine species. C4 photosynthesis is based on similar biochemistry but carboxylation steps are spatially separated in the leaf rather than temporally as in CAM. This biochemical pathway is most commonly associated with a specialized leaf anatomy known as Kranz anatomy; however, there are exceptions. The ultimate selective factor driving the evolution of this pathway is excessively high photorespiration that inhibits normal C3 photosynthesis under high light and high temperature in both terrestrial and aquatic habitats. CAM is an ancient pathway that likely has been present since the Paleozoic era in aquatic species from shallow-water palustrine habitats. While atmospheric CO2 levels have undoubtedly affected the evolution of terrestrial plant carbon-concentrating mechanisms, there is reason to believe that past atmospheric changes have not played as important a selective role in the aquatic milieu since palustrine habitats today are not generally carbon sinks, and the selective factors driving aquatic CAM are autogenic. Terrestrial CAM, in contrast, is of increasing selective value under extreme water deficits, and undoubtedly, high Mesozoic CO2 levels reduced the amount of landscape perceived by plants as water limited. Late Tertiary and Quaternary reductions in atmospheric CO2, coupled with increasing seasonality, were probably times of substantial species radiation and ecological expansion for CAM plants. C4 photosynthesis occurs in only about half as many families as CAM, and three-fourths of C4 species are either grasses or sedges. Molecular phylogenies indicate C4 is a more recent innovation than CAM and that it originated in the mid-Tertiary, 20–30 Ma, although some data support an earlier origin. While the timing of the origin of C4 remains controversial, the nearly explosive increase in C4 species is clearly documented in the late Miocene, 4–7 Ma. Increasing seasonality has been widely suggested as an important climatic stimulus for this C4 expansion. Alternatively, based on models of photosynthetic quantum yield at different temperatures and CO2 concentration, it has been hypothesized that the late Miocene C4 expansion resulted from declining atmospheric CO2 levels. This model is most appropriate for explaining the transition from C3 grasslands to C4 grasslands but by itself may not be sufficient to explain the more likely scenario of a late Miocene transition from C3 woodland/ savanna to C4 grasslands. A largely unexplored hypothesis is that climatic changes in late Miocene altered disturbance regimes, in particular the incidence of fires, which today are often associated with maintenance of C4 grasslands. Oceanic charcoal sediments that appear to represent Aeolian deposits from continental wildfires follow a strikingly similar pattern of explosive increase in late Miocene. Climate, CO2, and disturbance are not mutually exclusive explanations and probably all acted in concert to promote the expansion of C4 grasslands. More recently, late Quaternary changes in CO2 may have been responsible for driving major changes in the landscape distribution of C4 species. The theory is sound; however, many of the studies cited in support of this model are open to alternative interpretations, and none has eliminated climatic factors as important selective agents. CAM and C4 evolution required coupling of biochemical pathways with structural changes in photosynthetic tissues, succulence in CAM and Kranz in C4. This was apparently accomplished by piecemeal evolution beginning with mechanisms for recapturing respiratory CO2, although this need not have been so in aquatic CAM species. It has been proposed that the extreme rarity of both pathways in the same plant results from biochemical and structural incompatibilities (Sage 2002). Equally important is the fact that the selective environments are quite different, with CAM evolution thriving on stressful sites inhospitable to C3 species whereas C4 evolution has selected for rapid growth capable of outcompeting associated C3 plants.

International Journal of Plant Sciences↗
USGS2003

Fire and grazing impacts on plant diversity and alien plant invasions in the southern Sierra Nevada

Patterns of native and alien plant diversity in response to disturbance were examined along an elevational gradient in blue oak savanna, chaparral, and coniferous forests. Total species richness, alien species richness, and alien cover declined with elevation, at scales from 1 to 1000 m 2 . We found no support for the hypothesis that community diversity inhibits alien invasion. At the 1-m 2 point scale, where we would expect competitive interactions between the largely herbaceous flora to be most intense, alien species richness as well as alien cover increased with increasing native species richness in all communities. This suggests that aliens are limited not by the number of native competitors, but by resources that affect establishment of both natives and aliens. Blue oak savannas were heavily dominated by alien species and consistently had more alien than native species at the 1-m 2 scale. All of these aliens are annuals, and it is widely thought that they have displaced native bunchgrasses. If true, this means that aliens have greatly increased species richness. Alternatively, there is a rich regional flora of native annual forbs that could have dominated these grasslands prior to displacement by alien grasses. On our sites, livestock grazing increased the number of alien species and alien cover only slightly over that of sites free of livestock grazing for more than a century, indicating some level of permanency to this invasion. In chaparral, both diversity and aliens increased markedly several years after fire. Invasive species are rare in undisturbed shrublands, and alien propagules fail to survive the natural crown fires in these ecosystems. Thus, aliens necessarily must colonize after fire and, as a consequence, time since fire is an important determinant of invasive presence. Blue oak savannas are an important propagule source for alien species because they maintain permanent populations of all alien species encountered in postfire chaparral, and because the vegetation mosaic in this region places them in proximity to chaparral. The speed at which alien propagules reach a burned site and the speed at which the shrublands return to their former closed-canopy condition determine alien invasion. Frequent burning of this vegetation alters the balance in favor of alien invasion. In the higher-elevation coniferous forests, species diversity was a function of fire severity and time since fire. High-intensity fires create gaps that decrease canopy coverage and increase light levels and nutrients for an ephemeral successional flora. Few species have persistent seed banks, so the time since fire is an important determinant of colonization success. There was a highly significant interaction between fire severity and time since fire for understory cover, species richness, and alien richness and cover. Understory was sparse in the first year after fire, particularly in low-severity burns, and increased substantially several years after fire, particularly on high-severity burns. Both fire severity and time since fire affected alien species richness and dominance. Coniferous forests had about one-third as many alien species as the foothill oak savannas, and fewer than half of the species were shared between these communities. Unburned coniferous forests were largely free of alien species, whereas some burned sites had a significant alien presence, which presents a challenge for fire restoration of these forests.

Ecological Applications↗
USGS2003

Historical fire regime in southern California

The historical variability in fire regime is a conservative indicator of ecosystem sustainability. Understanding the natural role of fire in chaparral ecosystems is therefore necessary for effective fire management.

Fire Management Today↗
USGS2002

History and management of crown-fire ecosystems: A summary and response

Some ecosystems, such as yellow pine forests, have had a long history of frequent surface fires, but because of fire suppression policy, fires have been largely excluded from them during the last century (Covington 2000). Unnatural fuel accumulation in these forests has increased the potential for large, catastrophic crown fires, and re-introduction of prescribed fire is one remedy for this critical fire hazard. But fire ecologists and fire managers need to be cautious in transferring this model to all western ecosystems (Anderson et al. 1999; Gutsell et al. 2001). Although large, catastrophic crown fires are apparently unnatural in yellow pine forests (but cf. Shinneman & Baker 1997 ), this is not so in other western forests and shrub-lands, and widespread prescription burning is not warranted everywhere.

Conservation Biology↗
USGS2002

Native American impacts on fire regimes of the California coastal ranges

Aim Native American burning impacts on California shrubland dominated landscapes are evaluated relative to the natural lightning fire potential for affecting landscape patterns. Location Focus was on the coastal ranges of central and southern California. Methods Potential patterns of Indian burning were evaluated based upon historical documents, ethnographic accounts, archaeological records and consideration of contemporary land management tactics. Patterns of vegetation distribution in this region were evaluated relative to environmental factors and the resilience of the dominant shrub vegetation to different fire frequencies. Results Lightning fire frequency in this region is one of the lowest in North America and the density of pre-Columbian populations was one of the highest. Shrublands dominate the landscape throughout most of the region. These woody communities have weak resilience to high fire frequency and are readily displaced by annual grasses and forbs under high fire frequency. Intact shrublands provided limited resources for native Americans and thus there was ample motivation for using fire to degrade this vegetation to an open mosaic of shrubland/grassland, not unlike the agropastoral modification of ecologically related shrublands by Holocene peoples in the Mediterranean Basin. Alien-dominated grasslands currently cover approximately one-quarter of the landscape and less than 1% of these grasslands have a significant native grass presence. Ecological studies in the Californian coastal ranges have failed to uncover any clear soil or climate factors explaining grassland and shrubland distribution patterns. Main conclusions Coastal ranges of California were regions of high Indian density and low frequency of lightning fires. The natural vegetation dominants on this landscape are shrubland vegetation that often form dense impenetrable stands with limited resources for Native Americans. Natural fire frequencies are not high enough to maintain these landscapes in habitable mixtures of shrublands and grasslands but such landscape mosaics are readily produced with additional human subsidy of ignitions. It is hypothesized that a substantial fraction of the landscape was type converted from shrubland to grassland and much of the landscape that underwent such type conversion has either been maintained by Euro-American land management practices or resisted recolonization of native shrublands. It appears that these patterns are disturbance dependent and result from anthropogenic alteration of landscapes initiated by Native Americans and sustained and expanded upon by Euro-American settlers.

Journal of Biogeography↗