Geology topics
Publications and source records attributed to C. J. Fotheringham.
The effects of fire on seed germination and plant regeneration are discussed. Among the topics considered are the triggering of opening of serotinous fruits or cones by fire, the breaking of dormancy in seeds in the soil seed bank, the effects of smoke on germination, and the role of fire in initiating seedling recruitment by opening gaps in closed vegetation.
Although many residents of southern California have long recognised that wildfires in the region are an ongoing, constant risk to lives and property, the enormity of the regional fire hazard caught the world’s attention during the southern California firestorms of 2003 (Figure 5.1). Beginning on 21 October, a series of fourteen wildfires broke out across the five-county region under severe Santa Ana winds, and within two weeks, more than 300,000 ha had burned (Keeley et al., 2004). The event was one of the costliest in the state’s history, with more than 3,600 homes damaged or destroyed and twenty-four fatalities. Suppression costs for the 12,000 firefighters have been estimated at US$120 million, and the total response and damage cost has been estimated at more than US$3 billion (COES, 2004). [Excerpt]
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.
In the paper 'NO News', Preston et al. (2004) make a number of erroneous assumptions regarding nitrogen oxide chemistry. These authors also present some very significant misinterpretations of previous research into the effects of various nitrogen oxides on germination of post-fire followers. Methodological differences between the study by Preston et al. (2004) and previous work are also problematic, such as using NO-donors in solution versus the use of direct application of various nitrogen oxides in the gaseous phase. A closer review of these studies, with the proper understanding of nitrogen oxide chemistry, and interpretations of the available literature, would lead to the conclusion that, contrary to the authors' assertions, the Preston et al. (2004) study supports, rather than refutes, earlier findings by Keeley and Fotheringham (1997, 1998a, b, 2000). ?? CAB International 2005.
Wildfires elicit a perceived need for emergency action to stabilize denuded landscapes. Aerial seeding of rapidly growing nonnative grasses is used routinely in an attempt to control postfire erosion, despite limited scientific basis for its effectiveness and with little consideration for its unintended ecological impacts. As fire size and magnitude have increased in recent decades, so has the prevalence and cost of postfire seeding and the potential footprint of its unintended impacts. We see a growing consensus in the research community on two important points: this management practice often is not cost-effective and it appears to create more problems than it solves.
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No abstract available at this time
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.
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.
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.
Historical variability in fire regime is a conservative indicator of ecosystem sustainability, and thus understanding the natural role of fire in chaparral ecosystems is necessary for proper fire management. It has been suggested that the “natural” fire regime was one of frequent small fires that fragmented the landscape into a fine-grained mixture of age classes that precluded large, catastrophic fires. Some researchers claim that this regime was lost because of highly effective fire suppression and conclude that if fire managers could “restore” a regime of frequent fires with widespread prescription burning, they could eliminate the hazard of catastrophic fires. The primary evidence in support of this model is a study that compared contemporary burning patterns in southern California, U.S.A., a region subject to fire suppression, with patterns in northern Baja California, Mexico, where there is less effective fire suppression. We found that differences in fire regime between these two regions are inconclusive and could not be ascribed conclusively to differences in fire suppression. Historical records suggest that the natural fire regime in southern California shrublands was rather coarse-grained and not substantively different from the contemporary regime. There is no evidence that fire-management policies have created the contemporary fire regime dominated by massive Santa Ana wind-driven fires. Increased expenditures on fire suppression and increased loss of property and lives are the result of human demographic patterns that place increasing demand on fire-suppression forces.
1 Smoke-stimulated germination in the post-fire flora of California chaparral does not appear to be triggered by nitrate. Application of freshly prepared unbuffered KNO 3 solutions (pH c. 6.2) failed to enhance germination of five populations of Emmenanthe penduliflora or one Phacelia grandiflora population, regardless of light or stratification conditions. 2 KNO 3 buffered at acidic pH (or unbuffered solutions equilibrated with atmospheric CO 2 ) did induce germination, but KNO 3 solutions at pH 7 failed to induce germination. Induction of germination is therefore not due to the nitrate ion per se , but rather to high [H + ], although buffered controls gave weak germination at low pH, suggesting a role for H + plus nitrate. However, other anions such as sulphate were equally as effective as nitrate at breaking dormancy. 3 The germination response to KNO 3 was affected by the type of filter paper used and this may be linked to differences in pH. 4 NO 2 , at concentrations present in biomass smoke, was highly effective at inducing germination, and other oxidizing agents also induced germination. 5 Several growth regulators, including nitrite and gibberellin, were stimulatory only at acidic pH, but KCN was stimulatory across a broad pH range. 6 Germination decreased at smoke exposures longer than a few minutes. Also, smoked water samples effective at breaking dormancy were acidic and were less effective when buffered to pH >7. 7 Physical scarification of the seed coat induced germination but the effect was not due to penetration of a water barrier, or to enhanced oxygen uptake or to wound responses such as CO 2 or ethylene production. 8 Different effects of the gibberellin inhibitor CCC (chlorocholine chloride) suggested that the mechanisms of scarification-induced and smoke-induced germination may differ. 9 We conclude that either oxidizing gases in smoke and/or acids generated on burnt sites play a role in germination of post-fire annuals in chaparral.
Obligate seeding species are highly specialized to fire disturbance and many conifers such as cypress, which are adapted to high intensity stand-replacing fires, have canopy seed banks stored in serotinous cones. Resilience of these trees to fire disturbance is a function of disturbance frequency and one focus of this study was to determine the effect of patch age on postfire recruitment. A second focus was to determine the extent to which fire induced a landscape level change in the location of the forest boundary. Prior to a fire in 1994, a large Cupressus sargentii forest was a mosaic landscape of different aged patches of nearly pure cypress bordered by chaparral. Patches less than 60 years of age were relatively dense with roughly one tree every 1–2 m 2 but older patches had thinned to one tree every 3–15 m 2 . Older trees had substantially greater canopy cone crops but the stand level seed bank size was not significantly correlated with stand age. Fire-dependent obligate seeding species are sensitive to fire return interval because of potential changes in the size of seed banks – facing both a potential `immaturity risk' and a `senescence risk'. At our site, C. sargentii regeneration was substantial in stands as young as 20 years, suggesting that fire return interval would need to be shorter than this to pose any significant risk. Reduced seedling recruitment in stands nearly 100 years of age may indicate risk from senescence is greater, however, even the lowest density seedling recruitment was many times greater than the density of mature forests – thus this cypress would appear to be resilient to a wide range of fire return intervals. Changes in landscape patterning of forest and chaparral are unlikely except after fire. Factors that inhibit tree establishment within the shrubland, as well as factors that affect shrub establishment within the forest border likely affect the `permeability' of this ecotone. After the 1994 fire this boundary appeared to be stable in that cypress recruited best within the shadow of burned canopies and cypress were weak invaders of adjacent shrublands.
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The California chaparral community has a rich flora of species with different mechanisms for cuing germination to postfire conditions. Heat shock triggers germination of certain species but has no stimulatory effect on a great many other postfire species that are chemically stimulated by combustion products. Previous reports have shown that charred wood will induce germination, and here we report that smoke also induces germination in these same species. Smoke is highly effective, often inducing 100% germination in deeply dormant seed populations with 0% control germination. Smoke induces germination both directly and indirectly by aqueous or gaseous transfer from soil to seeds. Neither nitrate nor ammonium ions were effective in stimulating germination of smoke-stimulated species, nor were most of the quantitatively important gases generated by biomass smoke. Nitrogen dioxide, however, was very effective at inducing germination in Caulanthus heterophyllus (Brassicaceae), Emmenanthe penduliflora (Hydrophyllaceae), Phacelia grandiflora (Hydrophyllaceae), and Silene multinervia (Caryophyllaceae). Three species, Dendromecon rigida (Papaveraceae), Dicentra chrysantha, and Trichostema lanatum (Lamiaceae), failed to germinate unless smoke treatment was coupled with prior treatment of 1 yr soil storage. Smoke-stimulated germination was found in 25 chaparral species, representing 11 families, none of which were families known for heat-shock-stimulated germination. Seeds of smoke-stimulated species have many analogous characteristics that separate them from most heat-shock-stimulated seeds, including: (1) outer seed coats that are highly textured, (2) a poorly developed outer cuticle, (3) absence of a dense palisade tissue in the seed coat, and (4) a subdermal membrane that is semipermeable, allowing water passage but blocking entry of large (molecular mass > 500) solutes. Tentative evidence suggests that permeability characteristics of this subdermal layer are altered by smoke. While the mechanism behind smoke-induced germination is not known, it appears that smoke may be involved in overcoming different blocks to germination in different species. For example, in Emmenanthe penduliflora, NO 2 in smoke was sufficient to induce germination, and most forms of physical or chemical scarification also induced germination. For Romneya coulteri, NO 2 alone failed to induce germination, and scarified seeds required addition of gibberellic acid. In Dicentra chrysantha, none of these treatments, nor smoke alone, induced germination, but germination was triggered by a combination of soil burial followed by smoke treatment. Smoke-stimulated species differed substantially in the duration of smoke exposure required to induce germination, and this was inversely correlated with tolerance to smoke exposure. We suggest that such differences in response may affect postfire community structure.
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No abstract available at this time
Dormant seeds of a California chaparral annual were induced to germinate by smoke or vapors emitted from smoke-treated sand or paper. Nitrogen oxides induced 100 percent germination in a manner similar to smoke. Smoke-treated water samples inducing germination were comparable in acidity and concentration of nitrate and nitrite to nitrogen dioxide (NO 2 )–treated samples. Vapors from smoke-treated and NO 2 -treated filter paper had comparable NO 2 flux rates. Chaparral wildfires generate sufficient nitrogen oxides from combustion of organic matter or from postfire biogenic nitrification to trigger germination of Emmenanthe penduliflora . Nitrogen oxide–triggered germination is not the result of changes in imbibition, as is the case with heat-stimulated seeds.