Fire ecology

Fire ecology
The Old Fire burning in the San Bernardino Mountains (image taken from the International Space Station)

Fire ecology is concerned with the processes linking the natural incidence of fire in an ecosystem and the ecological effects of this fire. Many ecosystems, such as the North American prairie and chaparral ecosystems, and the South African savanna, have evolved with fire as a natural and necessary contributor to habitat vitality and renewal. [1][2][3] Many plant species in naturally fire-affected environments require fire to germinate. Fire suppression can lead to the build-up of inflammable debris and the creation of less frequent but much larger and destructive wildfires.

Campaigns in the United States have historically molded public opinion to believe that wildfires are always harmful to nature. This view is based on the outdated belief that ecosystems progress toward an equilibrium and that any disturbance, such as fire, disrupts the harmony of nature. More recent ecological research has shown, however, that fire is an integral component in the function and biodiversity of many natural habitats, and that the organisms within these communities have adapted to withstand, and even to exploit, natural wildfire.

Fire suppression, in combination with other human-caused environmental changes, has resulted in unforeseen consequences for natural ecosystems. Some uncharacteristically large wildfires in the United States have been caused as a consequence of years of fire suppression and the continuing expansion of people into fire-adapted ecosystems. Land managers are faced with tough questions regarding where to restore a natural fire regime.

Color-coded map of the different types of wildfire by location in the United States
Map of the different regimes of natural burning with location in the United States. Colors denote both the frequency of wildfires and their style of burning. Before European colonization, wildfires occurred most frequently in the tan, yellow, blue, pink, and light green areas.


Fire Components

A fire regime describes the pattern that fire follows in a particular ecosystem. Its "severity" is a term that ecologists use to refer to the impact that a fire has on an ecosystem. Ecologists can define this in many ways, but one way is through an estimate of plant mortality. Fire can burn at three levels. Ground fires will burn through soil that is rich in organic matter. Surface fires will burn through dead plant material that is lying on the ground. Crown fires will burn in the tops of shrubs and trees. Ecosystems may experience predominantly one of these fire regimes, or a mix of all three.[4]

Fires will often break out during a dry season, but in some areas wildfires may also commonly occur during a time of year when lightning is prevalent. The frequency over a span of years at which fire will occur at a particular location is a measure of how common wildfires are in a given ecosystem. It is either defined as the average interval between fires at a given site, or the average interval between fires in an equivalent specified area.[4]

Defined as the energy released per unit length of fireline (kW m-1), wildfire intensity can be estimated either as the product of the linear spread rate (m s-1), the low heat of combustion (kJ kg-1) and the combusted fuel mass per unit area, or it can be estimated from the flame length.[5]

Radiata Pine forest burnt during the 2003 Bogong Bushfires, Australia

Abiotic responses

Fire has important effects on the abiotic (non-living) components of an ecosystem, particularly the soil. Fire can affect the soil by direct contact with it and by its effects on the plant community associated with it.[6] By removing overhead vegetation, fire can lead to increased solar radiation on the soil surface by day, resulting in greater warming, and to greater cooling through the loss of radiative heat at night. Fewer leaves left to intercept rain will allow more moisture to reach the soil surface. In addition, plant transpiration (the process by which water travels through plants and evaporates through pores in the leaves) will be reduced following a fire, allowing the soil to retain more moisture. Exposure to sunlight, wind and evaporation, however, will work in the other way, to dry the soil. The fire may have created an impermeable crust at the soil surface, if organic matter on the ground was heated by the fire into a waxy residue, and if this has happened, it may lead to increased soil erosion through surface run-off.

Fire may cause nutrient loss through a variety of mechanisms, including oxidation, volatilization, and increased erosion and leaching by water. Temperatures must be very high, however, to cause a significant loss of nutrients, which are often replaced by organic matter left behind in the fire. Charcoal is able to counteract some nutrient and water loss because of its absorptive properties.

Overall, soils become more basic (higher pH) following fires because of acid combustion. By driving novel chemical reactions at high temperatures, fire can even alter the texture and structure of soils by affecting the clay content and the soil's porosity.

Biotic responses and adaptations


Lodgepole pine cones

Plants have evolved many adaptations to cope with fire. In chaparral communities in Southern California, for example, some plants have leaves coated in flammable oils that encourage an intense fire. This heat causes their fire-activated seeds to germinate and the young plants can then capitalize on the lack of competition in a burnt landscape. Other plants have smoke-activated seeds, or fire-activated buds. The serotinous cones of the Lodgepole pine (Pinus contorta) are sealed with a resin that a fire melts away, releasing the seeds.[7] Many plant species, including the shade-intolerant giant sequoia (Sequoiadendron giganteum), require fire to make gaps in the vegetation canopy that will let in light, allowing their seedlings to compete with the more shade-tolerant seedlings of other species, and so establish themselves.[8] Because their stationary nature precludes any fire avoidance, plant species may only be fire-intolerant, fire-tolerant or fire-resistant.[9]

Fire intolerance

Fire-intolerant plant species tend to be highly flammable and are destroyed completely by fire. Some of these plants and their seeds may simply fade from the community after a fire and not return, others have adapted to ensure that their offspring survives into the next generation. “Obligate seeders” are plants with large, fire-activated seed banks that germinate, grow, and mature rapidly following a fire, in order to reproduce and renew the seed bank before the next fire.[9][10]

Fire tolerance. Typical regrowth after an Australian bushfire

Fire tolerance

Fire-tolerant species are able to withstand a degree of burning and continue growing despite damage from fire. These plants are sometimes referred to as “resprouters.” Ecologists have shown that some species of resprouters store extra energy in their roots to aid recovery and re-growth following a fire.[9][10] For example, after an Australian bushfire, the Mountain Grey Gum tree (Eucalyptus cypellocarpa) starts producing a mass of shoots of leaves from the base of the tree all the way up the trunk towards the top, making it look like a black stick completely covered with young, green leaves.

Fire resistance

Fire-resistant plants suffer little damage during a characteristic fire regime. These include large trees whose flammable parts are high above surface fires. Mature Ponderosa Pine (Pinus ponderosa) is an example of a tree species that suffers virtually no crown damage under a naturally mild fire regime, because it sheds its lower, vulnerable branches as it matures.[9][11]

Animals, birds and microbes

Wildfire in the Bitterroot National Forest in Montana, United States, on August 6, 2000

Like plants, animals display a range of abilities to cope with fire, but they differ from plants in that they must avoid the actual fire to survive. Although birds are vulnerable when nesting, they are generally able to escape a fire; indeed they often profit from being able to take prey fleeing from a fire and to recolonize burned areas quickly afterwards. Mammals are often capable of fleeing a fire, or seeking cover if they can burrow. Amphibians and reptiles may avoid flames by burrowing into the ground or using the burrows of other animals. Amphibians in particular are able to take refuge in water or very wet mud.[9] Some arthropods also take shelter during a fire, although the heat and smoke may actually attract some of them, to their peril.[12] Microbial organisms in the soil vary in their heat tolerance but are more likely to be able to survive a fire the deeper they are in the soil. A low fire intensity, a quick passing of the flames and a dry soil will also help. An increase in available nutrients after the fire has passed may result in larger microbial communities than before the fire.[6]

Long term impacts

Ecological succession

Fire behavior is different in every ecosystem and the organisms in those ecosystems have adapted accordingly. One sweeping generality is that in all ecosystems, fire creates a mosaic of different habitat patches, with areas ranging from those having just been burned to those that have been untouched by fire for many years. This is a form of ecological succession in which a freshly burned site will progress through continuous and directional phases of colonization following the destruction caused by the fire.[13] Ecologists usually characterize succession through the changes in vegetation that successively arise. After a fire, the first species to re-colonize will be those whose seeds are already present in the ground, or those whose seeds are able to travel into the burned area quickly. These are generally fast-growing herbaceous plants that need lots of light and are poor competitors in crowded areas. As time passes, more slowly growing, shade-tolerant, and competitive, woody species will crowd out the herbaceous plants. These woody plants may be shrubs or trees.[14]

Different species of plants, animals, and microbes specialize in exploiting different stages in this process of succession, and by creating these different types of patches, fire allows a greater number of species to exist within a landscape. Soil characteristics will be a factor in determining the specific nature of a fire-adapted ecosystem, as will climate and topography.


Mild to moderate fires burn in the forest understory, removing small trees and herbaceous groundcover. Only high-intensity fires will burn into the crowns of the tallest trees. Crown fires may require support from ground fuels to maintain the fire in the forest canopy (passive crown fires), or the fire may burn in the canopy independently of any ground fuel support (an active crown fire). Fires used in the management of woodlands will typically aim for low to moderate intensity, whereas wildfires can evolve into crown fires. When a forest burns frequently and thus has less plant litter build-up, below-ground soil temperatures rise only slightly and will not be lethal to roots that lie deep in the soil.[12] Although other characteristics of a forest will influence the impact of fire upon it, factors such as climate and topography play an important role in determining fire severity and fire extent.[15] Fires spread most widely during drought years, are most severe on upper slopes and are influenced by the type of vegetation that is growing.

Forests in British Columbia

In Canada, forests cover about 10% of the land area and yet harbor 70% of the country’s bird and terrestrial mammal species. Natural fire regimes are important in maintaining a diverse assemblage of vertebrate species in up to twelve different forest types in British Columbia.[16] Different species have adapted to exploit the different stages of succession, regrowth and habitat change that occurs following an episode of burning, such as downed trees and debris. The characteristics of the initial fire, such as its size and intensity, cause the habitat to evolve differentially afterwards and influence how vertebrate species are able to use the burned areas.[16]



Shrub fires typically concentrate in the canopy and spread continuously if the shrubs are close enough together. Shrublands are typically dry and are prone to accumulations of highly volatile fuels, especially on hillsides. Fires will follow the path of least moisture and the greatest amount of dead fuel material. Surface and below-ground soil temperatures during a burn are generally higher than those of forest fires because the centers of combustion lie closer to the ground, although this can vary greatly.[12] Common plants in shrubland or chaparral include manzanita, chamise and Coyote Brush.

California shrublands

California shrubland, commonly known as chaparral, is a widespread plant community of low growing species, typically on arid sloping areas of the California Coast Ranges or western foothills of the Sierra Nevada. There are a number of common shrubs and tree shrub forms in this association, including salal, toyon, coffeeberry and Western poison oak.[17] Regeneration following a fire is usually a major factor in the association of these species.

South African Fynbos shrublands

Fynbos shrublands occur in a small belt across South Africa. The plant species in this ecosystem are highly diverse, yet the majority of these species are obligate seeders, that is, a fire will cause germination of the seeds and the plants will begin a new life-cycle because of it. These plants may have evolved into obligate seeders as a response to fire and nutrient-poor soils.[18] Because fire is common in this ecosystem and the soil has limited nutrients, it is most efficient for plants to produce many seeds and then die in the next fire. Investing a lot of energy in roots to survive the next fire when those roots will be able to extract little extra benefit from the nutrient-poor soil would be less efficient. It is possible that the rapid generation time that these obligate seeders display has led to more rapid evolution and speciation in this ecosystem, resulting in its highly diverse plant community.[18]


Grasslands burn more readily than forest and shrub ecosystems, with the fire moving through the stems and leaves of herbaceous plants and only lightly heating the underlying soil, even in cases of high intensity. In most grassland ecosystems, fire is the primary mode of decomposition, making it crucial in the recycling of nutrients.[12]

South African savanna

In the savanna of South Africa, recently burned areas have new growth that provides palatable and nutritious forage compared to older, tougher grasses. This new forage attracts large herbivores from areas of unburned and grazed grassland that has been kept short by constant grazing. On these unburned "lawns", only those plant species adapted to heavy grazing are able to persist; but the distraction provided by the newly burned areas allows grazing-intolerant grasses to grow back into the lawns that have been temporarily abandoned, so allowing these species to persist within that ecosystem.[3]

Fire suppression

Fire serves many important functions within fire-adapted ecosystems. Fire plays an important role in nutrient cycling, diversity maintenance and habitat structure. The suppression of fire can lead to unforeseen changes in ecosystems that often adversely affect the plants, animals and humans that depend upon that habitat. Wildfires that deviate from a historical fire regime because of fire suppression are called “uncharacteristic fires.”

Chaparral communities

In 2003, southern California witnessed powerful chaparral wildfires. Hundreds of homes and hundreds of thousands of acres of land went up in flames. Extreme fire weather (low humidity, low fuel moisture and high winds) and the accumulation of dead plant material from 8 years of drought, contributed to a catastrophic outcome. Although some have maintained that fire suppression contributed to an unnatural buildup of fuel loads,[19] a detailed analysis of historical fire data has showed that this may not have been the case.[20] Fire suppression activities had failed to exclude fire from the southern California chaparral. Research showing differences in fire size and frequency between southern California and Baja has been used to imply that the larger fires north of the border are the result of fire suppression, but this opinion has been challenged by numerous investigators and is no longer supported by the majority of fire ecologists.

One consequence of the fires in 2003 has been the increased density of invasive and non-native plant species that have quickly colonized burned areas, especially those that had already been burned in the previous 15 years. Because shrubs in these communities are adapted to a particular historical fire regime, altered fire regimes may change the selective pressures on plants and favor invasive and non-native species that are better able to exploit the novel post-fire conditions.[21]

Fish impacts

The Boise National Forest is a US national forest located north and east of the city of Boise, Idaho. Following several uncharacteristically large wildfires, an immediately negative impact on fish populations was observed, posing particular danger to small and isolated fish populations.[22] In the long term, however, fire appears to rejuvenate fish habitats by causing hydraulic changes that increase flooding and lead to silt removal and the deposition of a favorable habitat substrate. This leads to larger post-fire populations of the fish that are able to recolonize these improved areas.[22] But although fire generally appears favorable for fish populations in these ecosystems, the more intense effects of uncharacteristic wildfires, in combination with the fragmentation of populations by human barriers to dispersal such as weirs and dams, will pose a threat to fish populations.

Ponderosa pine forests

Prior to European settlement, a typical Ponderosa pine forest in the southwest United States was mostly savanna, a grassland ecosystem with scattered stands of trees, resulting in less than 25–30% of the ground shaded by an interlocking crown cover of Ponderosa Pine trees. Fires occurred at intervals of perhaps less than ten years or so in any one place, with average burns covering 3,000 acres (12 km2).[23] These forests historically suffered mild to moderate fires that generally did not reach the crown and left most of the trees alive.[24]

Ponderosa pine forests now face severe damage under harsher fire regimes brought on by fire suppression and aggravated by natural drought cycles.[25] Fires in these forests now result in crown fires that cause extensive tree-mortality. Fire suppression may also lead to increased defoliation of the trees by herbivorous insects, whose populations might otherwise be controlled by more regular outbreaks of wildfire.[24]

An important factor in the increased intensity and destructive power of wildfire in the Ponderosa pine forests is the impact of cattle grazing on public land. Prior to the arrival of railroads in the American West in the 1880s and 1890s, cattle and sheep grazing had a relatively low impact in this sort of terrain because livestock could not be transported to markets on the east and west coasts. Before the railroads, the average Ponderosa Pine stand density was 40–60 trees per acre (100–150 per hectare), based on evidence from the analysis of old photographs. Historically, throughout the west of the United States, Ponderosa pine forests experienced low-temperature, rapidly moving surface fires every ten years or so, which burned the grasses and herbaceous vegetation on the savanna floor, along with seedling pines, maintaining a low stand density and an open canopy. The introduction of cattle grazing, however, significantly reduced the fuel load on the ground, which effectively halted this natural wildfire regime. Over the next century, photographic evidence reveals that the stand density of the Ponderosa Pine increased from 40–60 to 600–800 stems per acre (from 100–150 to 1500–2000 per hectare). These trees are crowded together with a closed canopy, competing for both water and light. The lower branches are dead and bare.

As a result of this, the present-day fire regime is predominantly characterized by intense, devastating crown fires which volatilize nutrients, create tremendous erosion problems and pose an increased risk to firefighters and communities alike. US Forest Service Ranger Bill Armstrong of Los Alamos, New Mexico, prior to the 2000 Cerro Grande Fire, thinned an area of Pajarito Plateau to 40 stems per acre. The Cerro Grande Fire burned all the forest around the thinned area, but only burned ground cover within the experimental area.[attribution needed] Similar effects of thinning back tree density to that seen before the coming of the railroad show similar results.[attribution needed]

Fire as a management tool

Restoration ecology is the name given to an attempt to reverse or mitigate some of the changes that humans have caused to an ecosystem. Controlled burning is one tool that is currently receiving considerable attention as a means of restoration and management. Applying fire to an ecosystem may create habitats for species that have been negatively impacted by fire suppression, or fire may be used as a way of controlling invasive species without resorting to herbicides or pesticides. But what should managers aim to restore their ecosystems to? Does “natural” mean pre-human? Pre-European? Native American use of fire, not natural fires, historically maintained the diversity of the savannas of North America.[26][27] When, how, and where managers should use fire as a management tool is a subject of debate.

The Florida everglades

Everglades landscape

The Florida Everglades is one example of an ecosystem with a historical regime of frequent fires. Currently, the everglades are undergoing long-term and large-scale restoration.[28][citation needed] A problem that ecologists and managers have is how frequently to prescribe burns. There is a strong relationship between climate and fire in Florida and it may be that climate holds the key to this question.[28] The El Niño Southern Oscillation increases the frequency of lighting strikes, opening up a window for fire before there is too much precipitation. It is possible, however, that human-induced climate change may result in a perpetual El Niño that never allows conditions to become dry enough for fire in the future.

The Great Plains shortgrass prairie

A combination of heavy livestock grazing and fire-suppression has drastically altered the structure, composition, and diversity of the shortgrass prairie ecosystem on the Great Plains, allowing woody species to dominate many areas and promoting fire-intolerant invasive species. In semi-arid ecosystems where the decomposition of woody material is slow, fire is crucial for returning nutrients to the soil and allowing the grasslands to maintain their high productivity.

Although fire can occur during the growing or the dormant seasons, managed fire during the dormant season is most effective at increasing the grass and forb cover, biodiversity and plant nutrient uptake in shortgrass prairie.[2] Managers must also take into account, however, how invasive and non-native species respond to fire if they want to restore the integrity of a native ecosystem. For example, fire can only control the invasive spotted knapweed (Centaurea maculosa) on the Michigan prairie grasslands in the summer, because this is the time in the knapweed’s life cycle that is most important to its reproductive growth.[29]

Mixed conifer forests in the Sierra Nevada

Mixed conifer forests in the United States Sierra Nevada used to have fire return intervals that ranged from 5 years up to 300 years, depending on the local climate.[30] Lower elevations had more frequent fire return intervals, whilst higher and wetter elevations saw much longer intervals between fires. Native Americans tended to set fires during fall and winter, and land at a higher elevation was generally occupied by Native Americans only during the summer.[31]

In areas that formerly had frequent fire return intervals, fire suppression has modified the previous fire regime, resulting in the heavy accumulation of forest fuels such as downed coarse woody debris and ladder fuels composed of shade tolerant species such as white fir (Abies concolor).[32] Forests whose burning is to be managed, usually have their ground fuels and forest structure modified beforehand, so that wildland fire burning can be more as it used to be. Treatments are done either by hand crews, with mechanical equipment or through a combination of the two. Hand thinning has the advantage of being very "light on the land" but has the disadvantages of high cost and can be ineffective when the ladder fuels are above 14 inches (360 mm) in diameter measured at breast height. While mechanical treatment methods can remove valuable small diameter logs and biomass, they cannot easily treat steep slopes and they can compact soils if great care is not taken. Both hand thinning and mechanical treatments can leave in place the fuel that is important for creating wildlife habitats, such as large fallen trees and snags.

The material removed from the woodland by both hand thinning and mechanical treatments must be properly disposed of before managed or natural wildland fires can be reintroduced into the stand.[33] In the case of hand thinning, piles are usually created and burned. Mechanical treatments can remove the slash and then burn or crush it. Regardless of the method used, once the forest has been treated, fire can be reintroduced.[34]

Management policies

United States

Fire policy in the United States involves the federal government, individual state governments, tribal governments, interest groups, and the general public. The new federal outlook on fire policy parallels advances in ecology and is moving towards the view that many ecosystems depend on disturbance for their diversity and for the proper maintenance of their natural processes. Although human safety is still the number one priority in fire management, new US government objectives include a long-term view of ecosystems. The newest policy allows managers to gauge the relative values of private property and resources in particular situations and to set their priorities accordingly.[7]

Techniques such as sophisticated risk assessment strategies, that integrate the latest in ecological research with the social and economic consequences of a particular outcome, are one way to make the most informed fire policy decisions based on the interests of many stakeholders.[35] The US government now recognizes that the longer fuel accumulates in fire-suppressed areas, the greater will be the damage when an unexpected fire burns out of control.[7] One of the primary goals in fire management is to improve public education in order to suppress the “Smokey Bear” fire-suppression mentality and introduce the public to the benefits of regular natural fires.

Fire reintroduction will need to be mindful of regulations set by the Clean Air Act and the Environmental Protection Agency concerning wildfire emissions, limited fire professionals, potential property damage from escaped fire and complaints about smoke and destruction of scenic views.[7]

See also


  1. ^ California Chaparral Institute: Wildfire in the Chaparral . accessed 9/29/2010
  2. ^ a b Brockway et al. (2002)
  3. ^ a b Archibald et al. 2005
  4. ^ a b Bond and Keeley 2005
  5. ^ Byram, 1959
  6. ^ a b Hart et al. 2005
  7. ^ a b c d USDA Forest Service
  8. ^ US National Park Service
  9. ^ a b c d e Kramp et al. 1986
  10. ^ a b Knox and Clarke 2005
  11. ^ Pyne 2002
  12. ^ a b c d DeBano et al. 1998
  13. ^ Begon et al. 1996, pg. 692
  14. ^ Begon et al. 1996, pg 700
  15. ^ Beaty and Taylor (2001)
  16. ^ a b Bunnell (1995)
  17. ^ C.Michael Hogan (2008) "Western poison-oak: Toxicodendron diversilobum", GlobalTwitcher, ed. Nicklas Strömberg
  18. ^ a b Wisheu et al. (2000)
  19. ^ Minnich 1983
  20. ^ Keeley et al. 1999
  21. ^ Keeley et al. 2005
  22. ^ a b Burton (2005)
  23. ^ Covington, W. Wallace; Margaret M. Moore (1994-01). "Southwestern ponderosa forest structure: Changes since Euro-American Settlement". Journal of Forestry (Society of American Foresters) 92 (1): 39–47. 
  24. ^ a b McCullough et al. (1998)
  25. ^ Savage and Mast 2005
  26. ^ MacDougall et al. (2004)
  27. ^ Williams, Gerald W. (2003-06-12). "REFERENCES ON THE AMERICAN INDIAN USE OF FIRE IN ECOSYSTEMS". Retrieved 2008-07-31. 
  28. ^ a b Beckage et al. (2005)
  29. ^ Emery and Gross (2005)
  30. ^ Fire Regimes in Sierrian Mixed-Conifer Forests
  31. ^ ANDERSON, M. KAT; MICHAEL J. MORATTO (1996). "9: Native American Land-Use Practices and Ecological Impacts". Sierra Nevada Ecosystem Project: Final report to Congress, vol. II, Assessments and scientific basis for management options. Davis: University of California, Centers for Water and Wildland Resources. pp. 191, 197, 199.,accessed=2008-08-09. 
  32. ^ An Overview of Fire in the Sierra Nevada, KEVIN S. McKELVEY, Carl N. Skinner, Chi-ru Chang, Don C. Et-man, Susan J. Husari, David J. Parsons, Jan W. van Wagtendonk, C. Phillip Weatherspoon, In Sierra Nevada Ecosystem Project: Final report to Congress, vol. II, Assessments and scientific basis for management options: Davis: University of California, Centers for Water and Wildland Resources, 1996.
  33. ^ Performance Based Hazard Reduction, 2006, Proposed Rule Language for 45 Day Notice, Published August 26, 2005
  34. ^ Fire-Silviculture Relationships in Sierra Forests, C. PHILLIP WEATHERSPOON, Pacific Southwest Research Station, U.S. Forest Service, Redding, California
  35. ^ Dellasala et al. 2004, Fairbrother and Turnley 2005


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  • Pyne, S.J. “How Plants Use Fire (And Are Used By It).” 2002. PBS NOVA Online. 1 January 2006.
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  • Stephens, S. L., and J. J. Moghaddas. 2005. Fuel treatment effects on snags and coarse woody debris in a Sierra Nevada mixed conifer forest. Forest Ecology and Management 214:53-64.
  • United States Department of Fish and Agriculture (USDA) Forest Service.
Federal Wildland Fire Management Policy and Program Review (FWFMP).
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Sequoia and King’s Canyon National Parks. 13 February 2006. “Giant Sequoias and Fire.”
  • Wisheu, I.C., M.L. Rosenzweig, L. Olsvig-Whittaker, A. Shmida. 2000. What makes nutrient-poor Mediterranean heathlands so rich in plant diversity? Evolutionary Ecology Research 2: 935-955.

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