Fire ecologyFire ecology is a scientific discipline concerned with naturalprocesses involving fire in an ecosystem and the ecological effects,the interactions between fire and the abiotic and biotic components ofan ecosystem, and the role of fire as an ecosystem process. Manyecosystems, particularly prairie, savanna, chaparral and coniferousforests, have evolved with fire as an essential contributor to habitatvitality and renewal.[1] Many plant species in fire-affectedenvironments require fire to germinate, establish, or to reproduce.Wildfire suppression not only eliminates these species, but also theanimals that depend upon them.[2]

Campaigns in the United States have historically molded publicopinion to believe that wildfires are always harmful to nature. Thisview is based on the outdated belief that ecosystems progress towardan equilibrium and that any disturbance, such as fire, disrupts the harmony of nature.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 withinthese communities have adapted to withstand, and even to exploit, natural wildfire. More generally, fire is now regarded as a 'naturaldisturbance', similar to flooding, wind-storms, and landslides, that has driven the evolution of species and controls the characteristicsof ecosystems.[3] The map below right, shows one view of how ecosystem type in the United States has a characteristic frequency offire, ranging from once every 10 years to once every 500 years. Natural disturbances can be described by key factors such asfrequency, intensity and area.[3] The map also shows intensity, since some fires are understory fires (light burns that affect mostlyunderstory plants) while others are stand replacement fires (intense fires that tend to kill the adult trees as well.)

Fire suppression, in combination with other human-caused environmental changes, may have resulted in unforeseen consequences fornatural ecosystems. Some large wildfires in the United States have been blamed on years of fire suppression and the continuingexpansion of people into fire-adapted ecosystems, but climate change is more likely responsible.[4] Land managers are faced withtough questions regarding how to restore a natural fire regime, but allowing wildfires to burn is the least expensive and likely mosteffective method.[5]

The Old Fire burning in the San BernardinoMountains (image taken from the InternationalSpace Station)

1 Fire components

2 Abiotic responses

3 Biotic responses and adaptations3.1 Plants

3.1.1 Fire intolerance3.1.2 Fire tolerance3.1.3 Fire resistance

3.2 Animals, birds and microbes

4 Fire and ecological succession

5 Some examples of fire in different ecosystems5.1 Forests

5.1.1 Forests in British Columbia

5.2 Shrublands5.2.1 California shrublands5.2.2 South African Fynbos shrublands

5.3 Grasslands5.3.1 South African savanna

5.4 Longleaf pine savannas5.5 Fire in wetlands

Map of potential different regimes of natural burning in natural ecosystems of 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. This map shouldnot be considered definitive.

Contents

6 Fire suppression6.1 Chaparral communities6.2 Fish impacts

7 Fire as a management tool7.1 The Great Plains shortgrass prairie7.2 Mixed conifer forests in the US Sierra Nevada7.3 Finnish boreal forests7.4 Australian eucalypt forests

8 Management policies8.1 United States

9 See also

10 Footnotes

11 Bibliography

12 External links

A fire regime describes the characteristics of fire and how it interacts with a particular ecosystem.[6] Its "severity" is a term thatecologists use to refer to the impact that a fire has on an ecosystem. Ecologists can define this in many ways, but one way is throughan estimate of plant mortality. Fire can burn at three levels. Ground fires will burn through soil that is rich in organic matter. Surfacefires will burn through dead plant material that is lying on the ground. Crown fires will burn in the tops of shrubs and trees.Ecosystems generally experience a mix of all three.[7]

Fires will often break out during a dry season, but in some areas wildfires may also commonly occur during a time of year whenlightning is prevalent. The frequency over a span of years at which fire will occur at a particular location is a measure of howcommon wildfires are in a given ecosystem. It is either defined as the average interval between fires at a given site, or the averageinterval between fires in an equivalent specified area.[7]

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.[8]

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 effectson the plant community associated with it.[9] By removing overhead vegetation, firecan lead to increased solar radiation on the soil surface by day, resulting in greaterwarming, and to greater cooling through the loss of radiative heat at night. Fewerleaves left to intercept rain will allow more moisture to reach the soil surface. Inaddition, plant transpiration (the process by which water travels through plants andevaporates through pores in the leaves) will be reduced following a fire, allowing thesoil to retain more moisture.

Fire components

Radiata Pine forest burnt during the2003 Bogong Bushfires, Australia

Abiotic responses

Fire may cause changes in soil nutrients through a variety of mechanisms, including oxidation, volatilization, erosion, and leachingby water. Temperatures must be very high, however, to cause a significant loss of nutrients, which are often replaced by organicmatter left behind in the fire. Also, the minerals which were formerly part of slowly decaying plant matter become more soluble andavailable in the ash. 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 athigh temperatures, fire can even alter the texture and structure of soils by affecting the clay content and the soil's porosity.

Plants have evolved many adaptations to cope with fire. Of these adaptations, one ofthe best-known is likely pyriscence, where maturation and release of seeds istriggered, in whole or in part, by fire or smoke; this behaviour is often erroneouslycalled serotiny, although this term truly denotes the much broader category of seedrelease activated by any stimulus. All pyriscent plants are serotinous, but not allserotinous plants are pyriscent (some are necriscent, hygriscent, xeriscent, soliscent,or some combination thereof). On the other hand, germination of seed activated bytrigger is not to be confused with pyriscence; it is known as physiological dormancy.

In chaparral communities in Southern California, for example, some plants haveleaves coated in flammable oils that encourage an intense fire [10]. This heat causestheir fire-activated seeds to germinate (an example of dormancy) and the young plants can then capitalize on the lack of competitionin a burnt landscape. Other plants have smoke-activated seeds, or fire-activated buds. The cones of the Lodgepole pine (Pinuscontorta) are, conversely, pyriscent: they are sealed with a resin that a fire melts away, releasing the seeds.[11] Many plant species,including the shade-intolerant giant sequoia (Sequoiadendron giganteum), require fire to make gaps in the vegetation canopy that willlet in light, allowing their seedlings to compete with the more shade-tolerant seedlings of other species, and so establishthemselves.[12] Because their stationary nature precludes any fire avoidance, plant species may only be fire-intolerant, fire-tolerant orfire-resistant.[13]

Fire-intolerant plant species tend to be highly flammable and are destroyed completely by fire. Some of these plants and their seedsmay simply fade from the community after a fire and not return, others have adapted to ensure that their offspring survives into thenext generation. "Obligate seeders" are plants with large, fire-activated seed banks that germinate, grow, and mature rapidlyfollowing a fire, in order to reproduce and renew the seed bank before the next fire.[13][14] Seeds may contain the receptor proteinKAI2, that is activated by the growth hormones karrikin released by the fire.[15]

Fire-tolerant species are able to withstand a degree of burning and continue growingdespite damage from fire. These plants are sometimes referred to as "resprouters."Ecologists have shown that some species of resprouters store extra energy in theirroots to aid recovery and re-growth following a fire.[13][14] For example, after anAustralian bushfire, the Mountain Grey Gum tree (Eucalyptus cypellocarpa) startsproducing a mass of shoots of leaves from the base of the tree all the way up thetrunk towards the top, making it look like a black stick completely covered withyoung, green leaves.

Biotic responses and adaptations

Plants

Lodgepole pine cones

Fire intolerance

Fire tolerance. Typical regrowth afteran Australian bushfire

Fire tolerance

Fire-resistant plants suffer little damage during a characteristic fire regime. These include large trees whose flammable parts are highabove surface fires. Mature ponderosa pine (Pinus ponderosa) is an example of a tree species that suffers virtually no crown damageunder a naturally mild fire regime, because it sheds its lower, vulnerable branches as it matures.[13][16]

Like plants, animals display a range of abilities to cope with fire, but they differfrom plants in that they must avoid the actual fire to survive. Although birds arevulnerable when nesting, they are generally able to escape a fire; indeed they oftenprofit from being able to take prey fleeing from a fire and to recolonize burned areasquickly afterwards. Some birds are potentially responsible for intentional firepropagation to flush out prey.[17] Mammals are often capable of fleeing a fire, orseeking cover if they can burrow. Amphibians and reptiles may avoid flames byburrowing into the ground or using the burrows of other animals. Amphibians inparticular are able to take refuge in water or very wet mud.[13] Some arthropods alsotake shelter during a fire, although the heat and smoke may actually attract some ofthem, to their peril.[18] Microbial organisms in the soil vary in their heat tolerancebut are more likely to be able to survive a fire the deeper they are in the soil. A lowfire intensity, a quick passing of the flames and a dry soil will also help. An increase in available nutrients after the fire has passedmay result in larger microbial communities than before the fire.[9] The generally greater heat tolerance of bacteria relative to fungimakes it possible for soil microbial population diversity to change following a fire, depending on the severity of the fire, the depth ofthe microbes in the soil, and the presence of plant cover.[19] Certain species of fungi, such as Cylindrocarpon destructans appear tobe unaffected by combustion contaminants, which can inhibit re-population of burnt soil by other microorganisms, and therefore havea higher chance of surviving fire disturbance and then recolonizing and out-competing other fungal species afterwards.[20]

Fire behavior is different in every ecosystem and the organisms in those ecosystems have adapted accordingly. One sweepinggenerality is that in all ecosystems, fire creates a mosaic of different habitat patches, with areas ranging from those having just beenburned to those that have been untouched by fire for many years. This is a form of ecological succession in which a freshly burnedsite will progress through continuous and directional phases of colonization following the destruction caused by the fire.[21]

Ecologists usually characterize succession through the changes in vegetation that successively arise. After a fire, the first species tore-colonize will be those with seeds are already present in the soil, or those with seeds are able to travel into the burned area quickly.These are generally fast-growing herbaceous plants that require light and are intolerant of shading. As time passes, more slowlygrowing, shade-tolerant woody species will suppress some of the herbaceous plants.[22] Conifers are often early successional species,while broad leaf trees frequently replace them in the absence of fire. Hence, many conifer forests are themselves dependent uponrecurring fire.[3]

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

Fire resistance

Animals, birds and microbes

A mixed flock of hawks hunting inand around a bushfire

Fire and ecological succession

Some examples of fire in different ecosystems

Forests

Mild to moderate fires burn in the forest understory, removing small trees and herbaceous groundcover. High-severity fires will burninto the crowns of the trees and kill most of the dominant vegetation. Crown fires may require support from ground fuels to maintainthe fire in the forest canopy (passive crown fires), or the fire may burn in the canopy independently of any ground fuel support (anactive crown fire). High-severity fire creates complex early seral forest habitat, or snag forest with high levels of biodiversity. When aforest burns frequently and thus has less plant litter build-up, below-ground soil temperatures rise only slightly and will not be lethalto roots that lie deep in the soil.[18] Although other characteristics of a forest will influence the impact of fire upon it, factors such asclimate and topography play an important role in determining fire severity and fire extent.[23] Fires spread most widely duringdrought years, are most severe on upper slopes and are influenced by the type of vegetation that is growing.

In Canada, forests cover about 10% of the land area and yet harbor 70% of the country’s bird and terrestrial mammal species. Naturalfire regimes are important in maintaining a diverse assemblage of vertebrate species in up to twelve different forest types in BritishColumbia.[24] Different species have adapted to exploit the different stages of succession, regrowth and habitat change that occursfollowing 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.[24]

Shrub fires typically concentrate in the canopy and spread continuously if the shrubsare close enough together. Shrublands are typically dry and are prone toaccumulations of highly volatile fuels, especially on hillsides. Fires will follow thepath of least moisture and the greatest amount of dead fuel material. Surface andbelow-ground soil temperatures during a burn are generally higher than those offorest fires because the centers of combustion lie closer to the ground, although thiscan vary greatly.[18] Common plants in shrubland or chaparral include manzanita,chamise and Coyote Brush.

California shrubland, commonly known as chaparral, is a widespread plantcommunity of low growing species, typically on arid sloping areas of the CaliforniaCoast Ranges or western foothills of the Sierra Nevada. There are a number of common shrubs and tree shrub forms in thisassociation, including salal, toyon, coffeeberry and Western poison oak.[25] Regeneration following a fire is usually a major factor inthe association of these species.

Fynbos shrublands occur in a small belt across South Africa. The plant species in this ecosystem are highly diverse, yet the majorityof these species are obligate seeders, that is, a fire will cause germination of the seeds and the plants will begin a new life-cyclebecause of it. These plants may have coevolved into obligate seeders as a response to fire and nutrient-poor soils.[26] Because fire iscommon in this ecosystem and the soil has limited nutrients, it is most efficient for plants to produce many seeds and then die in thenext 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 thenutrient-poor soil would be less efficient. It is possible that the rapid generation time that these obligate seeders display has led tomore rapid evolution and speciation in this ecosystem, resulting in its highly diverse plant community.[26]

Grasslands burn more readily than forest and shrub ecosystems, with the fire moving through the stems and leaves of herbaceousplants and only lightly heating the underlying soil, even in cases of high intensity. In most grassland ecosystems, fire is the primarymode of decomposition, making it crucial in the recycling of nutrients.[18]

Forests in British Columbia

Shrublands

Lightning-sparked wildfires arefrequent occurrences on shrublandsand grasslands in Nevada.

California shrublands

South African Fynbos shrublands

Grasslands

In the savanna of South Africa, recently burned areas have new growth that provides palatable and nutritious forage compared toolder, tougher grasses. This new forage attracts large herbivores from areas of unburned and grazed grassland that has been kept shortby constant grazing. On these unburned "lawns", only those plant species adapted to heavy grazing are able to persist; but thedistraction provided by the newly burned areas allows grazing-intolerant grasses to grow back into the lawns that have beentemporarily abandoned, so allowing these species to persist within that ecosystem.[27]

Much of the southeastern United States was once open longleaf pine forest with a rich understory ofgrasses, sedges, carnivorous plants and orchids. The above maps shows that these ecosystems (coded aspale blue) had the highest fire frequency of any habitat, once per decade or less. Without fire, deciduousforest trees invade, and their shade eliminates both the pines and the understory. Some of the typical plantsassociated with fire include Yellow Pitcher Plant and Rose pogonia. The abundance and diversity of suchplants is closely related to fire frequency. Rare animals such as gopher tortoises and indigo snakes alsodepend upon these open grasslands and flatwoods.[28] Hence, the restoration of fire is a priority to maintainspecies composition and biological diversity.[29]

Although it may seem strange, many kinds of wetlands are also influenced by fire. This usually occursduring periods of drought. In landscapes with peat soils, such as bogs, the peat substrate itself may burn,leaving holes that refill with water as new ponds. Fires that are less intense will remove accumulated litterand allow other wetland plants to regenerate from buried seeds, or from rhizomes. Wetlands that areinfluenced by fire include coastal marshes, wet prairies, peat bogs, floodplains, prairie marshes andflatwoods.[30] Since wetlands can store large amounts of carbon in peat, the fire frequency of vast northern peatlands is linked toprocesses controlling the carbon dioxide levels of the atmosphere, and to the phenomenon of global warming.[31]

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

In 2003, southern California witnessed powerful chaparral wildfires. Hundreds of homes and hundreds of thousands of acres of landwent up in flames. Extreme fire weather (low humidity, low fuel moisture and high winds) and the accumulation of dead plantmaterial from 8 years of drought, contributed to a catastrophic outcome. Although some have maintained that fire suppressioncontributed to an unnatural buildup of fuel loads,[32] a detailed analysis of historical fire data has showed that this may not have beenthe case.[33] Fire suppression activities had failed to exclude fire from the southern California chaparral. Research showingdifferences in fire size and frequency between southern California and Baja has been used to imply that the larger fires north of theborder are the result of fire suppression, but this opinion has been challenged by numerous investigators and is no longer supportedby 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 quicklycolonized burned areas, especially those that had already been burned in the previous 15 years. Because shrubs in these communitiesare adapted to a particular historical fire regime, altered fire regimes may change the selective pressures on plants and favor invasiveand non-native species that are better able to exploit the novel post-fire conditions.[34]

South African savanna

Longleaf pine savannas

Yellow pitcherplant isdependent uponrecurring fire incoastal plainsavannas andflatwoods.

Fire in wetlands

Fire suppression

Chaparral communities

The Boise National Forest is a US national forest located north and east of the city of Boise, Idaho. Following severaluncharacteristically large wildfires, an immediately negative impact on fish populations was observed, posing particular danger tosmall and isolated fish populations.[35] In the long term, however, fire appears to rejuvenate fish habitats by causing hydraulicchanges 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.[35] But although fire generally appears favorable for fishpopulations in these ecosystems, the more intense effects of uncharacteristic wildfires, in combination with the fragmentation ofpopulations by human barriers to dispersal such as weirs and dams, will pose a threat to fish populations.

Restoration ecology is the name given to an attempt to reverse or mitigate some of the changes that humans have caused to anecosystem. Controlled burning is one tool that is currently receiving considerable attention as a means of restoration andmanagement. 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. However, there is debate asto what state managers should aim to restore their ecosystems to, especially as to whether "natural" means pre-human or pre-European. Native American use of fire, not natural fires, historically maintained the diversity of the savannas of NorthAmerica.[36][37] When, how, and where managers should use fire as a management tool is a subject of debate.

A combination of heavy livestock grazing and fire-suppression has drastically altered the structure, composition, and diversity of theshortgrass prairie ecosystem on the Great Plains, allowing woody species to dominate many areas and promoting fire-intolerantinvasive species. In semi-arid ecosystems where the decomposition of woody material is slow, fire is crucial for returning nutrients tothe 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 atincreasing the grass and forb cover, biodiversity and plant nutrient uptake in shortgrass prairies.[38] Managers must also take intoaccount, however, how invasive and non-native species respond to fire if they want to restore the integrity of a native ecosystem. Forexample, fire can only control the invasive spotted knapweed (Centaurea maculosa) on the Michigan tallgrass prairie in the summer,because this is the time in the knapweed's life cycle that is most important to its reproductive growth.[39]

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. Lower elevations had more frequent fire return intervals, whilst higher and wetter elevations sawmuch longer intervals between fires. Native Americans tended to set fires during fall and winter, and land at a higher elevation wasgenerally occupied by Native Americans only during the summer.[40]

The decline of habitat area and quality has caused many species populations to be red-listed by the International Union forConservation of Nature. According to a study on forest management of Finnish boreal forests, improving the habitat quality of areasoutside reserves can help in conservation efforts of endangered deadwood-dependent beetles. These beetles and various types offungi both need dead trees in order to survive. Old growth forests can provide this particular habitat. However, most Fennoscandianboreal forested areas are used for timber and therefore are unprotected. The use of controlled burning and tree retention of a forestedarea with deadwood was studied and its effect on the endangered beetles. The study found that after the first year of management thenumber of species increased in abundance and richness compared to pre-fire treatment. The abundance of beetles continued toincrease the following year in sites where tree retention was high and deadwood was abundant. The correlation between forest firemanagement and increased beetle populations shows a key to conserving these red-listed species.[41]

Fish impacts

Fire as a management tool

The Great Plains shortgrass prairie

Mixed conifer forests in the US Sierra Nevada

Finnish boreal forests

Much of the old growth eucalypt forest in Australia is designated for conservation. Management of these wet tropical forests isimportant because species like Eucalyptus grandis rely on fire to survive. There are a few eucalypt species that do not have alignotuber, a root swelling structure that contains buds where new shoots can then sprout. During a fire a lignotuber is helpful in thereestablishment of the plant. Because some eucalypts do not have this particular mechanism, forest fire management can be helpfulby creating rich soil, killing competitors, and allowing seeds to be released.[42]

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 thatmany ecosystems depend on disturbance for their diversity and for the proper maintenance of their natural processes. Althoughhuman safety is still the number one priority in fire management, new US government objectives include a long-term view ofecosystems. The newest policy allows managers to gauge the relative values of private property and resources in particular situationsand to set their priorities accordingly.[11]

One of the primary goals in fire management is to improve public education in order to suppress the "Smokey Bear" fire-suppressionmentality and introduce the public to the benefits of regular natural fires.

FynbosResprouter

Crown sproutingLignotuber

WildfireWildfire ecology indexEvolutionary history of plants

California chaparral and woodlands

Peat bog fire

1. "The Ecological Importance of Mixed-Severity Fires - ScienceDirect" (http://www.sciencedirect.com/science/book/9780128027493). www.sciencedirect.com. Retrieved 2016-08-26.

2. Hutto, Richard L. (2008-12-01). "The Ecological Importance of Severe Wildfires: Some Like It Hot" (http://onlinelibrary.wiley.com/doi/10.1890/08-0895.1/abstract). Ecological Applications. 18 (8): 1827–1834. ISSN 1939-5582 (https://www.worldcat.org/issn/1939-5582). doi:10.1890/08-0895.1 (https://doi.org/10.1890%2F08-0895.1).

3. Keddy 2007, Chapter 6

4. Westerling, A. L.; Hidalgo, H. G.; Cayan, D. R.; Swetnam, T. W. (2006-08-18). "Warming and Earlier Spring IncreaseWestern U.S. Forest Wildfire Activity" (http://science.sciencemag.org/content/313/5789/940). Science. 313 (5789):940–943. ISSN 0036-8075 (https://www.worldcat.org/issn/0036-8075). PMID 16825536 (https://www.ncbi.nlm.nih.gov/pubmed/16825536). doi:10.1126/science.1128834 (https://doi.org/10.1126%2Fscience.1128834).

5. Noss, Reed F.; Franklin, Jerry F.; Baker, William L.; Schoennagel, Tania; Moyle, Peter B. (2006-11-01). "Managingfire-prone forests in the western United States" (http://onlinelibrary.wiley.com/doi/10.1890/1540-9295(2006)4%5b481:MFFITW%5d2.0.CO;2/abstract). Frontiers in Ecology and the Environment. 4 (9): 481–487. ISSN 1540-9309 (https://www.worldcat.org/issn/1540-9309). doi:10.1890/1540-9295(2006)4[481:MFFITW]2.0.CO;2 (https://doi.org/10.1890%2F1540-9295%282006%294%5B481%3AMFFITW%5D2.0.CO%3B2).

Australian eucalypt forests

Management policies

United States

See also

Footnotes

6. Whitlock, Cathy; Higuera, P.E.; McWethy, D.B.; Briles, C.E. (2010). "Paleoecological Perspectives on Fire Ecology:Revisiting the Fire-Regime Concept" (http://www.benthamscience.com/open/toecolj/articles/V003/S10001TOECOLJ/6TOECOLJ.pdf) (PDF). Open Journal of Ecology. 3: 6–23. doi:10.2174/1874213001003020006 (https://doi.org/10.2174%2F1874213001003020006). Retrieved 13 June 2013.

7. Bond and Keeley 2005

8. Byram, 1959

9. Hart et al. 2005

10. https://www.nps.gov/fire/wildland-fire/learning-center/fire-in-depth/different-ecosystems/chaparral-sw.cfm

11. USDA Forest Service

12. US National Park Service

13. Kramp et al. 1986

14. Knox and Clarke 2005

15. "Smoke signals: How burning plants tell seeds to rise from the ashes" (http://www.salk.edu/news/pressrelease_details.php?press_id=613). Salik researchers. Salk Institute for Biological Studies. April 29, 2013. Retrieved 2013-04-30.

16. Pyne 2002

17. Gosford, Robert (Nov 2015). "Ornithogenic Fire: Raptors as Propagators of Fire in the Australian Savanna" (http://www.raptorresearchfoundation.org/files/2015/11/2015_conference_program.pdf) (PDF). 2015 Raptor ResearchFoundation Annual Conference, Nov 4 – 8 , Sacramento, California. Retrieved 23 February 2017.

18. DeBano et al. 1998

19. Andersson, Michael (5 May 2014). "Tropical savannah woodland: effects of experimental fire on soil microorganismsand soil emissions of carbon dioxide" (http://www.sciencedirect.com/science/article/pii/S0038071704000513). SoilBiology and Biochemistry. 36 (5): 849–858. doi:10.1016/j.soilbio.2004.01.015 (https://doi.org/10.1016%2Fj.soilbio.2004.01.015). Retrieved 2 June 2016.

20. Widden, P (March 1975). "The effects of a forest fire on soil microfungi". Soil Biology and Biochemistry. 7 (2): 125–138. doi:10.1016/0038-0717(75)90010-3.) (https://doi.org/10.1016%2F0038-0717%2875%2990010-3.%29).

21. Begon et al. 1996, pg. 692

22. Begon et al. 1996, pg 700

23. Beaty and Taylor (2001)

24. Bunnell (1995)

25. C.Michael Hogan (2008) "Western poison-oak: Toxicodendron diversilobum" (http://globaltwitcher.auderis.se/artspec_information.asp?thingid=82914) Archived (https://web.archive.org/web/20090721044257/http://globaltwitcher.auderis.se/artspec_information.asp?thingid=82914) July 21, 2009, at the Wayback Machine., GlobalTwitcher, ed. NicklasStrömberg

26. Wisheu et al. (2000)

27. Archibald et al. 2005

28. Means, D. Bruce. 2006. Vertebrate faunal diversity in longleaf pine savannas. Pages 155-213 in S. Jose, E. Jokelaand D. Miller (eds.) Longleaf Pine Ecosystems: Ecology, Management and Restoration. Springer, New York. xii +438 pp.

29. Peet, R. K. and Allard, D. J. (1993). Longleaf pine vegetation of the southern Atlantic and eastern Gulf Coastregions: a preliminary classification. In The Longleaf Pine Ecosystem: Ecology, Restoration and Management, ed. S.M. Hermann, pp. 45–81. Tallahassee, FL: Tall Timbers Research Station.

30. Keddy 2010, p. 114-120.

31. Vitt et al. 2005

32. Minnich 1983

33. Keeley et al. 1999

34. Keeley et al. 2005

35. Burton (2005)

36. MacDougall et al. (2004)

37. Williams, Gerald W. (2003-06-12). "REFERENCES ON THE AMERICAN INDIAN USE OF FIRE IN ECOSYSTEMS"(https://web.archive.org/web/20080706160607/http://www.blm.gov/heritage/docum/Fire/Bibliography%20-%20Indian%20Use%20of%20Fire.pdf) (PDF). Archived from the original (http://www.blm.gov/heritage/docum/Fire/Bibliography%20-%20Indian%20Use%20of%20Fire.pdf) (PDF) on 2008-07-06. Retrieved 2008-07-31.

38. Brockway et al. 2002

39. Emery and Gross (2005)

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