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Jon E. Keeley

Publications and source records attributed to Jon E. Keeley.

At least 145 records · Page 8Linked to original sources

Historic fire regime in southern California shrublands

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.

Baja California, California

Mechanism of smoke-induced seed germination in a post-fire chaparral annual

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.

Journal of Ecology

Fire management of California shrubland landscapes

Fire management of California shrublands has been heavily influenced by policies designed for coniferous forests, however, fire suppression has not effectively excluded fire from chaparral and coastal sage scrub landscapes and catastrophic wildfires are not the result of unnatural fuel accumulation. There is no evidence that prescribed burning in these shrublands provides any resource benefit and in some areas may negatively impact shrublands by increasing fire frequency. Therefore, fire hazard reduction is the primary justification for prescription burning, but it is doubtful that rotational burning to create landscape age mosaics is a cost effective method of controlling catastrophic wildfires. There are problems with prescription burning in this crown-fire ecosystem that are not shared by forests with a natural surface-fire regime. Prescription weather conditions preclude burning at rotation intervals sufficient to effect the control of fires ignited under severe weather conditions. Fire management should focus on strategic placement of prescription burns to both insure the most efficient fire hazard reduction and to minimize the amount of landscape exposed to unnaturally high fire frequency. A major contributor to increased fire suppression costs and increased loss of property and lives is the continued urban sprawl into wildlands naturally subjected to high intensity crown fires. Differences in shrubland fire history suggest there may be a need for different fire management tactics between central coastal and southern California. Much less is known about shrubland fire history in the Sierra Nevada foothills and interior North Coast Ranges, and thus it would be prudent to not transfer these ideas too broadly across the range of chaparral until we have a clearer understanding of the extent of regional variation in shrubland fire regimes.

Environmental Management

A new species of Ceanothus from northern Baja California

Ceanothus bolensis S. Boyd & J. Keeley is a new species in the subgenus Cerastes from northwestern Baja California, Mexico. It is well represented at elevations above 1000 m on Cerro Bola, a basaltic peak approximately 35 km south of the U.S./Mexican border. It is characterized by small, obovate to oblanceolate, cupped, essentially glabrous leaves with sparsely toothed margins, pale blue flowers, and globose fruits lacking horns. Principal components analysis on morphological traits shows it to be distinct from other members of Cerastes which are distributed away from the coast in southern California and Baja California, Mexico. These phenetic comparisons also suggest that Ceanothus otayensis should not be subsumed under C. crassifolius , as treated in the Jepson Manual, but rather should be retained at specific rank as well.

Cerro Bola;Baja California

Comparison of post-fire seedling establishment between scrub communities in mediterranean and non-mediterranean climate ecosystems

I Both fire regimes and the conditions under which fires occur vary widely. Abiotic conditions (such as climate) in combination with fire season, frequency and intensity could influence vegetation responses to fire. A variety of adaptations facilitate post-fire recruitment in mediterranean climate ecosystems, but responses of other communities are less well known. We evaluated the importance of climate by comparing sites with mediterranean and subtropical climates. 2 We used paired burned and mature sites in chamise chaparral, mixed chaparral and coastal sage scrub (California), and rosemary scrub, sand pine scrub and sand-hill (Florida), to test whether (i) patterns of pre-fire and post-fire seedling recruitment are more similar between communities within a region than between regions, and (ii) post-fire stimulation of seedling establishment is greater in regions with marked fire-induced contrasts in abiotic site characteristics. 3 Post-fire seedling densities were more similar among sites within climatic regions than between regions. Both seedling densities and proportions of species represented by seedlings after fires were generally higher in California. 4 The only site characteristic showing a pre-fire-post-fire contrast was percentage open canopy, and the effect was greater in California than in Florida. Soil properties were unaffected by fire. 5 Mediterranean climate ecosystems in other regions have nutrient-poor soils similar to our subtropical Florida sites, but show post-fire seedling recruitment patterns more similar to the nutrient-rich sites in California. Climate therefore appears to play a more major role than soil characteristics.

Journal of Ecology

Varied ecosystems need different fire protection

Covington states in his Commentary 1 that the open ponderosa pine forests of the western United States are "in widespread collapse" because fire suppression by humans has eliminated the low-intensity surface fire regime that maintained the open, park-like structure of these forests. He fears this will lead to an "unprecedented" crown fire regime that will eliminate forests.

Nature

On incorporating fire into our thinking about natural ecosystems: A response to Saha and Howe

Ecologists long have had a fascination with fire impacts, although they have been slow to incorporate this ecological factor into serious thinking about the structure of communities and evolution of species (Bond and van Wilgen 1996). The remarks by Saha and Howe (2001, in this issue) illustrate some of the problems ecologists have in trying to apply fire to their thinking about natural ecosystems. Fire is commonly perceived qualitatively in terms of presence or absence, and the variation in frequency and intensity, as well as other components of the fire regime (e.g., fig. 1), is ignored. Often not considered is the fact that plant life histories are fine-tuned to particular fire regims, and in this regard, landscapes present a range of selective peaks and valleys, both figuratively and literally. Because landscape patterns affect the propagation of natural fires, it is imperative that the degree of human disturbance be considered (e.g. Marsh [1864] 1965; Gadgil and Guhu 1993) and the limitations of basing evolutionary arguments on anthropogenically derived landscapes be recognized (e.g., Janzen and Martin 1982).

American Naturalist

We still need Smokey Bear!

It was gratifying to see articles in recent issues of Fire Management Today clarifying the role of Smokey Bear in wildland fire management strategies (Baily 1999; Brown 1999). These articles clearly spelled out Smokey’s importance in reducing unplanned human-ignited wildland fires and rightly criticized attempts to detract from Smokey’s campaign (Williams 1995; see also Vogl 1973).

Fire Management Today

2nd interface between ecology and land development in California

The 2nd Interface Between Ecology and Land Development Conference was held in association with Earth Day 1997, five years after the first Interface Conference. Rapid population growth in California has intensified the inevitable conflict between land development and preservation of natural ecosystems. Sustainable development requires wise use of diminishing natural resources and, where possible, restoration of damaged landscapes. These Earth Week Celebrations brought together resource managers, scientists, politicians, environmental consultants, and concerned citizens in an effort to improve the communication necessary to maintain our natural biodiversity, ecosystem processes and general quality of life. As discussed by our keynote speaker, Michael Soule, the best predictor of habitat loss is population growth and nowhere is this better illustrated than in California. As urban perimeters expand, the interface between wildlands and urban areas increases. Few problems are more vexing than how to manage the fire prone ecosystems indigenous to California at this urban interface. Today resource managers face increasing challenges of dealing with this problem and the lead-off section of the proceedings considers both the theoretical basis for making decisions related to prescribed burning and the practical application. Habitat fragmentation is an inevitable consequence of development patterns with significant impacts on animal and plant populations. Managers must be increasingly resourceful in dealing with problems of fragmentation and the often inevitable consequences, including susceptibility to invasive oganisms. One approach to dealing with fragmentation problems is through careful landplanning. California is the national leader in the integration of conservation and economics. On Earth Day 1991, Governor Pete Wilson presented an environmental agenda that promised to create between land owners and environmentalists, agreements that would guarantee the protection of -endangered species and out of this grew the pioneering initiative, known as the Natural Communities Conservation Planning (NCCP) program. California's vast expanse of seemingly endless resources has traditionally been viewed as justification for abusive land use practices. The modem day recognition that resources are finite has led to greater concern, not only for conserving what is left, but for restoring abused landscapes. Ecological restoration is a new science devoted to returning disturbed environments to a semblance of their 'pristine' state. Based on principles of 'revegetation,' restoration goes far beyond simple replanting, rather the ambition of ecological restoration is to return landscapes to functioning ecosystems and is the focus of the last section.

Open-File Report

Restoration with smoke-dependent species

Examination of blood films from wood ducks (Aix sponsa) from several northeastern states revealed Haemoproteus, Leucocytozoon, Plasmodium and a typanosome. Haemoproteus occurred in all areas sampled and birds of the year from Massachusetts demonstrated the highest incidence during the last 2 weeks in August. Leucocytozoon was most prevalent in more northern areas. P. circumflexum and a trypanosome are reported for the first time from this host.

Ecological Restoration

Patch to landscape patterns in post fire recruitment of a serotinous conifer

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.

California

C4 photosynthetic modifications in the evolutionary transition from land to water in aquatic grasses

Cladistic analysis supports the conclusion that the Orcuttieae tribe of C 4 grasses reflect evolution from a terrestrial ancestry into seasonal pools. All nine species in the tribe exhibit adaptations to the aquatic environment, evident in the structural characteristics of the juvenile foliage, which persist submerged for 1–3 months prior to metamorphosis to the terrestrial foliage. Aquatic leaves of the least derived or basal genus Neostapfia have few morphological and anatomical characteristics specialized to the aquatic environment and have retained full expression of the C 4 pathway, including Kranz anatomy. Orcuttia species have many derived characteristics and are more specialized to the aquatic environment. These latter species germinate earlier in the season and persist in the submerged stage longer than Neostapfia and evidence from the literature indicates length of submergence is positively correlated with fitness components. Aquatic leaves of Orcuttia species lack Kranz or PCR bundle sheath anatomy, yet 14 C-pulse chase studies indicate >95% malate + aspartate as the initial products of photosynthesis and these products turn over rapidly to phosphorylated sugars, indicating a tight coupling of the C 4 and C 3 cycles. Presence of the C 4 pathway is further supported by enzymological data. Contemporary dogma that Kranz anatomy is a sine qua non for operation of the C 4 pathway is contradicted by the patterns in Orcuttia ; however, it is unknown whether the pathway acts as a CO 2 concentrating mechanism in these aquatic plants.

California

Diel acid fluctuations in C4 amphibious grasses

Orcuttieae is a small tribe of C4 grasses endemic to seasonal pools in the southwestern U.S., comprising the basal genus Neostapfia, Tuctoria, and the most derived group, Orcuttia. Growth is initiated underwater, and when pools dry, species undergo a metamorphosis replacing aquatic foliage with terrestrial foliage. O. californica and O. viscida exhibit CAM-like diel fluctuations in acidity in the aquatic foliage. Pulse-chase studies showed that although CO2 was fixed into malic acid in the dark, an overnight chase in the dark revealed that most label was not retained in organic acids, indicating a role other than CAM. Terrestrial foliage exhibited a very different diel fluctuation; acids accumulated during the day, and diminished overnight. Malic acid predominated and was secreted on the surface of the leaf in a manner similar to another arid land species. This terrestrial daytime acid accumulation may not be related to photosynthetic pathway but may play an anti-herbivore function. No acid fluctuations were observed in either N. colusana or T. greenei.

Photosynthetica

Smoke-induced seed germination in California chaparral

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.

California