United States

Department of

Agriculture

Forest Service

Pacific Southwest

Research Station

General Technical

Report

PSW-GTR-224

September 2009

Ecological Effects of Prescribed

Fire Season: A Literature Review

and Synthesis for Managers

Eric E. Knapp, Becky L. Estes, and Carl N. Skinner
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Authors

Eric E. Knapp is a research ecologist, Becky L. Estes is a research ecologist, and

Carl N. Skinner is a research geographer, U.S. Department of Agriculture, Forest

Service, Pacific Southwest Research Station, 3644 Avtech Parkway, Redding, CA

96002.

Cover photos from left to right by Eric Knapp, Quinn Long, and Ron Masters.
Abstract

Knapp, Eric E.; Estes, Becky L.; Skinner, Carl N. 2009. Ecological effects of

prescribed fire season: a literature review and synthesis for managers. Gen.

Tech. Rep. PSW-GTR-224. Albany, CA: U.S. Department of Agriculture,

Forest Service, Pacific Southwest Research Station. 80 p.

Prescribed burning may be conducted at times of the year when fires were infrequent historically, leading to concerns about potential adverse effects on vegetation

and wildlife. Historical and prescribed fire regimes for different regions in the

continental United States were compared and literature on season of prescribed

burning synthesized. In regions and vegetation types where considerable differences

in fuel consumption exist among burning seasons, the effects of prescribed fire

season appears, for many ecological variables, to be driven more by fire-intensity

differences among seasons than by phenology or growth stage of organisms at the

time of fire. Where fuel consumption differs little among burning seasons, the effect

of phenology or growth stage of organisms is often more apparent, presumably

because it is not overwhelmed by fire-intensity differences. Most species in ecosystems that evolved with fire appear to be resilient to one or few out-of-season

prescribed burn(s). However, a variable fire regime including prescribed burns at

different times of the year may alleviate the potential for undesired changes and

maximize biodiversity.

Keywords: Fire effects, fire intensity, fire season, fuel consumption, historical

fire regime, phenology, prescribed fire, pyrodiversity.
Contents

1 Chapter 1: Overview

5 Chapter 2: Introduction

5 The Fire Season Issue

9 Chapter 3: Western Region

9 Climate, Vegetation, and Fire

9 Humid Temperate

11 Dry Interior

15 Fuel Consumption and Fire Intensity

15 Ecological Effects of Burning Season in Forested Ecosystems

15 Trees

18 Understory Vegetaton

20 Soils

21 Wildlife

23 Ecological Effects of Burning Season in Chaparral and Grasslands

23 Chaparral

25 Western Grasslands

26 Implications for Managers

29 Chapter 4: Central Region

29 Climate, Vegetation, and Fire

29 Historical Fire Regime

32 Prescribed Fire Regime

33 Fuel Consumption and Fire Intensity

35 Ecological Effects of Burning Season

35 Grassland Vegetation

38 Soils

38 Wildlife

40 Implications for Managers

43 Chapter 5: Eastern Region

43 Climate, Vegetation, and Fire

43 Subtropical

48 Hot Continental and Warm Continental

50 Fuel Consumption and Fire Intensity

50 Ecological Effects of Burning Season

50 Trees

53 Understory Vegetation

57 Soils

57 Wildlife

60 Implications for Managers

61 Acknowledgments

62 Metric Equivalents

62 Literature Cited
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Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

Chapter 1: Overview

In some areas of the United States, most fires historically occurred when plants were dormant and animals had

reproduced and dispersed. This includes the Western

United States, where fires were historically most abundant

during the months of the year with the driest fuels and after

senescence of surface vegetation, and the forests of the

Northeast, where fallen leaves of deciduous trees are the

main carrier of fire. On the other hand, in the Southwestern

United States, the main historical fire season was toward

the end of the dry season (late spring/early summer), in

association with the first thunderstorms, which ignited

the fires but also provided moisture for plants to initiate

growth. In the Southeastern United States, historical fires

were once common throughout the summer and peaked

in May at the transition from the dry spring period to the

wet summer period, when lightning incidence was at its

highest, vegetation was growing, and animals were active.

Prescribed fires may not only differ from natural fires in

their timing relative to phenology (seasonal growth or life

history stage) of organisms that live in the ecosystem, but

may also often differ in their intensity. For example, in the

Western United States, prescribed burns are increasingly

conducted in the spring, when many of the larger surface

fuels are still somewhat moist from the winter and spring

precipitation. Because of the higher moisture, prescribed

burns at this time of year tend to consume less fuel and

therefore release less heat. Thus, to evaluate the effect of

burn season, both the role of differences in intensity and

timing between prescribed fire and natural fire need to be

considered. Although burn season research results that

have controlled for fire intensity have often shown an effect

Prescribed burning is a tool for reducing fuels and restoring

a disturbance process to landscapes that historically experienced fire. It is often assumed, or at least desired, that

the effects of prescribed burns mimic those of natural fires.

However, because of operational and liability constraints,

a significant proportion of prescribed burning is, in many

ecosystems, conducted at different times of the year than

when the majority of the landscape burned historically.

This has brought into question the extent to which prescribed fire mimics effects of the historical fire-disturbance

regime, and whether there are any negative impacts of such

out-of-season burning.

Most plant and animal species that exist in areas with

a history of relatively frequent low- to moderate-intensity

fire are resilient to its effects. However, burning season

may influence the outcome in a number of ways. For example, many plant species recover quickly from fire, either

through resprouting or fire-stimulated seed germination,

but it is believed that the recovery can differ depending

on the timing of the fire. When aboveground parts are

consumed or killed by the fire, resprouting depends on

stored resources, such as carbohydrates. These carbohydrates are typically at their lowest annual levels early in

the growing season. Thus, plants may recover more slowly

from fire that occurs during the active growing season than

fire that occurs after plants have gone dormant. Animal

species can often avoid the flames; however, they may be

more vulnerable to fire at times of reduced mobility, such

as during nesting or breeding season. The influence of fire

season can also be indirect, through differences in habitat

created, or competitive release of some species owing to

damage to or mortality of others.
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GENERAL TECHNICAL REPORT PSW-GTR-224

of fire timing, the latest research suggests that, in many

cases, variation in fire intensity exerts a stronger influence

on the ecosystem than variation in fire timing.

Given the potential importance of fire intensity to fire

effects, a useful means of evaluating the outcome of prescribed burn season relative to what might have been expected under a natural fire regime would be to consider

the amount of fuel consumed by prescribed burns and the

intensity of those burns at different times of the year, in

relation to the amount of fuel that was likely consumed by

and the intensity of historical fires (both lightning ignited

and anthropogenic) (table 1).

In forest ecosystems of the Western United States,

prescribed burns are often conducted in areas with very

heavy fuel loads resulting from decades of fire exclusion.

Although spring prescribed burns typically consume less

fuel than those that are ignited in other seasons, prescribed

burns in any season can conceivably consume more fuel

than historical burns would have under a natural fire regime. Several recent papers have shown that late summer

or fall prescribed burns often lead to higher tree mortality

and set back herbaceous understory vegetation more than

spring burns, even though late summer and early fall fire

was the historical norm. The difference in fuel consumption and fire intensity between the prescribed burn seasons apparently overwhelmed the effect of phenology of

the organisms. Many coniferous forest ecosystems of the

Southwest also typically have unnaturally high fuel loads,

but times of the year with lower fuel moisture and higher

consumption differs, owing to monsoon rains in the

summer. Until fuels are reduced to historical levels, any

prescribed burn under higher fuel moisture conditions may

have effects more similar to historical burns, because the

amount of fuel consumed, and fire intensity are closer to

that noted for historical burns. A different situation exists

in chaparral shrub lands of the West, where prescribed

burns are usually conducted under more benign conditions

in the winter or spring, and are therefore often less intense

and consume less fuel than historical fires would have.

With organisms in these shrub ecosystems presumably

adapted to high-severity stand-replacing fire, reduced

intensity over what might have been experienced historically also means that the outcomes sometimes have not

met objectives. For example, several authors have noted

that shrubs and herbs requiring intense heat to stimulate

germination emerge in lesser numbers following spring

burns.

Grasslands are composed of fine fuels that dry readily

and are likely to be nearly completely consumed with prescribed fire in any season (table 1). Grass thatch also breaks

down relatively rapidly, so there is not a large buildup of

fuels relative to historical levels. Because the difference

Table 1—Historical and prescribed fire seasons plus potential fuel consumption differences between dormant- and

growing-season prescribed burnsa

Main historical Main prescribed Typical potential fuel consumption difference

Region fire season fire season between dormant and growing season burns

Western forests Dormant Dormant/growing Very high

Southwestern forests Growing/dormantb Dormant High

Central grasslands Dormant/growing Dormant Low

Southeastern pine forests Growing Dormant/growing Moderate

Eastern hardwood forests Dormant Dormant Low to moderate

a

 Much variation in conditions at the time of burning exists within both the historical and prescribed fire regimes for each region—the listed text is simply a rough

average.

b

 When multiple seasons are reported, the order indicates the most likely.
3

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

in total fuel consumption and fire intensity between burn

seasons is relatively low, the effect of timing of the fire is

generally more evident in grasslands than in other vegetation types. Numerous examples of alterations to grassland

plant communities with prescribed burning in different

seasons are found in the literature.

In the Southeastern United States, prescribed burns are

typically conducted in late winter/early spring when many

plants and other organisms are dormant, and in the late

spring/early summer, during the historical peak period of

lightning-ignited fire. Burning during the dormant season

became standard practice in order to reduce direct impacts

to nesting birds and other wildlife species. However, in

many cases, the prescribed burns during the late spring/

early summer growing season have been shown to better

meet longer term management objectives for pine forests

by reducing competition from competing hardwoods.

Furthermore, concerns about negative effects to wildlife

from late spring/early summer growing-season burns have

generally not been supported by research.

In eastern forests, burn intensity does not generally

vary predictably with season, with fuel consumption influenced more by time since previous rainfall and year-toyear climatic variability. Differences in fuel consumption

among burning seasons is often much less in eastern forests (particularly deciduous forests) than in western forests,

where because of a long history of fire exclusion and a

slower decomposition rate, surface fuel loads are typically

much higher. Therefore, differences among burn seasons

related to fire intensity are expected to be considerably less

in eastern forests than in western forests (table 1).

Many species show strong resilience to fire in either

season, with the majority of studies reporting relatively

minor differences, if any. Differences in the timing of a

single or even several applications of prescribed fire do

not appear likely to substantially change the plant or

animal community. In most ecosystems studied, the change

associated with either burning or not burning is much

greater than differences in the outcome with burning in

different seasons. This should not be interpreted as burning

season not mattering. Burning season has been shown to

affect community composition, particularly with repeated

application of fire in the same time of year. Many authors

have therefore stressed the importance of incorporating

variability in prescribed fire timing (along with variability

in other aspects of the fire regime) into long-term burn

management plans. Because response to burning season

differs a great deal among species, a heterogeneous fire

regime is likely to maximize biodiversity.

One recurring problem in fire management and fire

science is the inconsistency in terminology. Fire timing

may be referred to as a spring burn, fall burn, early-season

burn, late-season burn, wet-season burn, dry-season burn,

growing-season burn, dormant-season burn, or lightningseason burn, each of which may have different meanings

across ecosystems. Furthermore, the phenological status of

target species often differs with latitude and yearly climate.

This creates a serious impediment to truly understanding

and synthesizing the literature on season of burning. To

maximize what can be learned, we recommend that authors

and practitioners should, whenever possible, provide information on exact burn dates, as well as variables such as

weather conditions and year-to-year climatic variation (was

it a drought year?), fuel moistures at the times of burns, fire

behavior (including fire-line intensity), plus the phenological or life-history status of target species.
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GENERAL TECHNICAL REPORT PSW-GTR-224

Key Points

Both fire intensity and burn season can influence fire effects. To evaluate the expected outcome of prescribed burning

season, managers may need to ask the following questions: (1) What is the phenological or life-history stage of

organisms at the time of the prescribed burn and how does this differ from our best approximation of historical

conditions? (2) What are the loading, composition, and architecture of fuels at the site to be burned and how do

these compare with historical conditions? (3) How different will fire intensity be for prescribed burns conducted in

different seasons, and does this vary from historical fire intensity?

• Effects related to the phenology or life history stage of organisms at the time of prescribed burning

are more likely to be noticed if differences in fuel consumption or fire intensity between seasons are

low. If differences in consumption or intensity are substantial, these factors will likely drive fire

effects.

• The burn season leading to an amount of fuel consumed and fire intensity closest to or within the

historical range of variability will often have the best outcome.

• A prescribed burn timed to occur within the historical burn season will often have the best outcome.

• A single prescribed burn (or even a few prescribed burns) outside of the historical fire season

appear(s) unlikely to have strong detrimental effects. Substantial shifts in community composition

often require multiple cycles of prescribed burning. In many ecosystems, the importance of burning

appears to outweigh the effect of burn season.

• Variation in the timing of prescribed burns will help to ensure biodiversity is maintained.
5

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

Chapter 2: Introduction

seed dispersal; resistance to rotting; modified seedling

structure; and thick heat-resistant buds (Abrams 1992,

Bond and van Wilgen 1996, Myers 1990, Wade et al. 2000)

(fig. 1). Understory herbaceous plant species survive fire

through various mechanisms including resprouting from

underground structures such as rhizomes or stolons that are

located deeply enough in the soil to avoid the lethal heat

pulse (Bond and van Wilgen 1996, Flinn and Wein 1977),

or establishing from seeds that are stimulated to germinate

by heat (Kauffman and Martin 1991, Keeley 1987). Other

organisms survive in microenvironments where fire is less

frequent as a result of lower fuel accumulation or where

fuels have higher moisture levels. Among animals, less

mobile species may use stump holes, cracks, or burrows as

refuges when fire passes through, whereas more mobile

species can flee, returning when the danger has passed. The

type of adaptations depends on the fire regime, with, for

example, frequent low-severity regimes requiring a different suite of characteristics than high-severity regimes such

as lodgepole pine (Pinus contorta Dougl. ex Loud.) forest

or chaparral shrublands, where the aboveground stems

typically do not survive.

Fire adaptations may interact with burning season in

several ways. In plants, carbohydrate reserves necessary to

sustain growth are often at their lowest levels shortly after

breaking dormancy (de Groot and Wein 2004, Harrington

1989). Stored carbohydrates help fuel this rapid burst of

growth, and these reserves are generally replenished by

products of photosynthesis during the growing season. It

is thought that plants may have a harder time recovering

from tissue loss to fire during the period when carbohydrate reserves are low than at other times of the year

(Garrison 1972, Hough 1968, Volland and Dell 1981). In

addition, tender early-season tissues may be more sensitive

to heat (Bond and van Wilgen 1996, DeBano et al. 1998).

Fire in the early season can also kill aboveground flowering parts prior to seed production and seed fall, limiting

reproductive capacity. With animals, vulnerability to

The Fire Season Issue

Fire is being reintroduced to many ecosystems that historically experienced frequent fire to reduce hazardous fuels

that have accumulated and to restore important ecological

functions. This reintroduction often occurs through prescribed burning, the assumption being that the disturbance

produced by such fires approximates the disturbance

historically produced by wildfire. However, prescribed

burns are sometimes ignited outside of the historical fire

season. Reasons for this include the following: (1) Safety

concerns. Igniting during times of more benign weather

and fuel moisture conditions lessens the chance of an

escape. (2) Smoke management. Certain times of the year

may be better for smoke dispersal than others. (3) Operational constraints. There may be a lack of resources during

the historical fire season because personnel are being used

to fight wildfires. (4) Biological management. Certain

seasons may reduce the chance of injury and death of

target species.

There has been concern that “out-of-season” burning

might be harmful to some species because the ecosystem

did not evolve with fire during these times. For example,

across much of the Western United States, prescribed burns

are frequently ignited in the spring and early summer,

during the period of active growth of many organisms,

although wildfires were historically uncommon during this

time. In the Southeastern United States, the peak season for

wildfires was historically during the active growth phase of

trees and other vegetation, but prescribed burning is now

more commonly conducted during the late winter when the

majority of vegetation is dormant. Burning in the dormant

season may not effectively control competing midstory

vegetation, thereby reducing the establishment of fireadapted overstory conifers.

Organisms of fire-adapted ecosystems have evolved

and thrive with fire in a multitude of ways. For example,

many trees have one or more of the following characteristics: thick bark; fire-stimulated sprouting, germination or
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GENERAL TECHNICAL REPORT PSW-GTR-224

prescribed fire can differ depending on the time of year.

For example, birds are potentially more strongly impacted

by spring and early summer burns because this coincides

with the nesting season (Reinking 2005). Reptiles and

amphibians may be more active or more likely to be at the

surface at certain times of the year where they are less able

to survive flaming combustion (Griffiths and Christian

1996, Pilliod et al. 2003). Both plant and animal species

may depend on unburned patches to persist (Martin and

Sapsis 1992), and creation of these refugia often differs

among seasons, varying with fuel moisture levels and fuel

continuity.

The response of organisms to prescribed fire depends

on complex interactions between factors such as the timing

of prescribed burning relative to the historical fire season,

phenological stage of the organisms at the time of fire, differences in fire severity among burn seasons, and variation

in climate within and among burn seasons. Many studies

on the timing of prescribed fire only broadly describe the

season of burning (i.e., spring burn), which allows for some

variation with respect to the growth stage of plants and

other organisms (Svejcar 1990). For example, a prescribed

burn very early in the spring, prior to bud break, may have

entirely different effects on vegetation than a prescribed

burn later in the spring after leaves have flushed. In addition, no two prescribed burns are the same, even those

conducted within the same season. Among the limitations

of studies comparing different seasons of burning is that

the timing of treatment is often confounded with other

factors that affect fire intensity and severity at different

times of the year. To best understand the effect of burn season, we present associated data on fire severity, phenology

of vegetation, and activity level/vulnerability of the fauna

of interest at the time of the burns, whenever available.

A B

Figure 1—Adaptations to fire in two pine species of the Southern United States. (a) Young longleaf pine seedlings in the “grass” stage

resemble a tuft of grass, with height growth suppressed and the apical growing points protected from the frequent surface fires. As shown

in the photograph, seedlings can recover from needle scorch during this stage. After development of the tap root, the seedling enters the

candle stage where rapid height growth occurs, moving the terminal bud above average flame height. (b) Shortleaf pine can resprout from

the base following disturbance, increasing resilience to fire. The ability of shortleaf pine to resprout is dependent on tree age and intensity

of the fire.

Stephen Hudson

Ron Masters
7

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

Because of differences in historical and prescribed fire

regime (timing, intensity, vegetation type, spatial scale),

research findings from studies conducted in one area or

vegetation type may not apply to others. In this synthesis,

we therefore cover three broad regions of the continental

United States, adapted roughly from groupings of ecoregional divisions outlined by Bailey (1983), which are

based on both climatic zones and potential natural vegetation. Our regions consider differences in vegetation with

the strongest influence on fuel loading and the fire regime

(fig. 2). The Western region is everything west of the central

grasslands, and consists of both a humid temperate division along the Pacific Coast as well as the non-grassland

portions of the dry interior division. The Central region is

composed of both dry temperate to subtropical steppe

(shortgrass prairie) and humid temperate prairie (tallgrass).

The Eastern region consists of mainly a warm continental

and a hot continental division (boreal and deciduous

forest, respectively), plus a subtropical division (Bailey

1983), dominated by pine and mixed pine-oak forests, and

a savanna division in south Florida. Alaska and Hawaii are

not covered, as little or no information on seasonal differences of prescribed fire is available for either of these two

areas.

Figure 2—Three broad fire regions of the continental United States roughly adapted from ecosystem divisions outlined by

Bailey (1980).
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GENERAL TECHNICAL REPORT PSW-GTR-224
9

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

Chapter 3: Western Region

Historical fire regime—

Prior to fire exclusion, the historical fire-return interval

averaged across all forest types in Washington was 71

years, whereas the fire-return interval in Oregon forests

was estimated to be 42 years (Agee 1993). A great deal

of variability existed among forest types, with mesic

cedar/spruce/hemlock forests burning in mixed to standreplacing fire every 400 to 500+ years (Agee 1993, Brown

2000), whereas drier ponderosa pine forests burned in lowto mixed-severity fires every 15 or so years (Agee 1993).

Many forested regions in California burned even more

frequently in low- to mixed-severity fires at approximately

8- to 30-year intervals, depending on forest type (Skinner

and Chang 1996). In general, the shorter the interval, the

less fuel accumulated between fires, and the lower severity

the average fire. This gradient in fire regime from north to

south is a function of precipitation and temperature patterns. Chaparral shrublands found in central and southern

California typically burned in high-severity standreplacing events at moderate intervals (Keeley 2006).

Owing to the lack of historical records, actual number

of years between fires in chaparral shrub ecosystems is

somewhat uncertain, but estimated to have typically

ranged from 30 to 100 years.1

The wildfire season generally lasts from June until

September in the north, with this period expanding as

one moves south (Schroeder and Buck 1970). Although

wildfires in southern California are most common from

May through November, they can occur in nearly every

month of the year when conditions are dry. In forested

1

Keeley, J.E. 2008. Personal communication. Research ecologist,

U.S. Geological Survey, Sequoia and Kings Canyon Field Station,

47050 Generals Highway, Three Rivers, CA 93271-9651.

Climate, Vegetation, and Fire

Large differences in topography and climate in the Western region naturally lead to a great deal of variation in

fire regime. For the purpose of this synthesis, the Western

region was split into two zones–the Humid Temperate

zone with maritime influence from the Pacific Ocean lying

mainly closer to the coast, and the Dry Interior zone to the

east, with the crest of the Cascade Range and the Sierra

Nevada forming the approximate boundary.

Humid Temperate

This zone is characterized by seasonality in precipitation, with a distinct wet period between approximately

October and April and dry summers (fig. 3 a, c). Because

the warmest months of the year also have the least amount

of precipitation, surface fuels do not decompose as readily

as in some other regions. In the north, average yearly rainfall is high, with the moisture and moderate temperatures

resulting in very productive coniferous forest ecosystems

with heavy fuel accumulation (Schroeder and Buck 1970).

Some summer rains reduce fire hazard in all but the driest

years. The average yearly rainfall generally declines and

temperatures increase as one moves south through this

zone (fig. 3). From approximately Roseburg, Oregon, south,

the climate becomes increasingly mediterranean, with a

defined cool winter rainy season followed by hot, dry

summers. In California, summer rainfall is rare, and fire

hazard is correspondingly higher.

Vegetation within the Humid Temperate zone is highly

complex, varying from mesic hemlock (Tsuga Endl. Carr.),

western redcedar (Thuja plicata Donn ex D. Don), and

Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) forests

in the north to drier mixed-conifer forests and shrublands

in the south.
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GENERAL TECHNICAL REPORT PSW-GTR-224

Figure 3—Climographs (monthly average temperature and precipitation) and the average time of the year of the peak

historical and prescribed fire seasons from four representative locations within the Western region: (a) Crater Lake National

Park, Oregon; (b) Missoula, Montana; (c) Yosemite National Park, California; and (d) Flagstaff, Arizona.
11

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

regions throughout the Humid Temperate zone, growth

ring records from fire-scarred trees indicate that the majority of acres historically burned late in the growing season

or after trees had ceased growth for the year and were dormant (table 2). Late growing season would correspond

approximately to late July through August, whereas

dormancy typically occurs by September in most years

(Fowells 1941). Early to mid growing-season fires (approximately May through July) also occurred, but mainly

in unusually dry years (Norman and Taylor 2003). It is

believed that Native Americans made use of spring burns to

manage vegetation (Lewis 1973), but such fires were likely

less extensive than later lightning-ignited fires under drier

conditions.

Prescribed fire regime—

Prescribed burns are typically conducted in two seasons

either before or after the main period of summer drought

(fig. 3). Early season burns are ignited after the cessation

of winter and spring precipitation or snowmelt, as soon

as the fuels have dried enough to burn (typically mid

April until about July 1), until conditions become too

dry and wildfire season begins in the summer (fig. 3). At

lower elevations below the snowline, prescribed burning

can sometimes also be successfully done during dry

periods within the winter and early spring rainy season

(McCandliss 2002). In black oak (Quercus kelloggii

Newb.)-dominated forests below the snowline, periods

during tree dormancy when the leafless canopy allows

sunlight to dry the leaf litter on the forest floor are often

ideal for burning.2 Spring or early summer prescribed

burning can be problematic because surface fuels are

drying and temperatures warming. Thus, fires may continue to creep and smolder, sometimes for months. The

second prescribed fire season typically occurs in the fall,

after temperatures have cooled and often after the fuels

have moistened with the first rains. In many areas of the

2

 Skinner, C.N. 1995. Using fire to improve wildlife habitat near

Shasta Lake. 26 p. Unpublished report. On file with: USDA Forest

Service, Pacific Southwest Research Station, 3644 Avtech Parkway,

Redding, CA 96002.

West, the fall prescribed fire season coincides with inversions and poor air quality (McCandliss 2002). The spring

and early summer prescribed burning period is generally

earlier than the main historical fire season, and the fall

prescribed burning period is often later than the historical

fire season (fig. 3). Few prescribed burns are conducted

in mid to late summer, the main historical fire season,

because of fire control concerns that can result from the

heavy fuels that characterize many contemporary forest

landscapes. In addition, the summer wildfire season uses

a significant proportion of available firefighting resources,

meaning that fire crews are often unavailable for prescribed burns at this time of year.

The range of ecological conditions under which prescribed burns occur is quite broad. In the coniferous forest

zone, early spring prescribed burns (prior to May) usually

happen prior to active tree and plant growth as well as

other significant biological activity. Burns conducted in

late spring (May to June) occur during the main period

of seasonal growth of vegetation and significant wildlife

activity such as bird nesting (fig. 4a). Late summer and fall

prescribed burns (September to October) typically occur

during the dormant season after biological activity has

slowed or ceased for the year (fig. 4b). Because of the

nearly precipitation-free summers, soils are typically drier

in the late summer and early fall than in the spring or early

summer. However, this is not always the case, and much

depends upon rainfall patterns for that year in relation to

the prescribed burning period. Concerns about prescribed

burning conducted outside of the historical season include

(1) less-than-desired fuel consumption owing to high fuel

moisture levels, and (2) potentially detrimental impacts to

organisms if burns coincide with periods of peak growth/

activity.

Dry Interior

Although the average yearly precipitation is lower in the

Dry Interior zone than in most parts of the Humid Temperate zone, distinct seasonality is also apparent. The western

and northern sections are in the rain shadow of the Cascade
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GENERAL TECHNICAL REPORT PSW-GTR-224

Table 2—Position of fire scars within annual growth rings at different locations in the Western region (from north

to south)a

Approximate time

Before Sept.–

May May June July Aug. Oct.

Location Dormant Early early Mid early Late early Late Dormant Author

Percent of all scars

Pacific northwest:

East Cascades,

Washington 0 19 32 49 Wright and Agee 2004

southwest Montana 0 3 97 Heyerdahl et al. 2006

Blue Mtns., Washington

and Oregon 0 8 20 72 Heyerdahl et al. 2001

California

Shasta Trinity National

Forest 0 1 2 4 17 76 Taylor and Skinner 2003

Whiskeytown National

Recreation Area 0 0 0 7 57 36 Fry and Stephens 2006

Lassen National Forest 0 0 1 10 18 71 Bekker and Taylor 2001

Lassen National Park 0 1 7 8 1 83 Taylor 2000

Plumas National Forest 0 0 1 15 31 53 Moody et al. 2006

Southern Sierra Nevada 0 1 10 12 67 10 Swetnam et al. 1992 b

Sequioa National Park 0 2 3 6 89 Schwilk et al. 2006

San Jacinto Mountains 0 2 2 0 33 63 Everett 2008

Arizona, New Mexico,

and Texas:

Grand Canyon, Arizona 12 12 43 24 19 0 Fulé et al. 2003

Camp Navajo, Arizona 19 21 45 15 0 0 Fulé et al. 1997

Santa Rita Mtns. Arizona 9 30 34 25 2 0 Ortloff 1996

Rincon Mtns., Arizona 12 87 1 0 Baisan and Swetnam 1990

U.S./ Mexico border: 20 41 30 8 1 0 Swetnam et al., in press

Guadalupe Mtns., Texas 6 67 24 1 2 0 Sakulich and Taylor 2007

a

 Timing of the fire (month) is approximate and based on studies of period of radial growth in trees (Fowells 1941, Ortloff 1996, Swetnam et al., in press), which

can vary with elevation, tree species, and yearly climatic differences. Giant sequoia (Sequoiadendron giganteum (Lindl.) J. Buchholz) is thought to have somewhat

later phenology. At sites in Arizona and New Mexico, scars at the ring boundary (dormant) were assumed to have occurred in the spring, prior to tree growth,

whereas at the remainder of sites, scars at the ring boundary were assumed to have occurred in the fall after tree growth was done for the year. b

 Swetnam, T.W.; Baisan, C.H.; Caprio, A.C.; Touchan, R.; Brown, P.M. 1992. Tree ring reconstruction of giant sequoia fire regimes. 173 p. Unpublished report.

On file with: National Park Service, Sequoia and Kings Canyon National Parks, 47050 Generals Highway, Three Rivers, CA 93271.
13

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

Range and the Sierra Nevada, and as a result are characterized by lighter precipitation than the Humid Temperate

zone to the west (fig. 3). The southwest and eastern portions of the Dry Interior are influenced by the summer

monsoon, with two peak times of precipitation—winter

and summer (fig. 3). This monsoonal rainfall is often accompanied by thunderstorms. The monsoon typically

starts out with more scattered high-based storms, which

start fires, whereas the later storms are often wetter

(Schroeder and Buck 1970).

Vegetation is strongly associated with precipitation,

usually along elevation gradients. Forests consisting of

ponderosa pine (Pinus ponderosa Dougl. ex Laws.), or

ponderosa pine mixed with Douglas-fir, and white fir (Abies

concolor (Gord. and Glend.) Lindl. ex Hildebr.) or spruce

(Picea A. Dietr.) at the higher elevations are found on

mountain ranges, whereas the vegetation in the valleys

is often composed of shrubs such as sagebrush, or even

desert vegetation. Pinyon pines (Pinus edulis Engelm.)

or junipers (Juniperus L.) may be found in between.

Historical fire regime—

In the western and northern areas of this zone, such as the

Great Basin, the lightning fire season generally starts in

June and runs through September or October (Schroeder

and Buck 1970) (fig. 3b). The main fire season is somewhat earlier in areas influenced by the monsoon, with area

burned historically peaking in May and June (GrissinoMayer and Swetnam 2000) (table 2, fig. 3d). These fires are

typically ignited by dry high-based thunderstorms that are

common this time of year. As the summer progresses,

thunderstorms begin to be accompanied by more rainfall,

limiting fire spread. Although the fall may be dry enough

for fire as well, thunderstorms are less common and thus

sources of ignition are fewer. Native Americans also surely

contributed to the historical fire regime, and may have

burned at times that did not necessarily coincide with

peak lightning activity.

The peak of the historical fire season in parts of the Dry

Interior zone not strongly affected by the summer monsoon

was similar to the Humid Temperate zone to the west, with

Figure 4—(a) Late spring prescribed burn (June 3, 2008) and (b) fall prescribed burn (October 30, 2008) at Blacks Mountain Experimental Forest, Lassen National Forest, California. Note the phenological stage of the vegetation at the time of the fires. Wildfires in this

area were historically uncommon in the early season, but did occur, especially in dry years. Ten-hour and 1,000-hour fuel moistures were

19 percent, and 52 percent, respectively, at the time of the June burn and 7 percent, and 8 percent, respectively, at the time of the October

burn. Moisture of the top inch of soil was 24 percent in June and 4 percent in October. Both burns were halted prematurely because

objectives were unlikely to be met, with high fuel moisture in June causing too little fuel to be consumed and low fuel moisture in October

leading to unpredictable fire behavior.

A B Eric Knapp Todd Hamilton
14

GENERAL TECHNICAL REPORT PSW-GTR-224

most of the fire occurring when most plants were past the

peak of growth or dormant, and animals presumably less

active. The peak of the historical fire season in areas

strongly influenced by the summer monsoon was approximately the time at which trees begin growth for the year.

Cool-season grasses in the understory are often actively

growing at this time. May and June fires also coincide with

bird nesting.

Prescribed fire regime—

Prescribed burns in juniper or pinyon-juniper woodlands

of Nevada, as well as forested areas farther east and north,

are generally conducted either in the spring or fall (fig.

3b). More days of weather and fuel conditions within

the usual prescription conditions occur during the spring

(Klebenow and Bruner 1977). Cool conditions in either

season moderate fire behavior and reduce crown scorching. However, such prescribed burns typically occur before

or well after the typical historical fire season. In areas influenced by the monsoon in the Southwest, the majority

of prescribed burns are conducted in the cool conditions

of fall (mid-September into December or even later in

years without early snow) (Sackett et al. 1996) (fig. 3d).

Fuels at this time of year are usually fairly dry, but moister

conditions may also occur in some years. Prescribed burns

can also be ignited when the weather is cool in early

spring. Little prescribed burning is done during the

peak historical fire season (late spring to early summer),

because windier and drier weather make fire more difficult

to control, especially when fuel loading is high (Fulé et al.

2007).

Fall is recommended for the initial prescribed burn

after a long period of fire exclusion and fuel accumulation

(Sackett et al. 1996). Once fuels have been reduced to near

historical levels, the prescribed burning window of opportunity is a bit broader, with good results even when conditions are warmer, such as in the late spring, early fall, or

even the summer (Sackett et al. 1996). Summer prescribed

burns are possible depending on weather conditions, but

ignition is generally limited by the availability of fire

crews, which are often on assignment this time of year.

Both early spring and fall prescribed burns occur during the period of plant dormancy for many species (fig. 5).

One of the main issues with prescribed burns during these

times is that because of the cool conditions, they are often

milder and therefore result in less ecological change than

historical fires.

Figure 5—Prescribed burns during the (a) early growing season (May 3, 2007), and (b) dormant season (October 17, 2007) at Fort Valley

Experimental Forest, Arizona. Understory vegetative growth in the Southwestern United States is influenced by moisture from the summer

monsoon.

A B both photos Walker Chancellor
15

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

Fuel Consumption and Fire Intensity

Because of the seasonal nature of precipitation in the

West, fuels are typically moister for prescribed burns

conducted in spring/early summer or later in the fall, than

for prescribed burns conducted in late summer/early fall

(Kauffman and Martin 1989, Knapp et al. 2005). As a result, such burns often consume less fuel, are less intense,

and are patchier (Kauffman and Martin 1989, Knapp et al.

2005, Monsanto and Agee 2008, Perrakis and Agee 2006).

Kauffman and Martin (1989) reported that total fuel consumption ranged from 15 percent in early spring burns to

92 percent in early fall burns at three mixed-conifer forest

sites in northern California (fig. 6). Duff moisture (as a percentage of dry weight) was 135 percent in early spring and

only 15 percent in early fall.

In the Southwest, conditions at the time of fall prescribed burns are often dry, leading to nearly complete

consumption of the forest floor (Covington and Sackett

1992). However, fuel consumption does not differ predictably with season and is often more of a function of time

since the last rainfall event; conditions often vary substantially within both prescribed burning periods, and consumption is largely controlled by fuel moisture content.

Many prescribed burns in the Western region are conducted in forested areas where fire has been suppressed for

long periods. Because of this, the amount of fuel consumed

by burns in either season may be much greater than the

amount of fuel typically consumed historically (Knapp et

al. 2005). The elevated fuel loading also means that the

difference in total fuel consumption and the resulting fire

intensity among burns in different seasons may be inflated

compared to what was once the case.

Ecological Effects of Burning Season in

Forested Ecosystems

Trees

Differential tree mortality among burning seasons has been

attributed to both phenology (seasonal growth stage) and

variation in fire intensity. In a study of ponderosa pine in

southwestern Colorado, Harrington (1987) reported mortality of trees in different crown scorch categories after

spring (June) and summer (August) prescribed fires conducted during the active growth period, and fall prescribed

fires (October) conducted when the trees were already dormant. By comparing trees that experienced similar fire

intensity, the effect of phenology could be isolated. Trees

with >90 percent of crown scorched were more likely to die

after the spring (54 percent) and summer fires (42 percent)

than after the fall fires (13 percent). Mortality in trees with

crown scorch less than 90 percent was quite low in all seasons. For example, mortality of trees with 67 to 89 percent

of the crown scorched was 12, 11, and 0 percent, for spring,

summer, and fall burns, respectively. When crown scorch

was 66 percent or less, the differences in mortality between

seasons was not statistically significant. Because the goal

of operational prescribed burns is generally to avoid high

levels of scorching of larger trees, any difference in mortality between burning seasons may end up not being biologically meaningful. Indeed, ponderosa pines greater than

12 in diameter, which managers are most likely to want to

retain, had equally low (< 8 percent) mortality rates after

fires in all three seasons (Harrington 1993). Differential

mortality among seasons was only witnessed for small size

classes. Younger trees of shorter stature are more likely to

have high levels of crown scorch, and as the objective of

prescribed burns is often to thin the forest of younger or

suppressed trees, greater mortality of this size class with

early or mid-season burns may be advantageous.

In a study of interior Douglas-fir, Ryan et al. (1988)

noted that overall mortality was nearly the same for spring

and fall prescribed burns (53 percent vs. 47 percent, respectively), although the spring burns were more intense. Fire

damage measures (proportion of cambium killed and crown

scorch) were predicted to contribute much more strongly to

mortality than the burning season.

Several recent prescribed fire studies (Perrakis and

Agee 2006, Sala et al. 2005, Schwilk et al. 2006, Thies et

al. 2005, all covered in the following paragraphs) reported

at least a tendency for higher tree mortality after fall burns.

Most, if not all, of the sites studied had not burned in some
16

GENERAL TECHNICAL REPORT PSW-GTR-224

time, and common to all was greater fuel consumption

in the fall. Although the spring and early summer burns

were conducted during the active growth phase when

loss of living material is expected to be more detrimental,

it appears that when the difference in fuel consumption

between spring and fall burns is substantial (such as after

a period of fire exclusion and fuel buildup), the effect of

fire intensity may overwhelm the effect of phenology.

Perrakis and Agee (2006) reported higher mortality

after fall burns (October) than spring burns (late June) in

mixed-conifer forests of Crater Lake National Park without

a recent history of fire. Fall burns were conducted when

fuels were drier, with burn coverage averaging 76 percent

and fuel consumption averaging 52 percent, as compared

to 37 percent and 18 percent, respectively, for the spring

burns. The authors concluded that the higher mortality was

best explained by the greater intensity of the fall burns,

which may have overwhelmed seasonal vulnerabilities.

Interestingly, an earlier less controlled study of prescribed

burning season nearby showed the opposite result (Swezy

and Agee 1991). These authors reported mortality of large

ponderosa pine after prescribed fires in June, July, and

September to be 38 percent, 32 percent, and 12 percent,

respectively. Although the effect of burning season was

significant, the relative importance of variables showed

fire severity measures (scorch height and ground char)

explained more of the variation in mortality than burning

season. The prescribed fires in this study were conducted

over a period of two decades, with all but one of the lateseason burns occurring in the 1970s and most of the earlyseason burns occurring in the 1980s. Therefore, mortality

results could have been confounded with longer term

climatic patterns. It is also possible that fuel consumption

differences among seasons were not as great as for the fires

studied by Perrakis and Agee (2006).

In a large replicated study of burning season in mixedconifer forests of the Southern Sierra Nevada, Schwilk

et al. (2006) did not find any significant differences in

tree mortality between early season (June) and late season

(September to October) prescribed burns (fig. 7). The June

burns were conducted shortly after trees had initiated

growth (bud break), whereas the September/October burns

were conducted after visual evidence suggested growth

had ceased for the year. The historical fire-return interval in

Figure 6—Average litter and duff consumption at

varying litter and duff moisture levels for burns

in the Sierra Nevada, California, conducted at

different times of the year. Data from Kauffman

and Martin (1989, 1990).
17

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

the study area was approximately 27 years (Schwilk et al.

2006), but as a consequence of fire exclusion, hadn’t

burned for over 125 years, and fuel loading was therefore

very high. Because of higher moisture levels, the June

burns consumed less of the available fuel; however, total

amount of fuel available and consumed was likely far

above historical levels for burns in both seasons. There

was a tendency for higher mortality in the small tree size

classes with the late-season burns (greater fuel consumption) than the early-season burns (less fuel consumption),

although the differences were not statistically significant.

Despite variation in fuel consumption, average crown

scorch height and bole char height did not differ between

seasons. For each tree size category, differences in mortality appeared to be largely a result of local variation in fire

intensity, with little effect of fire season.

In a study conducted in eastern Oregon, ponderosa

pine trees experienced less mortality after spring (June)

burns (11 percent) than after fall (October) burns (32 percent) (Thies et al. 2005). The amount of fuel consumed was

not quantified. However, the fuel at the base of the trees

burned more completely, and a higher proportion of trees

experienced crown scorch with the fall burns than spring

burns. The apparently greater fire intensity with fall burns

appeared to have a stronger impact than effects of phenology, which would have been expected to cause greater

mortality with the spring burns. A tree mortality model

developed using data from this study and burns in northern California did not find burn season to be a predictor

variable, with approximately the same level of delayed

mortality expected for a given level of fire damage,

regardless of the burn timing (Thies et al. 2008).

Other studies include Sala et al. (2005), who found

that physiological performance (net photosynthetic rate,

stomatal conductance, and xylem water potential) and

wood growth of ponderosa pine did not differ between trees

in units burned in the spring or the fall. As is often the case

with prescribed burns in the Western United States, the

spring burns consumed less fuel than the fall burns.

Comparing the outcome of a spring wildfire (May),

a summer wildfire (late June), and a fall prescribed fire

(September) in Arizona, McHugh and Kolb (2003) reported that mortality in all seasons was greatest on trees

most heavily damaged by fire. Total tree mortality averaged 32.4 percent, 13.9 percent, and 18.0 percent in spring,

summer, and fall, respectively. Although the spring wildfire

Figure 7—Mortality of fir (white fir (Abies

concolor) and red fir (Abies magnifica A.

Murr.)) trees in four size classes 2 years after

prescribed burns in the late spring/early summer and in the fall at Sequoia National Park,

California. This large-scale season-of-burning

experiment was initiated in 2001 as part of

the National Fire and Fire Surrogate study.

Although mortality of the 4- to 8-inch and 8- to

16-inch size category trees with burning differed

from background mortality in the unburned control, difference between burning-season treatments was not significant. Data based on

Schwilk et al. (2006).
18

GENERAL TECHNICAL REPORT PSW-GTR-224

occurred prior to bud break, conditions were dry and crown

scorch was also greater than for the other fires (55.3 percent) (McHugh et al. 2003). The summer fire burned during the active growth phase of trees but scorched the least

canopy of the three fires (27.3 percent) (McHugh et al.

2003). Crown scorch for the fall prescribed fire was intermediate, as was the mortality. Total crown damage and bole

char explained much more of the variation in tree mortality

than season of the fire (McHugh and Kolb 2003).

Secondary mortality in many western conifer species is

often attributed to bark beetles. Bark beetle attack probability is usually correlated to degree of tree injury, which

may differ among burning seasons as a result of differences

in fire intensity. The timing of fire may also influence bark

beetle populations directly (Schwilk et al. 2006). Bark

beetles are known to be attracted to volatiles released from

tissues injured by heat (Bradley and Tueller 2001, McHugh

et al. 2003). Bark beetle activity had likely already ceased

for the season by the time of the fall prescribed burning

period. By the time bark beetles become active again the

following spring, volatiles produced by injured tissue may

have already subsided. Early-season burns, on the other

hand, typically coincide with increasing bark beetle flight

activity (Fettig et al. 2004), and there is some concern that

this could lead to a buildup of bark beetle numbers.

Schwilk et al. (2006) did not find any difference

in bark beetle attack probability between June and

September/October prescribed burns on pine species,

but did note an increase in bark beetle attacks on smaller

diameter firs with the earlier burns. Because of the overabundance of small firs in many mixed-conifer forests

following logging and fire exclusion, favoring pines over

firs is a management goal of many prescribed fire projects.

Thus, if causing greater mortality of small firs relative to

small pines is an objective, early-season burns may prove

advantageous.

In a survey of bark beetle populations following fires

in ponderosa pine forests in Arizona, McHugh et al. (2003)

found some differences in attack probabilities among seasons, with a May wildfire leading to greater probability of

attack (41 percent), compared to a June wildfire (19 percent), or a September prescribed burn (11 percent). The

May wildfire also was the most intense, causing the most

crown scorch, and overall attack probability was associated

with degree of fire-caused damage. However, attack probability was somewhat greater for the June fire than the

September prescribed burn although crown scorch was less.

This suggests that the timing of fire relative to periods of

bark beetle activity may play a role. Still, studies to date

all point to degree of crown damage being the overriding

contributing factor to bark beetle attack, regardless of

season of burn.

Understory Vegetation

Steele and Beaufait (1969) found no important differences in the cover of understory vegetation between

areas treated with either early- or late-season broadcast

burning treatments in Montana. In southwestern ponderosa

pine systems, fall prescribed burns often lead to a greater

abundance of understory vegetation such as cool-season

perennial grasses. Sackett and Haase (1998) suggested

that burning during the natural fire season (May through

early July) might lead to an even greater increase in grass

production, because grasses that are growing and green are

less readily consumed by such fires. In addition, seed heads

are possibly less likely to be consumed with late spring/

early summer burns than with fall burns (Sackett and Haase

1998). Certain species that grow later in the year, such as

the warm-season grass mountain muhly (Muhlenbergia

montana (Nutt.) Hitchc.) appear to be negatively affected

by repeated fall burns (Laughlin et al. 2008).

Kauffman and Martin (1990) reported much higher

shrub mortality after early fall burns (high fuel consumption), than after spring burns (low fuel consumption).

Overall, the greater the consumption of fuel, the greater

mortality of shrubs, regardless of burning season. Variability in mortality was also seen among sites within a burn

season treatment, with lesser mortality at sites that contained the least fuel, and therefore experienced lower total

heat flux upon burning. These authors hypothesized that
19

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

shrub phenology at the time of fire may have also played a

role, albeit a lesser one. At one site, mortality of black oak

was 31 percent following early spring burns conducted

prior to bud break and initiation of growth, and 55 percent

following late spring burns conducted during the period of

rapid growth following bud break, although fuel consumption with these two burn treatments was nearly identical

(77 percent for early spring vs. 79 percent for late spring

burns, respectively). Differences in plant carbohydrate

storage among seasons may have been one mechanism

for this observed difference (Kauffman and Martin 1990).

However, variation in mortality between seasons could also

be attributed to factors other than phenology. For example,

soil moisture at the time of early spring burns was nearly

double that of the late spring burns (Kauffman and Martin

1989, 1990), which may have also reduced the heat flux

into the soil.

For fire-following species, differential response among

burning seasons is also sometimes evident in the seed

germination phase. Enough heat is required to scarify the

seed, but not so much that the seeds are killed (Knapp et al.

2007, Weatherspoon 1988). Depth of lethal heating, which

is affected by both the amount of fuel consumed and the

moisture content of the soil, may determine how many

seeds are available to germinate. Kauffman and Martin

(1991) found that wet heat, simulating a heat pulse under

moist soil conditions, was more effective for scarifying

seeds of shrubs than dry heat, simulating fire in the fall

when soils were dry. The dry heat actually resulted in

higher seed mortality. In another study in an area with

low fuel loading (10 years after a fire), Harrod and Halpern

(2009) found that fall burns stimulated germination of

long-sepaled globe mallow (Iliamna longisepala (Torr.

Wiggins)), while spring burns did not. It is possible that

the soil heating generated by spring burns was, in this case,

insufficient.

Knapp et al. (2007) reported that understory vegetation

in a mixed-conifer forest in the Sierra Nevada of California

was resilient to prescribed fire conducted in either late

spring/early summer (June) when plants were in the midst

of active growth, or in the fall (September/ October) when

most plants were nearly to fully dormant. Several years

after treatment, total plant cover and species richness in the

spring/early summer- and fall-burned plots did not differ

significantly from each other or from an unburned control.

However, there was a difference in the rate of vegetation

recovery between burn season treatments. In the season

immediately following the burns, cover was initially reduced relative to the control in the fall burn treatment, but

not the spring/early summer burn treatment. Furthermore,

certain species, particularly ones most common under the

forest canopy where surface fuel loading is expected to be

the highest, such as whiteveined wintergreen (Pyrola picta

Sm.), were reduced in frequency by late-season burns but

not early-season burns. Because the late-season burns were

conducted when the fuels and soils were drier, the greater

fuel consumption and heat penetration into the soil (see

“Soils” section) may have killed more of the underground

structures than the late spring/early summer burns. Lateseason burns also covered a larger proportion of the forest floor, leaving fewer undisturbed patches. Vegetation

change was associated with variation in fire severity, and

the authors concluded that effects on vegetation suggested

a greater dependency on amount of fuel consumed and fire

intensity than on plant phenology.

In a longer term study of understory vegetation response to burning season in a ponderosa pine forest of

eastern Oregon, Kerns et al. (2006) reported no significant

difference in native perennial forb cover 5 years after earlyseason (June) and late-season (September/October) prescribed burns. The June burns occurred during the active

growth phase of many understory plant species, whereas

the September/October burns occurred when most species

were dormant. Harrod and Halpern (2009) also found few

effects of either spring (May) or fall (October) prescribed

burns on mature individuals of two native herbaceous

perennial plant species. In the Kerns et al. (2006) study,

exotic species, which often thrive with disturbance, were

more frequent following the higher severity (as evidenced

by greater bole char and higher tree mortality) late-season
20

GENERAL TECHNICAL REPORT PSW-GTR-224

burning treatments. Exotic species were also concentrated

in patches within burns where local severity was the

highest. This study is another example of plants responding more strongly to fire intensity and degree of environmental change than the plant phenology at the time of the

fire. A similar trend, with greater numbers of exotic species

in plots that burned at higher severity in the fall was noted

by Knapp et al. (2007); however, in this latter study, the

difference was too small to be statistically significant.

By timing prescribed burns for when plants are most

vulnerable, fire can be used to control vegetation or target

certain species. Harrington (1985) reported that a Gambel

oak (Quercus gambelii Nutt.) understory of a ponderosa

pine forest resprouted vigorously following single prescribed burns conducted in the spring (June), summer

(August), or fall (October). The spring burns occurred

3 to 4 weeks after bud break and leaf emergence, the

summer burns occurred while vegetation was still actively

growing, and the fall burns occurred after plants had gone

dormant and leaves had fallen. A second summer fire 2

years later significantly reduced the frequency of resprouting stems, whereas spring and fall fires did not. However,

differences in sprout number among treatments were relatively small. The effect was attributed to reduced root

carbohydrate reserves in the summer following a second

flush of growth, which suppressed the energy available for

resprouting following fire (Harrington 1989).

Several studies have been conducted to investigate

whether burning in different seasons might be used to control bear clover (Chamaebatia foliolosa Benth.), a vigorous highly flammable shrub with rhizomatous roots that

can compete strongly with conifer seedlings. Fires in May

(prior to the growing season) and October (after the growing season) stimulated growth of C. foliolosa relative to

the control, whereas prescribed burn in July (mid growing

season) resulted in growth comparable to the control after

2 years (Rundel et al. 1981). Weatherspoon et al. (1991)

reported that a single prescribed burn in any season (May

through October) was ineffective for reducing the cover

of this plant, but a second treatment during the growing

season, where all tops were removed, simulating the effect

of a followup prescribed burn, did slow regrowth. Studies

on chamise (Adenostoma fasciculatum Hook. & Arn.) also

have shown top removal during the growing season to slow

regrowth compared to top removal during the dormant season (Jones and Laude 1960). Results suggest that carbohydrate reserves at the time of treatment may play a role in

regrowth.

Burning in different seasons has been attempted as a

means of controlling shrubs with seed banks stimulated

to germinate by fire (such as Ceanothus sp. or Manzanita

(Arctostaphylos sp.)). Hotter burns that consumed the

entire duff layer under dry soil conditions in the fall killed

more seeds by pushing critical temperatures deeper into

the soil than burns in the spring that consumed less fuel

(Weatherspoon 1988). However, so many seeds were found

in the soil that sufficient seeds remained to regenerate a

vigorous shrub layer no matter the burn season

(Weatherspoon 1988).

Soils

Soil heating during the process of combustion can cause

biological and physical changes such as root mortality

or increased water repellency. The magnitude of change

depends at least partially on three factors that may differ

with burning season: amount of fuel consumed, duration

of combustion (residence time), and soil moisture at the

time of burning.

Fuel moisture largely dictates how much organic

material is consumed, and therefore the residence time

of combustion. Likewise, the extent to which the heat

penetrates into the soil is determined by soil moisture

(Campbell et al. 1995). Water has a high specific heat and

therefore substantial energy is required to drive off the

moisture before the temperature of that soil will exceed

212 o

F, the boiling point of water. Because of this, moist

soils are much less likely to heat up than dry soils. Soils

are largely protected from excessive heating, even under

high fuel loading conditions if they contain sufficient

moisture (Busse et al. 2005, Frandsen and Ryan 1986,

Hartford and Frandsen 1992). Plant roots are killed starting

at soil temperatures between 118 and 129 o

F, microbes are
21

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

killed between 122 and 250 oF, and buried seeds have been

reported to die at temperatures between 158 and 194 oF

(Neary et al. 1999). Busse et al. (2005) found that the temperature at 1-inch depth in the soil below a laboratory burn

that consumed a very high load of masticated wood chips

(69.9 tons/ac) reached a maximum of 595 oF in dry soils

and only 241 oF in moist soils.

Effects on soil physical properties and soil biota

largely mirror the intensity and severity of the fire (Neary

et al. 1999). In a study in mixed-conifer forest of the Southern Sierra, California, Hamman et al. (2008) reported soil

temperature, moisture and pH, plus mineral soil carbon

levels and microbial activity following late spring/early

summer (June) prescribed burns to be generally intermediate between fall (September/October) prescribed burns and

unburned controls. A similar result was reported from ponderosa pine forests in eastern Oregon, with October prescribed burns decreasing soil carbon and nitrogen, whereas

June burns had little impact (Hatten et al. 2008). The

magnitude of effects for both the Hamman et al. (2008)

and Hatten et al. (2008) studies were in line with the

greater fuel consumption and intensity of the late-season

burns. In the same study plots as Hatten et al. (2008), Smith

et al. (2004) found that the October prescribed burns significantly reduced fine root biomass to a depth of 4 in and

depressed the number of ectomycorrhizal species, relative

to units burned in June. Fine root biomass and ectomycorrhizal species richness following the June burns did not

differ from the unburned control. Soil moisture values were

not provided, but given the rainfall patterns, it was likely

considerably higher at the time of the June burns. Other

studies corroborate findings of a greater loss in soil

microbes following burns when soils were dry than when

soils were moist (Klopatek et al. 1988, 1990), corresponding to the amount of soil heating. Filip and Yang-Erve

(1997) reported a reduction in root disease causing fungi

following fall burns but not spring burns; however, soil

moisture and fuel consumption were not reported.

In addition to changes within the soil, other variables

that frequently differ with burning season may influence

soils indirectly through erosion. Such variables include the

percentage of the soil surface burned, and the depth of

burn (how much of the duff layer is removed). Burns when

soils and the fuels in contact with those soils are moist tend

to be patchier (Knapp and Keeley 2006). These unburned

patches may act as refugia from which fire-sensitive

organisms such as soil ectomycorrhizae can recolonize

burned areas (Smith et al. 2004), or act as barriers to soil

erosion (Knapp et al. 2005). Johansen et al. (2001) reported

an exponential increase in the amount of erosion once the

percentage of the forest floor burned exceeded 60 to 70

percent, presumably because as the percentage increases,

burned patches coalesce into larger and larger areas, leaving fewer unburned patches at a scale necessary to capture

sediment. Under the high fuel loading and high fuel continuity in landscapes common today, many prescribed

burns cover a greater percentage of the landscape than this,

particularly ones conducted when fuel conditions are dry.

Whether changes to soils as a result of fire are beneficial or detrimental will depend on the burn objectives.

Burns at times of the year when soils (and fuels) are still

moist may limit the amount of soil heating and leave a

greater amount of duff unconsumed, which could reduce

the threat of erosion. However, burns at drier times of the

year may be necessary if bare mineral soil exposure is

desired to produce an adequate seedbed for species that

don’t germinate well through a layer of organic material,

or if the objective is to heat scarify deeply buried seeds of

fire-following species.

Wildlife

Wildlife populations may be affected by fire either directly

by heat and flames, or indirectly through modification of

the habitat. In environments where fire was historically

common, there is little evidence that fires falling within the

range of historical intensities cause much direct mortality

of wildlife (Lyon et al. 2000b, Russell et al. 1999). Most

animals have presumably developed behavioral adaptations for escaping fire that enable population persistence,

and many, in fact, benefit from the habitat modifications

resulting from fire.
22

GENERAL TECHNICAL REPORT PSW-GTR-224

In the Western United States, most species have already

successfully produced young by peak fire season in late

summer to early fall. There has been some concern that

prescribed fires ignited outside of the season when historical fires were common might do harm to wildlife populations, especially for species with poor dispersal or species

that raise offspring in locations that are most likely to burn.

For example, small mammal young may be more vulnerable to early-season fire, because of lack of mobility prior

to maturity (Lyon et al. 2000a). Many of these species have

high reproductive rates, however, and recovery is likely

rapid.

Ground-nesting birds could be killed prior to fledging

(Reinking 2005) and forest floor arthropods in the egg or

larval stages may be more vulnerable to loss (Niwa and

Peck 2002). Amphibians are also likely to be more active

with the moister conditions under which prescribed fires

are typically conducted (Pilliod et al. 2003). On the other

hand, amphibians tend to live in the moister microsites

that are least likely to burn in prescribed fires, especially

in the early season (Lyon et al. 2000a). In the Southwestern

United States, the peak historical fire activity occurred

earlier, during the spring and early summer, when effects

on wildlife might be more severe. In this case, the impact

of prescribed fires in the spring or fall would be expected

to be less than those in the main historical fire season.

Much of the information about effects of season of

prescribed fire on wildlife in the Western United States is

anecdotal or has lacked a direct comparison among seasons. For example, many studies compared early-season

fire with no fire, or late-season fire with no fire. What has

been written generally has found very little influence of

fire season on populations. Wildlife may be affected by

fire both through direct mortality or habitat alteration

(Lyon et al. 2000b), but the latter appears to play a larger

role. In some cases, the magnitude of change in populations or communities has been associated with measures

of fire severity, which may differ with burning season. For

example, dark-eyed juncos (Junco hyemalis) often choose

nest sites in unburned patches within prescribed fire units

(Sperry et al. 2008), and burns in early season when fuels

are moist are more likely to create such unburned islands.

One of the most rigorous evaluations of burning season to date reported similar effects of early (June)- and

late (September/October)-season prescribed burns on

small mammal populations in a mixed-conifer forest of

the Southern Sierra Nevada (Monroe and Converse 2006).

Although the June burns occurred during the small mammal breeding season, the burns consumed less fuel and

were therefore less intense than later burns under dryer

conditions. June burns were also patchier (Knapp and

Keeley 2006), leaving more potential refuges and habitat

such as coarse woody debris where animals could have

escaped fire. Most of the variation in population numbers

in the Monroe and Converse (2006) study was attributed

to year-to-year differences in food availability tracking the

yearly seed production cycles of the overstory trees. This

further suggests that small mammals respond more strongly

to habitat conditions, including those created by the fires,

than to the burning season.

As is the case with small mammals, the effect of early

season prescribed fire on forest floor arthropods might also

be expected to differ with the life cycle of the organisms

because of seasonal vulnerabilities. However, using the

same plots as the Monroe and Converse (2006) study,

Ferrenberg et al. (2006) reported no significant differences

in forest arthropod community structure between the two

burning season treatments. Fire influenced the arthropod

community, reducing abundance but increasing diversity,

but changes appeared to be mediated by habitat alteration (amount of litter and duff, coarse woody debris, vegetation), and these habitat variables differed much more

strongly between the control and burn units than between

the June and September/October burning treatments.

Changes in the June burn treatment were generally intermediate between the control and September/October burn

treatments.

Adult birds are highly mobile and easily escape prescribed burns. Early-season burns may cause some direct

mortality of young, particularly for species nesting on

the ground, but the ultimate impact on bird populations

requires a longer term view. When nests are lost, many species will renest (Reinking 2005). In addition, like many
23

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

wildlife species, bird populations are capable of responding rapidly, with population size limited by food availability and shaped by habitat changes.

Unfortunately, experimental design flaws limit the

inference of many studies of the response of birds to fire

(Finch et al. 1997). Published literature comparing the effects of prescribed burns in different seasons on birds are

not available for the Western United States. Preliminary

data from the Sequoia National Park study on burning season suggest that effects one to three seasons after the burns

were minimal.3 Population sizes of the eight most common

species observed with point counts and bark foraging surveys did not differ significantly between burning season

treatments. Too few nests could be located to investigate

direct mortality from the June burns.

Besides direct mortality, another possible short-term

impact of spring or early-summer prescribed burns is a

temporary drop in food availability or cover because

understory vegetation in these systems may not resprout

until the following year. It is possible that lack of food

could reduce reproductive success. The longer term responses of many bird species are thought to be due primarily to structural changes of vegetation or changes

to food resources, as affected by fire severity (Huff and

Smith 2000, Kirkpatrick et al. 2006). For example, foliage

gleaners typically decline in abundance when more of the

tree crowns are lost to scorch, and woodpeckers increase in

abundance when fire-damaged trees are attacked by bark

beetles, an important food source (Huff and Smith 2000).

Variation in outcomes among prescribed burns early or late

in the season would therefore mainly be expected if crown

scorch or mortality of vegetation differed.

Ecological Effects of Burning Season in

Chaparral and Grasslands

Chaparral

Extensive chaparral shrublands are found in nondesert

areas of central and southern California and historically

3

Farris, K.; Zack, S. Unpublished data.

burned over a range of intervals, from every few decades

in montane sites with more frequent lightning, to 100 years

or more in areas closer to the coast. Most of the acres were

burned in late summer through the fall, often in high intensity stand-replacing events (Keeley and Fotheringham

2001) (fig. 8). Because of frequent human-caused ignitions

and seasonal hot and dry winds, the fire regime remains

similar today, despite fire-suppression efforts. Plant species

have evolved means of persisting under such burning conditions, from resprouting of lignotubers, to seeds requiring

substantial heating or exposure to chemicals found in char

for germination (Kauffman 1990, Keeley 1987, Odion

2000).

Prescribed burns are sometimes used to reduce fire

hazard in chaparral, but such burns are controversial

(Keeley 2002, Keeley and Fotheringham 2001, Parker

1987a). To avoid burning during times when the vegetation is most volatile and conditions are conducive to rapid

fire spread, many prescribed burns are conducted in the

winter or spring, outside of the historical fire season. Live

fuel moisture is typically higher and soils considerably

wetter at such times of the year, than would have been the

case for historical fires (Beyers and Wakeman 2000). As

a result, prescribed burns are usually considerably less

intense than the wildfires that this vegetation evolved

with. Observations suggest that vegetation response to

such prescribed burns often differs from response to natural

wildfires, with reduced germination of certain herbs and

potentially altered species composition (Le Fer and Parker

2005; Parker 1987a, 1987b). For example, Ceanothus L.

seeds require heat for germination (Keeley 1987), and

abundance of seedlings has been shown to be greater

following fall prescribed burns than spring burns (Biswell

et al. 1952, Gibbens and Schultz 1963).

Parker (1987a) and Le Fer and Parker (2005) attributed

the reduced germination of some obligate seeding chaparral species following spring prescribed burns to higher

seed mortality upon heating. It was thought that seeds

are particularly vulnerable when soils are moist and seeds

full of water, compared to when seeds are dry. Interestingly,

species producing hard seed with dormancy (such as
24

GENERAL TECHNICAL REPORT PSW-GTR-224

Ceanothus spp.) that do not imbibe water until dormancy

is broken, were not differentially affected by heating under

moist or dry conditions (Le Fer and Parker 2005). Given

that heat penetration is limited when soils are moist (Busse

et al. 2005, Frandsen and Ryan 1986), it is also possible

that the soil heating under prescribed burning conditions

typical for this vegetation type may be insufficient to

scarify seeds of hard-seeded species (Beyers and Wakeman

2000). However, Beyers and Wakeman (2000) reported no

decline in numbers of shrub seedlings or herbaceous species germinating from seed following late spring prescribed

burns (May) as compared to fall (October) wildfire. Although this result might seem contrary to the work of

Parker (1987b), the late spring prescribed burns in the

Beyers and Wakeman (2000) report were likely of higher

intensity, closer to the fire intensity expected with historical wildfires. Soil moisture was likely also less.

Out-of-season burns have the potential to reduce the

length of the growing season, and this could also potentially influence seedling survival.4

 Chaparral shrubs are

typically actively growing throughout the winter rainy

season—a seedling might have 6 months to grow after

germination following a typical fall wildfire, whereas a

4

Keeley, J.E. 2008. Personal communication.

winter or spring burn would considerably shorten the time

to establish prior to the summer dry period. With less time

to grow and put down deep roots, smaller seedlings may be

less likely to survive.

Reported responses of mature shrubs to burning season

have been variable. Shoot growth for resprouting chamise

(Adenostoma fasciculatum Hook. & Arn.) was not found

to be affected by prescribed burn season (Radosevich and

Conard 1980). Beyers and Wakeman (2000) also did not

note differences in mortality of resprouting shrubs with

spring or fall burns. Conversely, Parker (1987) found that

more than 70 percent of chamise plants had died one or

two years after spring burns, while nearly all plants successfully resprouted after early fall burns. Higher mortality

with spring burns was thought to have been due to the

timing of fire in relation to periods during which carbohydrate storage is lowest (Jones and Laude 1960).

The bottom line is that the potential for shifts in the

plant community exists when the heat generated by prescribed burning is dissimilar to what would have been experienced with the fire regime that species evolved with.

Seeds of species requiring heat to germinate are dependent

on receiving enough to break dormancy, but not so much

that they are killed. Seeds of species requiring chemical

(charate) cues rather than heat to germinate should not be

A B

Figure 8—Chaparral vegetation of the Western United States typically burns in high-intensity stand-replacing fires, and many plant species

possess adaptations to persist with such a fire regime. Intensity of prescribed burns is often less than that of wildfires, which could affect the

abundance of herbs and shrubs with seeds that are stimulated to germinate by heat. Recovery of chaparral and herbaceous species after the

McNally Fire, southern Sierra Nevada, California, (a) March 2003, 7 months after the fire, and (b) late May 2003, 9 months after the fire. both photos Eric Knapp
25

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

as strongly affected by fire season, unless they are killed

by excess heat. Excess heat is likely to be less in the winter or spring, when soils are moist. Thus, winter or spring

burning might be expected to favor species with charatestimulated seeds, whereas late summer or early fall burning

may create opportunities for a greater mix of species with

different strategies. Biswell et al. (1952) suggested that

some fall management burns, during the natural fire season,

may be necessary to perpetuate Ceanothus, the seeds of

which require heat to germinate.

Western Grasslands

Many western grasslands are highly altered as a result of

nonnative species invasion. Rather than fuel reduction,

the objective of prescribed burning is frequently to reduce the cover of nonnative species so that more desirable

native species may flourish. Such burns are usually timed

for periods where the nonnative species targeted may be

more vulnerable to fire than the native species (DiTomaso

et al. 2006, Meyer and Schiffman 1999, Pollak and Kan

1998). Prescribed burns are likely to be most effective at

reducing a target species if the seeds of that species are

still immature and on the plant, whereas seeds of desirable

species have dispersed to the ground where they may more

readily escape the heat of fire (DiTomaso et al. 2006). For

example, early summer prescribed burns have been effective for controlling yellow star-thistle (Centaurea

solstitialis L.) (DiTomaso et al. 1999)—burns occurred

when this late-flowering annual still contained immature

seeds, but much of the associated vegetation had senesced.

Controlling target herbaceous species with fire is likely to

be more effective in grasslands than many other vegetation

types found in the West, because of the relatively high importance of annuals in this vegetation type. Herbaceous

perennial species that emerge from underground structures

are typically more difficult to kill with fire.

Parsons and Stohlgren (1989) followed vegetation

in grasslands dominated by nonnative species that had

been burned one, two, and three times in successive years

in the spring (mid June, when grass had dried enough to

burn, but prior to the period when such grasslands would

have normally burned historically), and in the fall (late

October or early November, at the very end of the historical fire season). Although fire in both seasons reduced

the number of nonnative grass species and increased the

number of forb species, fire in the fall favored nonnative

forbs, whereas fire in the spring favored native and nonnative forbs equally. Meyer and Schiffman (1999) compared

late spring (June), fall (September), and winter (February)

burns, and reported that late spring fires suppressed non

native annual grasses more so than fall burns, presumably

because grass seeds were not completely mature at the time

of the late spring burns and therefore more vulnerable to

being killed by fire. Winter burns were less intense and

much less effective at altering nonnative grass cover than

either spring or fall burns. Therefore, both phenology and

intensity differences among burning seasons appeared to

have played a role in how grassland vegetation was

affected.

Owing to the presence of nonnative species, the

amount of fuel consumed and the nature of the fire may

differ from historical fires in some cases. However, because

grassland fuels are fine and dry quickly, the difference in

moisture and therefore consumption and aboveground fire

intensity between different burning seasons may not often

be as substantial as in forested ecosystems. Thus, with the

confounding effect of fire intensity lessened, differences

among seasons may more readily be attributed to timing

of the fire in relation to plant phenology.

Much more has been written about ecological effects

of burning season in grasslands from the Great Plains,

which may apply as well. This information is contained

in chapter 4—the Central region.
26

GENERAL TECHNICAL REPORT PSW-GTR-224

growth or during the breeding season. However, all else is

rarely equal. In many areas of the Western United States,

fall prescribed burns are generally conducted when fuels

and soils are drier, more fuel is consumed, and resulting

fire intensity is greater than at the time of spring or early

summer burns. Thus phenology or life history stage and

fire intensity can be seriously confounded. When the difference in fuel consumption between burns in different

seasons is substantial, response of many ecological variables appears to be influenced more by fire-intensity differences than by phenology or life history stage at the time

of the fire. When differences in fuel consumption between

fires in varying seasons are small or nonexistent, the influence of phenology or life history stage may become

Implications for Managers

The published literature on season of burning in western

ecosystems indicates that most species are quite resilient

to fire in any season. The majority of plants in forested

vegetation types here are perennial; loss of one season’s

growing structures in long-lived or readily resprouting

herbaceous species appears to have limited effects over

the long term. In wildlife studies, the large amount of yearto-year variability in population sizes caused by non-fire

factors makes detecting seasonal effects particularly

difficult.

All else being equal (fuel consumption, fire intensity,

etc.), evidence suggests that certain organisms might be

somewhat more affected by burns during times of peak

Key Points–Western region

• The effect of prescribed burning season appears to be relatively minor for many of the species that have

been studied.

• Although stage of plant growth (phenology) at the time of prescribed fire may have some influence on the

community trajectory in forested vegetation types, it appears that the intensity and resulting severity of the

fire often has a greater impact. This is likely to be especially the case in forests that contain heavy surface

fuel loads, where fuel moisture differences among seasons can lead to substantial differences in

consumption.

• In chaparral vegetation, prescribed burns conducted at times of the year with higher soil and fuel moistures

are often considerably less intense and may not stimulate the germination and growth of some species that

are adapted to the historical regime of high-severity fire.

• In predicting outcomes of prescribed burning, it may be useful to compare the prescribed fire intensity and

severity to historical intensity and severity. Burning prescriptions for producing historical or near-historical

intensity and severity could then be developed.

• Until heavy fuels are reduced to historical levels, out-of-season burns that consume less fuel may be useful

for reintroducing fire without causing severe effects.

• A single prescribed burn outside of the historical fire season appears unlikely to have major detrimental

impacts. However, the effect of multiple sequential out-of-season burns remains poorly understood.

• Variation in the timing of prescribed burns will reduce chances of selecting for certain species, thereby

helping to maintain biodiversity.
27

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

more apparent. Another factor that needs to be considered

is the fire intensity in relation to likely historical intensity.

Most prescribed fire studies in western forest ecosystems

have been conducted in areas where fire has long been suppressed and surface fuel loading is uncharacteristically

high. Therefore, prescribed burns in many cases consume

more fuel than wildfires burning every 10 to 15 years once

did. As a result, fire intensity and resulting severity may be

somewhat unnatural. In addition, when the total amount of

fuel consumed is large, the magnitude of potential differences in fuel consumption among seasons as a result of

fuel moisture variation, is also substantial.

If fire effects are driven by differences in intensity

among seasons, burning when fuels are moister may be

one means of limiting consumption and producing fire

effects more similar to those found historically. Higher fuel

moisture is more common in the spring or early summer.

Limiting consumption may be especially advantageous

under conditions of unnaturally high fuel loading. Once

fuels have been reduced to closer to historical levels, burning at times of the year with higher fuel moisture may lead

to less fuel consumed than was historically the norm (fig.

9a). In this case, prescribed burning may result in less ecological change than desired. Also, once fuels are reduced,

the difference in consumption between seasons will likely

not be as high, and the effect of phenology or life history

stage may become more apparent.

In contrast to forested ecosystems that historically experienced frequent low- to moderate-intensity fire, vegetation types where high-severity stand-replacing fire was the

historical norm (chaparral shrublands, for example) may

require hotter prescribed burns than is currently common.

Prescribed burns conducted under benign weather conditions of the late fall, winter, or spring likely consume less

fuel and are less intense than historical fires were (fig. 9b).

In addition, soils at the time of many of these burns are

generally moist, and heat penetration into moist soils

could possibly be insufficient to trigger germination of

heat-stimulated seeds of certain hard-seeded fire-following

species.

The take-home message is that early-season burns may

be a valuable tool for more gradually reducing high fuel

loads, especially for the first restoration burn(s) after a

period of fire exclusion. Once fuels are reduced to historical levels, early-season burns might then be followed by

late-season or a mix of late- and early-season burns. To

mimic the historical highly variable fire regime, timing of

prescribed burns should ideally also be variable. Shifting

the fire regime to entirely spring/early summer growing

season prescribed burning when the historical regime consisted of predominantly late summer/early fall dormant

season fire (much of the Western United States), or shifting

the fire regime to entirely fall dormant-season burning,

when the historical regime consisted of late spring/early

summer growing-season fire (as in areas of the Southwestern United States influenced by a monsoonal climate), may

eventually lead to demonstratable ecological change, even

if such change is not apparent today. Areas of the Western

United States have generally seen at most three cycles of

prescribed burning, and data from other parts of the United

States with a longer history of prescribed fire show that

numerous burn cycles may be required to dramatically shift

community composition. Some of the heterogeneity in the

prescribed fire regime will be produced from year-to-year

variation in climate alone. A prescribed burn in one year

may have entirely different effects than a fire on the same

date in another year, as climatic differences can influence

the phenology or life history stage.
28

GENERAL TECHNICAL REPORT PSW-GTR-224

Figure 9—Conceptual diagram showing expected fire effects under typical historical and contemporary fuel loading (dead and live) conditions with prescribed burning in different seasons. Fire effects could include variables such as amount of crown loss, percentage of

ground surface burned, or depth of soil heating. (a) In a western coniferous forest where fire has been excluded and fuel loading is unnaturally high, spring burns under moist conditions may consume an amount of fuel and produce fire effects closer to the historical norm

than a late summer (or early fall) burn under drier conditions. Once fuels are reduced/restored to historical levels, it is possible that the

opposite may occur, with late summer burns resulting in fuel consumption closer to the historical norm and early-season burns resulting

in fuel consumption (and fire effects) outside of the historical range of variability. (b) In western chaparral ecosystems, spring burns under

moist conditions might be expected to lead to fire effects below the historical range under both historical and contemporary fuel loading

conditions. This ecosystem historically most commonly burned in high-severity stand-replacing fires in the late summer or fall, and fuel

loading is today, in many areas, not greatly different from historical levels.
29

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

Chapter 4: Central Region

The dominant grasses in all of the four grassland types

are generally perennial with annuals becoming more

abundant after disturbance (table 3). Grass composition

varies within the three main grassland types. Tallgrass

prairies are mainly composed of warm-season grasses (C4

photosynthetic pathway), whereas mixed and shortgrass

prairies are composed of varying quantities of cool-season

(C3 photosynthetic pathway) and warm-season grasses.

Many perennial grasses have underground rhizomes or

growing points at or below the soil surface, protecting them

from fire, drought, and grazing. Forb abundance is dynamic

with patches affected by disturbances such as fire and grazing. Hardwood pockets and scattered oak savannas are also

found, especially in areas with higher precipitation, along

riparian corridors, and where fire has been excluded for

long periods. Another vegetation type covered in this

chapter is the mesquite savannah found from southeastern

Arizona through western Texas (fig. 10b). This system

contains more shrubs, which have invaded an arid grassland composed of a mix of cool- and warm-season species.

The growth period for many plants here is earlier than in

grasslands farther north.

Historically, fire played an integral role in maintaining North American grasslands by stimulating native

grass production and impeding succession to woody vegetation (Axelrod 1985; Collins and Wallace 1990; Hulbert

1969, 1988). Unless accumulated litter is periodically removed by fire, grazing, or haying, productivity and plant

diversity decline (Anderson 1990, Kansas Natural Heritage

Inventory 2007).

Historical Fire Regime

The central grasslands have developed and flourished in

an environment with recurrent fire from lightning ignitions and Native American activity (Abrams 1992, Axelrod

1985, Baker et al. 1996, Komarek 1967). Without physical

evidence such as fire scars, understanding how often grasslands burned historically is mostly anecdotal. Rate of fuel

The Central region encompasses the major grasslands of

the United States from the Rocky Mountains east to the

Great Lakes, and from eastern Montana, North Dakota, and

western Minnesota in the north, to the Mexican border in

Texas in the south (fig. 10). Over much of the area, native

grasslands have been replaced by agriculture, degraded by

overgrazing, or lost through hardwood encroachment and

now cover only a small portion of their former range. Many

are so fragmented that the fire regime has been seriously

disrupted. Reduction in fine fuels from grazing as well as

fire exclusion has limited the role of fire in the maintenance of grasslands.

Climate, Vegetation, and Fire

The Central region vegetation is composed of four major

grassland types: shortgrass prairie, northern mixed-grass

prairie, tallgrass prairie, and southern mixed-grass prairie,

with vegetation influenced by climate, topography, and

soil type. Precipitation is light to moderate and generally

ranges from 10 to 20 inches in the north and west to 20 to

40 inches in the south and east (Bailey 1980). Airmasses

from the Gulf and the Pacific trigger precipitation, but the

Pacific airmass is usually dry after passing over several

mountain ranges; thus the temperate steppe and substropical steppe grasslands directly east of the Rocky

Mountains receive less precipitation and are of shorter

stature (northern mixed-grass and shortgrass prairie,

respectively) (fig. 10a). The Gulf airmass originates in the

Gulf of Mexico, producing higher humidity and greater

precipitation, limiting the periods of drought in the mixed

and tallgrass prairie (Anderson 1990) (fig. 10c). Stature of

the grassland vegetation follows this moisture gradient,

with the shortgrass prairie transitioning to the southern

mixed-grass prairie and finally into the tallgrass prairie

from west to east. Gradients also exist from north to south,

with the polar airmass exerting a greater influence to the

north. This can result in more continuous snow cover,

which reduces periods of flammability.
30

GENERAL TECHNICAL REPORT PSW-GTR-224

Figure 10—Climographs (monthly average temperature and precipitation) and the average time of the year of the peak

historical and prescribed fire seasons from three representative locations within the Central region: (a) Medora, North

Dakota; (b) Big Bend National Park, Texas; and (c) Wichita, Kansas. Note that because the timing of anthropogenic fire

is poorly understood, the historical fire season reflects mainly lightning-ignited fires. Historical anthropogenic fires

likely extended farther into the dormant season.
31

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

accumulation in some grasslands is sufficient to carry fire

every year, but in others at least 1 year between fires may

be necessary for dead fuels to build up (Bragg 1982),

particularly if the grassland is grazed.

Historical timing of fire in the central grasslands was

dictated by phenology of the vegetation, sources of ignition, and other weather events such as precipitation and

wind. Grassland vegetation typically starts growing in

spring (March/April), senescing in late summer and fall, or

earlier if summer moisture is not available. In the dormant

season (fall and winter through early to mid spring), the

grassland consists of a higher dry component as thatch.

This thatch is more flammable than actively growing

vegetation, at least at times without recent precipitation.

In northern climates, snow cover limits drying of thatch,

and thus the duration of the fire season. In the more mesic

grasslands, fuels may also be too moist to burn during the

summer growing season, especially during wet years,

because of the low ratio of dead to live fuels (Engle and

Bidwell 2001). However, Bragg (1982) reported that

Table 3—Cool-season (C3

 photosynthetic pathway) and warm-season (C4 photosynthetic pathway) grasses and

forbs commonly found in tallgrass prairies (Howe 1994b)a

Cool-season grasses Warm-season grasses Cool-season forbs Warm-season forbs

Texas wintergrass Buffalograss Tall goldenrod Richardson’s alumroot

(Nassella leucotricha (Buchloe dactyloides (Solidago altissima L.) (Heuchera richardsonii

[Trin. & Rupr.] Pohl) (Nutt.) J.T. Columbus) R. Br.)

Scribner panicum Indiangrass Spotted trumpetweed Candle anemone

(Dichanthelium (Sorghastrum nutans Eupatoriadelphus (Anemone cylindrica

oligosanthes (Schult.) (L.) Nash) maculatus (L.) King & A. Gray)

Gould) H. Rob. var. maculatus)

Porcupine grass Switchgrass Flowering spurge Old man’s whiskers

(Hesperostipa spartea (Panicum virgatum L.) (Euphorbia corollata L.) (Geum triflorum Pursh)

(Trin.) Barkworth)

Kentucky bluegrass Big bluestem Canadian hawkweed Purple meadow-rue

(Poa pratensis L.) (Andropogon gerardii (Hieracium canadense (Thalictrum dasycarpum

Vitman) Michx.) Fisch. & Ave-Lall.)

Sweetgrass Sideoats gramma Roundhead lespedeza Bluejacket

 (Hierochloe odorata (Bouteloua (Lespedeza capitata (Tradescanta

(L.) P. Beauv.) curtipendula (Michx.) Torr.) Michx.) ohiensis Raf.)

Bluejoint Witchgrass Prairie blazing star

(Calamagrostis (Panicum capillare L. (Liatris pycnostachya

canadensis (Michx.) var. agreste Michx.)

P. Beauv.) Gattinger)

Reed canarygrass Little bluestem Wild bergamot

(Phalaris arundinacea (Schizachyrium (Monarda fistulosa L.)

L.) scoparium (Michx.)

Nash)

Quackgrass Canada wildrye 

(Elymus repens (L.) (Elymus

Gould) canadensis L.)

Dropseed

(Sporobolus R. Br.) a

 Cool-season species typically initiate growth and flower before warm-season species.
32

GENERAL TECHNICAL REPORT PSW-GTR-224

grasslands with 1 year of accumulated thatch could burn

anytime during a March-to-November study of flammability and consumption.

The majority of thunderstorms occur from April to

October, and the months in between comprise the typical

fire season. Of lightning-ignited fires in grasslands of the

Northern Great Plains from 1940 to 1981, nearly all occurred during the growing season from May through

September, with 73 percent occurring in July and August

alone (Higgins 1984) (fig. 11). Bragg (1982) noted that

over two-thirds of lightning fires in grasslands of Nebraska

during the years 1971 to 1975 occurred in July and August

(fig. 11). Lightning strikes may have ignited fires in advance of precipitation during thunderstorms, but could

also have occurred in conjunction with precipitation in

areas of higher fuel loading and thatch buildup (Bragg

1982). Native Americans also set fire to grasslands to

clear vegetation and to aid with hunting (Anderson 1990,

Axelrod 1985, Stewart 2002), and may have done so anytime the vegetation was dry enough to burn—i.e., during

both the growing season and the dormant season for vegetation (Reinking 2005). Higgins (1986b) wrote that

Native Americans “did not pattern their use of fire with

the seasonal patterns of lightning fires,” burning both in

the spring and fall dormant seasons, when lightning ignitions were infrequent. In Illinois, the preferred time for

igniting grassland fires for hunting purposes was apparently during warm dry spells in the fall, following the first

killing frosts (McClain and Elzinga 1994).

Prescribed Fire Regime

Recognition that fire plays an important role in maintaining grasslands has led to widespread use of prescribed fire,

initially to promote livestock forage and later for restoration goals such as reduction of woody vegetation. The season of prescribed burning differs, but for operational ease,

the majority of burns are typically conducted when vegetation is dormant in the early spring or late fall (Bragg 1982,

Ehrenreich and Aikman 1963, Howe 1994b). Spring burning (often late April) is the norm in tallgrass prairie remnants such as the Flint Hills (Seastedt and Ramundo 1990)

(fig. 12a and b), which extends from Kansas into northeastern Oklahoma. Fire at this time of year is thought to be

most beneficial to warm-season perennial grass species that

Figure 11—Percentage of lightning-ignited wildfires by month for grasslands of Nebraska, compiled for the period from 1971 to 1975 (data from

Bragg 1982), and for four grassland areas in the

Northern Great Plains, compiled for the period

from 1940 to 1981 (data from Higgins 1984).
33

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

are important for grazing (Towne and Kemp 2003). Prescribed burning of grasslands farther south may be conducted earlier (January to March) (fig. 13). Overall, the

majority of prescribed burns occur either earlier or later

in the season, and at a time of greater plant dormancy than

the majority of natural lightning-ignited fires. Greater use

of growing-season burns has been advocated in order to

mimic historical timing of lightning ignitions (Howe

1994a). However, there is some debate whether the goal

with grassland burning should be to re-create grassland

conditions representative of 30 million years of grassland

evolution (predominantly growing-season lightning fires),

or whether the goal should be to re-create conditions as

they existed immediately prior to Euro-American settlement, which is thought to have been a mixture of growingseason lightning fires augmented by growing- and

dormant-season fires, ignited by Native Americans (Howe

1994a).

Fuel Consumption and Fire Intensity

The total amount of fuel consumed is generally considerably less for grassland burns than for burns in forested ecosystems. In addition, because much of the fuel in grassland

ecosystems is fine and dries rapidly, the amount of fuel

consumed by burns in different seasons does not typically

differ much, relative to other vegetation types. For example, Howe (1994b) noted that growing-season burns in

the middle of the summer (July 15) consumed an average

of 96 percent of aboveground biomass, whereas dormantseason burns conducted on March 31 consumed 100

percent of the aboveground biomass. In another study,

consumption ranged from 84 percent in growing-season

(mid-June) burns to >99 percent in dormant-season (April)

burns (Bragg 1982). Copeland et al. (2002) reported that

late-growing-season burns (Sept. 3) consumed 91 percent

of the litter, whereas dormant-season burns (April 23) consumed 100 percent of the litter. In dry mesquite-savanna

A

B

Steven Apfelbaum

Quinn Long

Figure 12—(a) Many prescribed burns in the Central grasslands are conducted when grasses are dormant, such as this one in March 2009 at the

Stone Prairie Farm, Wisconsin. (b) Some prescribed burns are also conducted during the growing season, especially when the objective is to

control hardwood encroachment or approximate the historical disturbance

regime prior to human intervention. Summer burn (late August, 2005) at

the University of Kansas Nelson Environmental Studies Area, near

Lawrence, Kansas).
34

GENERAL TECHNICAL REPORT PSW-GTR-224

grassland in south Texas, both winter (December-February)

and summer (August) burns covered 100 percent of the

ground surface (Ruthven et al. 2008).

When actively growing, plant tissue contains moisture

that needs to be vaporized for complete consumption to

occur. Grasslands may still burn when they appear green

because accumulated thatch and litter underneath can provide ample fuel. Owing to the green component, growingseason fire is often of lesser intensity, with reduced flame

lengths and rates of spread, compared to dormant-season

fire (Copeland et al. 2002, Ford and Johnson 2006, Steuter

1987). Also potentially playing a role are weather differences. Although air temperature (and the initial heat of the

fuel) is typically higher during the growing season, relative

humidity is also often higher at this time of year, particularly for tallgrass prairie ecosystems. Therefore, the suppressing effect of live fuels (and higher relative humidity)

on fire behavior is apparently usually greater than the

enhancing effect of higher air temperature. Growing-season

burns can also result in greater variation in intensity (Howe

1999) and more burn patchiness compared to dormantseason burns (Komarek 1965, Steuter and McPherson

1995). This patchiness may be important for the persistence of many grassland species with fire. Historically,

large ungulates like bison (Bison bison) likely reduced the

amount of thatch and broke up the fuel complex by

preferentially grazing some areas over others, leading to

patchy burns (Fuhlendorf et al. 2006). Without historical

grazing patterns, burns today (especially in the dormant

season) may be more uniform in coverage.

In a mesquite savannah ecosystem in southern Texas,

Ansley and Castellano (2007a) reported that summer burns

(September 1) were higher intensity than winter burns

(February/early March). However, because this location is

Figure 13—Spring (March) prescribed burn at Sevilleta National Wildlife Refuge on the

western edge of the shortgrass prairie. Prescribed burns in south-central grasslands are often

ignited earlier in the spring than burns in grasslands farther north, where frost and snowfall limit

drying of fuels.

 Laura Calabrese
35

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

farther south and warmer than the sites of other comparable

grassland studies, some grass species were actively growing at the time of both winter and summer burns (coolseason species during the winter, and warm-season species

during the summer). With fire-behavior suppressing live

fuels present in both seasons, the higher air temperatures

apparently contributed to the greater intensity of summer

burns. In another mesquite savannah study, Drewa (2003)

did not find any difference in fire intensity between burns

in January or August. However, in this case, the January

burns occurred when both cool- and warm-season grass

species were dormant, whereas the warm-season grasses

were still actively growing in August. Overall, less variability in intensity is generally found within and among grassland fires than among fires in plant community types that

contain woody fuels (Bond and van Wilgen 1996).

Ecological Effects of Burning Season

Grassland Vegetation

In a review of the literature, Engle and Bidwell (2001) concluded that prairies are far more resilient to burning in any

season than previously thought. For example, Johnson et

al. (2008) noted that most prairie forbs are resilient to burning in any season, with 75 of 92 species studied unaffected

by burns in different seasons. However, timing of fire can

alter certain grassland species directly through injury or

mortality, especially during vulnerable phenological

stages. Fire during the period of most active growth is

thought to be most damaging, because new plant tissues

are more sensitive to heat (Bond and van Wilgen 1996)

and because carbohydrate reserves are lower this time

of year (Wright and Klemmedson 1965). Wright and

Klemmedson (1965) compared fire in June, July, and

August on four bunchgrass species and found plants to

be most resistant to fire later in the season, presumably

when carbohydrate reserves were again replenished.

Needle and thread (Hesperostipa comata (Trin. & Rupr.)

Barkworth) was damaged most by June fires, when plants

were greenest. Squirreltail (Elymus elymoides (Raf.)

Swezey), which was still green to partially green in both

June and July was damaged most by July fires, when

outside temperatures were the hottest. Data from this study

demonstrated that depending on the species, both timing

in relation to plant phenology, as well as the total heat

experienced (from fire plus starting air temperature) may

play a role in the response. In a different grassland type—

mesquite savannah—the yield of Texas wintergrass

(Nassella leucotricha (Trin. & Rupr.) Pohl) was reported to

be nearly twice as high after summer fires than after winter

fires (Ansley and Castellano 2007a). This cool-season grass

species grows in the early season (February to June). The

winter fires (February/early March) therefore coincided

with growth, whereas the summer fires (September) were

ignited after the species had finished growth. In a study of

burning season on a rare forb, either spring (mid April) or

fall burns (mid September) increased the germination of

Spalding’s catchfly (Silene spaldingii S. Watson), which

grows from May through September (flowering in July

and setting seed in August), but response was greater after

spring burns (Lesica 1999). Benning and Bragg (1993)

noted significant differences in response of big bluestem

(Andropogon gerardii Vitman) to burns just 4 days apart,

with fires shortly after initiation of spring growth increasing subsequent stem height and numbers of flowering

culms compared to fires prior to initiation of spring growth.

All of these studies highlight the importance of evaluating

the effect on individual species in context of the timing of

fire in relation to phenology of the plant at the time of the

fire.

Much of the research on season of burning in grasslands has looked at the impact on the plant community. In

addition to direct effects of fire on certain species, grassland vegetation can also be altered indirectly through

changes in competitive relationships that occur when

injury or mortality to some species is greater than to others.

Prairies are typically composed of varying amounts of two

groups of grass species: the cool-season grasses (C3

 photosynthetic pathway) that experience peak growth from

approximately March through May and the warm-season

grasses (C4 photosynthetic pathway) that have peak growth
36

GENERAL TECHNICAL REPORT PSW-GTR-224

from approximately April through October (table 3). Prescribed burns in the spring can kill, damage, or inhibit

growth of early cool-season species that are active at this

time of year, thereby favoring later warm-season species

that have not yet started to grow (Howe 1994a, 1994b).

Conversely, prescribed fire during the middle of the summer at the peak of lightning and historical fire frequency

are more detrimental to the dominant warm-season grass

species, thereby favoring early-flowering cool-season

species, many of which have already finished growth

and dropped seed by this time (Howe 1994a, 1994b, 1995;

Steuter 1987). For example, population size of the early

perennial forb Golden zizia (Zizia aurea (L.) W.D. Koch), a

species that sets seed in early summer, was greater following August burns than May burns (Howe 1999). The summer burns more effectively suppressed the canopy of the

taller dominant warm-season grasses, creating an environment free from shading by thatch.

Altering the fire regime of the Central and Northern

Great Plains from lightning-ignited summer wildfire to

spring prescribed fire has possibly shifted species composition toward a greater proportion of warm-season grasses

(Anderson et al. 1970; Howe 1994a, 1994b). The warmseason grasses favored by spring (dormant season) prescribed fire are generally taller and outcompete other

species for light; burning at this time of year is therefore

thought to have contributed to rarity of formerly abundant

species, and reduced overall diversity (Copeland et al.

2002, Howe 1994b). Conversely, summer burns, by reducing competition by dominant warm-season grasses, have

been shown to favor early-flowering cool-season grasses

and forbs (Howe 1995, 1999). In a study comparing midsummer (July 15) and early spring (March 31) burns, Howe

(1994b) reported that early cool-season flowering species

such as black-eyed Susan (Rudbeckia hirta L.) and quackgrass (Agropyron repens (L.) Gould) increased in abundance after the mid-summer burns, whereas the spring

burns caused both to decline or disappear. One census of

unburned prairies found that the guild of early-flowering

species covered only 2 to 15 percent of the ground; after a

single mid-July burn, the cover of early-flowering species

rose to 46 percent (Howe 1994b). Because lightning fires

historically occurred most often during the summer, it is

believed that such early-flowering species were once more

abundant. With more early-flowering species in place of a

few dominant warm-season grasses, tallgrass prairies managed with summer (growing season) burns have higher species diversity than prairies managed with spring or fall

(dormant season) burns (Biondini et al. 1989, Howe 2000).

The greater heterogeneity in intensity and effects with

growing-season burns may be another reason for higher

plant diversity (Howe 1999). If biodiversity management

in tallgrass prairies is the goal, burning during the summer

active phase of the dominant grasses may be preferred

(Towne and Kemp 2008). Howe (1994b) suggested that

greater biodiversity can be maintained with a “chaotic

array” of burn seasons, such as what might have occurred

historically.

Extent of community shifts caused by different burning

seasons is largely dependent upon the mix of species present. For example, major changes in the plant community

have not been noted for tallgrass prairies dominated by

warm-season species. In a study of burning at Konza prairie

in Kansas where cool-season species are only a minor

component, Towne and Kemp (2003, 2008) noted a high

degree of resilience to fire in any season. Canopy cover of

warm-season grasses increased with burning in the fall,

winter, or spring (Towne and Kemp 2003). Whereas some

cool-season grasses did decline with repeated spring burning, low initial abundance apparently did not lead to

differences in the competitive relationships between

cool- and warm-season species among burning season

treatments. Even repeated growing season (summer)

burning, which was expected to suppress warm-season

grasses and increase cool-season species, had few strong

effects, possibly because watershed-scale burns in this

season were patchy and incomplete (Towne and Kemp

2008). That repeated burning in different seasons led to

few and slow changes of most species suggests that in this

grassland type, the impact of one or a few out-of-season

burns is likely to be relatively minor.
37

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

Studies in shortgrass prairies have also not demonstrated dramatic shifts in species composition with burning

season. Owens et al. (2002) found that grass population

size and forb composition in a shortgrass savanna did

not differ after fires in the growing season (July through

August) or dormant season (January through February).

Based on data showing few changes in species composition, but large reductions in grass cover and biomass production, dormant-season burning (April) was deemed less

likely to put the shortgrass prairie at risk than growingseason (September) burning (Brockway et al. 2002).

Another common goal of burning in grasslands is to

increase forage for livestock (Engle and Bidwell 2001).

In one study, forage production of a tallgrass prairie was

greater after late-spring burning (May 1) than after earlyspring burning (March 20), with mid-spring burning (April

10) intermediate (Anderson et al. 1970). Late-spring burns

were timed for the start of growth of dominant warm-season

grasses and preferentially killed or reduced cool-season

species that initiated growth earlier. Similar results were

reported by Towne and Owensby (1984) from the same

site 20 years later, with greater forage production after late

spring burns (May 1), than after winter (December 1), earlyspring burns (March 20) or mid-spring burns (April 10).

These plots had been burned annually since 1928. It was

thought that the earlier fires led to greater duration of bare

mineral soil exposure, with evaporation drying out the soil

and reducing plant growth. In some grasslands, higher

productivity with late-spring burns is likely partially the

result of species shifts. The dominant warm-season grasses

that are favored by such a fire regime are generally more

robust and taller than the subdominant forbs and coolseason grasses favored by summer burns (Howe 2000).

In shortgrass prairies, biomass production following

growing-season (July) burning was shown to be substantially less than following dormant-season (April) burning

(Brockway et al. 2002, Ford and Johnson 2006). Grass

cover was also significantly reduced after growing-season

burns, owing primarily to a drop in cover of buffalograss

(Buchloe dactyloides (Nutt.) J.T. Columbus), a late warmseason species (Brockway et al. 2002). Another study in a

similar grassland type showed that spring burns enhanced

forage production more than fall burns (White and Currie

1983). In a marsh plant community, summer (August) burning decreased the biomass of common reed (Phragmites

australis (Cav.) Trin. ex Steud.), whereas spring (May)

burns increased biomass, and fall (October) burns resulted

in no change in biomass (Thompson and Shay 1989).

Nonnative vegetation—

Prescribed fire is sometimes used to control nonnative

species in grasslands. Shifting the plant community by

timing burns to coincide with the most vulnerable stage

of the target nonnative species while favoring native species, is seen as key to success (Emery and Gross 2005,

MacDonald et al. 2007, Simmons et al. 2007). Emery and

Gross (2005) found summer (July) burning at the time of

flowering reduced population growth rates of spotted

knapweed (Centaurea maculosa L. ssp. micranthos

(Gugler) Hayek), whereas spring (April) and fall (October)

burns had no significant effect. Summer burns killed the

flowering stalk, but not the adult plant. Spring burns

allowed surviving plants to flower, whereas fall burns

occurred after seeds had dropped. Summer burning is,

however, not the best time for native late-season grasses

that can keep spotted knapweed at bay through competition. In another study, mid-spring burning (late April to

late May), timed to kill newly germinating seedlings,

increased the dominance of native warm-season grasses

and reduced spotted knapweed abundance (MacDonald

et al. 2007). Because many of the nonnative species in

grasslands dominated by native warm-season grasses

germinate early, numbers can often be suppressed more

effectively with spring burning, which also tends to be

beneficial for the native grasses (Smith and Knapp 1999).

Unfortunately, because many nonnative species thrive

with disturbance, populations are also often enhanced by

fire (D’Antonio 2000). Hotter fall burns (September) were

found to increase numbers of the nonnative sulphur

cinquefoil (Potentilla recta L.) more than spring (April)

burns (Lesica and Martin 2003).
38

GENERAL TECHNICAL REPORT PSW-GTR-224

Trees and other woody vegetation—

The presence of trees and shrubs in grasslands is often

limited to areas that have experienced a break in the fire

regime. This is evident in oak savannas that occur along

the transition zone between the tallgrass prairie and the

deciduous eastern forests. It is thought that fire may have

once kept grasslands free of fire-sensitive tree species

(Axelrod 1985). For example, absence of fire in Texas

grasslands has contributed to an influx of mesquite and

scrub oak. Although growing-season burns in mesquite

savannas are sometimes higher intensity than dormantseason burns with greater flame length and faster rates of

spread, Ruthven et al. (2003) reported that either dormantor growing-season burns reduced the cover of mesquite

and other shrubs. However, these shrubs also resprout, and

Owens et al. (2002) found that neither growing-season nor

dormant-season burns were intense enough to kill them.

Intensity differences among burning seasons (higher intensity with summer burns) were associated with mortality

of prickly pear cactus (Opuntia spp.), a grassland invader

(Ansley and Castellano 2007b). Drewa (2003) manipulated fire intensity by adding fuels around the base of

some mesquite shrubs prior to growing- and dormantseason burns, and noted that although resprouting was

on average less vigorous after growing-season burns, resprouting was reduced in higher intensity patches within

burns in both seasons. Both intensity and fire season apparently play a role, influencing the outcome by different

mechanisms; intensity by damage to growing parts, and

season through changes in rate of recovery as a result of

seasonal differences in carbohydrate storage (Drewa 2003).

Soils

Grass fires move rapidly, and because the amount of fuel

consumed and heat produced when grasslands are burned

is relatively low, heating below the immediate soil surface

is generally minimal (Anderson 1990, Vogl 1979). However, burning is thought to influence soil heating in other

ways. Growth in the spring, especially in areas with cooler

winters, is limited by soil temperature (Ehrenreich and

Aikman 1963). Clearing the ground of litter and thatch,

whether through spring burning or other means, allows

sunlight to reach and heat the soil surface (Ehrenreich and

Aikman 1963, Knapp 1984, Seastedt and Ramundo 1990),

thereby promoting earlier growth of the warm-season

grasses (Howe 1994b, Vogl 1979). Earlier initiation of

growth and a longer growing season may be one reason

why spring prescribed fire has frequently been found to

increase vegetative production; however, Ehrenreich and

Aikman (1963) found that plants in burned areas also

senesced earlier without significantly greater production.

Burning in the early spring can lower soil moisture

compared to burning in late spring, because the ground is

exposed for a longer period of time, allowing more evaporation to occur (Anderson et al. 1970). Excess evaporation

may be detrimental to herbage production, especially in

low rainfall years when moisture is already limited. The

presence of litter and thatch is also thought to increase

snow accumulation, time required for snow to melt, and

rate of moisture infiltration into the soil (Ehrenreich and

Aikman 1963). Therefore, any burning regime that leaves

the soil uncovered during the winter and spring could

potentially reduce soil moisture.

Burning during the growing season minimized the

impact on biological soil crusts compared to burning

during the dormant season, presumably because of lower

fire severity (Ford and Johnson 2006). However, recovery

of soil crusts was rapid, regardless of the burn season (Ford

and Johnson 2006).

Wildlife

Wildlife is impacted by fire in two main ways: direct

mortality and indirect changes through alteration of their

habitat. However, few data on the effect of prescribed fire

season on wildlife in grasslands have been published.

Most studies have looked at either the difference between

growing-season fire and no fire or the difference between

dormant-season fire and no fire, without comparing among

burning seasons.
39

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

Birds and small mammals—

Some mortality of birds and small mammals is expected

with fire, especially those that nest above the ground.

Burns, stress, and asphyxiation are possible mechanisms

(Kaufman et al. 1990). However, prairie vegetation is composed of fine fuels that burn rapidly in a narrow band of

flame; this makes heating relatively transient and allows

animals to more easily escape (Vogl 1979).

Because of their mobility, adult grassland birds rarely

experience direct mortality with fire. Young birds still in

the nest are more vulnerable, and as a result, spring burning

during the nesting season may cause greater mortality than

burning in the summer or fall (Reinking 2005). Erwin and

Stasiak (1979) and Higgins (1986a) observed that nests

of ducks and other bird species were destroyed by spring

prescribed burns. However, if the nest is lost, many prairie

species will renest (Reinking 2005). In the Nebraska grassland, harvest mouse mortality was noted with a dormantseason burn occurring during the nesting season (Erwin

and Stasiak 1979). Even so, the high reproductive capability of rodents generally compensates for any impact of

different seasons of fire (Kaufman et al. 1988).

As in other vegetation types, longer term changes in

animal numbers owing to fire are thought to be caused

mainly by effects on habitat. Burning-season-mediated

shifts in grassland species composition can affect animal

populations, but lack of cover likely plays a stronger role

(Kaufman et al. 1990). In grasslands, fire removes all or

nearly all of the aboveground biomass, and it is thought

that amount of time without cover can affect wildlife

(either positively or negatively, depending on the species).

Because vegetative growth typically starts with warm

weather and precipitation in the spring, cover is generally

reduced for a longer period after a fall burn than a spring

burn (Kaufman et al. 1990). In a study comparing duck

nesting in plots burned in the spring (June) during the May

1 through July 31 nesting season, and plots burned in the

late summer (August-September), after the nesting season,

Higgins (1986a) noted that far fewer ducks initiated nests

the following spring in the sparse cover after late-summer

burns. In the following year, number of nests was the same

between treatments, but nest success was greater in the

late-summer burning treatment compared to the spring

burning treatment. This difference equalized over time,

and by year 4, no difference between burning seasons was

found. Westmeier (1973) found that nest densities of the

greater prairie chicken (Tympanuchus cupido L.) increased

after burns in either the late summer (August) or spring

(March).

Losses or gains in food sources may lead animals to

migrate, and loss of cover could increase predation rates

during migration. When managing for key animal species,

burning has sometimes been done to manipulate the

abundance of plant food sources, but as pointed out by

Sparks and Masters (1996), the optimal time of the year

differs, with no time best for all species. For example,

growing-season prescribed fire, which favors grasses over

forbs, may benefit species that feed on grass seeds but not

be ideal for species that eat seeds of legumes and other

forbs. If the goal is to increase the abundance of forb food

sources, dormant-season burns may be preferred.

A variable fire regime with burns in multiple seasons

may be necessary to maximize grassland biodiversity.

Fuhlendorf et al. (2006) and Reinking (2005) argued for

less uniform burn management and greater patchiness to

promote multiple grassland habitats and greater diversity

of birds.

Amphibians and reptiles—

In a mesquite savanna grassland in Texas, dormantseason burns had no effect on the diversity and abundance

of amphibians and reptiles, whereas diversity and abundance tended to be slightly greater in plots managed with

growing-season burns (Ruthven et al. 2008). One species

of lizard was 10 times more abundant in plots burned in

the growing season than in the unburned control; however,

burning season overall had few short-term effects on the

community. These authors recommended a varied fire

regime to maximize diversity of this group of species.

Arthropods—

Burning in different seasons has been used in attempts to

control arthropod pests, such as ticks, but results from
40

GENERAL TECHNICAL REPORT PSW-GTR-224

studies to date have been mixed because of differences in

fire intensity and timing in relation to periods of aboveground activity (Warren et al. 1987). For winged arthopods

that can escape the main heat pulse, a fire will tend to

favor those species that are mature at the time of the burn.

In a Kansas prairie, grasshoppers (Acrididae), which overwinter as eggs in the soil, were reduced by burns timed to

occur after the nonflying nymphs emerged (Knutson and

Campbell 1976). In another study, Vermeire et al. (2004)

found that the response to burning season differed among

grasshopper species, with two unaffected by prescribed

fire in any season, one reduced by both spring and fall

burns, and one reduced more by fall than spring burns.

The latter species lays eggs near the soil surface, where

they are presumably killed by fall burns. Developing

burning prescriptions to target vulnerabilities of each

species was suggested (Vermeire et al. 2004). Mortality

of other arthopods, such as centipedes and millipeds that

live in crevices in the soil, is likely minor with flaming

combustion, varying more with habitat modifications

(Warren et al. 1987) that may differ among burning seasons. Conditions as vegetation recovers following a fire

are beneficial to many arthopods. Total weight of insects

in a Texas grassland was greater in burned than unburned

areas one season after fire, with more insect biomass following spring burns than winter or fall burns (Chamrad

and Dodd 1973), presumably owing to attributes of the

postfire vegetation. In another study of a Texas grassland

burned three times in either the winter (dormant season) or

summer (growing season), and sampled 3 years after the

final burn, Johnson et al. (2008) found 170 percent more

individual insects in the summer burn plots. Although species richness was also 60 percent higher in the summer

burn plots, the difference between burn-season treatments

was only marginally significant.

Spring prescribed burning was shown to suppress

arthropod diversity in an Illinois tallgrass prairie (Harper

et al. 2000). Because of low survival in place, recovery of

burned landscapes may depend upon recolonization from

adjacent unburned areas. These authors suggested that to

avoid negative impacts to arthropods, managing for burn

patchiness and leaving unburned refuges would be beneficial. Burn seasons were not compared experimentally;

therefore the magnitude of potential burn season effects is

unknown.

Implications for Managers

Reviews of the literature on burning season in prairies of

North America highlight the wide range of outcomes that

are possible, making broad generalizations a challenge.

How fire interacts with the ecosystem depends on the

frequency of fire, time since previous fire or successional

stage of the grassland, grassland type (shortgrass, mixedgrass, or tallgrass), the evenness of cool-season and warmseason species within grassland type, herbivory, and

climatic conditions, most of which differ among the many

studies that have been done. Timing of fire in relation to

seasonal growth is key to understanding response in grassland species. For example, a greater increase in production

with spring than summer burns might be expected in areas

where warm-season grasses currently predominate (i.e., the

more mesic grasslands in the eastern portion of the Central

region), than in areas where cool-season grasses predominate, such as the more xeric western grasslands.

In many studies, the descriptions of when the fires were

conducted is vague (i.e., spring burn, early-season burn, fall

burn, late-season burn, etc.) and slight variation in timing

of fire in relation to plant phenology can produce different

results. Significant variation in outcomes has been reported

for burns conducted as little as 4 days to 3 weeks apart.

Effect of prescribed burning also depends on year-to-year

variation in rainfall, with greater expected plant mortality

from burning in drought years. Year-to-year variation in

precipitation or temperature can also alter the onset of

seasonal activity and growth. Thus lack of detail on the

exact phenological stage of organisms at the time of burning, inadequate description of burn timing, and climatic

variability limit syntheses and generalization of results.

Each year and each burn is potentially unique.

Although prairies are generally fairly resilient to

burning in any season, it is clear from the literature that

prescribed burns in different seasons can sometimes lead to
41

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

The optimal burn season depends on the objective: Is the

goal to improve forage biomass, or to enhance native

biodiversity? If the latter, is the goal to restore conditions/

processes to the time immediately prior to Euro-American

settlement, taking into account the impact that Native

American burning likely had, or is the goal to mimic

processes in place prior to anthropogenic manipulation

of vegetation?

Fine-tuning the timing of burns will depend on many

other factors including a complete understanding of the

phenology or periods of greatest vulnerability of key

species, the role of climatic variation, and the interaction

between phenology and climate. It is also important to

recognize that repeated burning in any one time of year

over large land areas may have the effect of simplifying

the system. With grassland species differing in response

to timing of fire, heterogeneity in the prescribed burning

regime, including a mix of fire seasons, may be necessary

to maintain prairie diversity.

Key Points—Central Region

• Grasslands exhibit resiliency to fire in any season, but substantial changes in community composition can

result from altering the burn season.

• Shifts in the plant community are caused by variation in phenology and susceptibility to fire among

species. Prairies with a mix of cool-season and warm-season species having different periods of growth

appear to be most susceptible to community shifts, whereas prairies dominated by one or the other appear

to be more resistant to change.

• Phenology of the vegetation at the time of burning appears to play a more important role in grasslands than

most other vegetation types, presumably because fuel consumption and fire intensity do not differ

substantially among burn seasons (assuming the same firing strategy: backing or heading). When intensity

is similar, the influence of phenology is more likely to be seen.

• Low fuel loading and rapid fire passage allows most mobile animal species to escape the flames in any

season. Although fire during periods of vulnerability, such as the nesting season, can cause short-term

losses, the effect on populations in the longer term is unclear.

• A burn program that promotes heterogeneity, including burning in multiple seasons within the historical

range of variability will likely benefit the greatest number of grassland species.

substantial ecological change. Population sizes of certain

species can shift from a single burn, but such changes are

usually ephemeral. The role of vegetation phenology on

response is perhaps stronger in grasslands than in any other

ecosystem. As with other ecosystems, response is tied to

both the phenology of organisms and intensity of the fire.

Because grassland fires consume much, if not all, of the

aboveground vegetation regardless of fire season, differences in intensity among seasons are likely of lesser

magnitude than in many other vegetation types. (Note,

however, that intensity does differ depending on whether

heading fires or backing fires are used, and this too can

affect the vegetation response [Bidwell et al. 1990]). Without strong differences in intensity, the role of phenology

becomes increasingly important.

Rather than managing fire intensity through prescribed

burning under different conditions or time of year, or using

different firing strategies, the issue for the fire manager is

mainly one of timing burning to achieve different goals.
42

GENERAL TECHNICAL REPORT PSW-GTR-224
43

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

Chapter 5: Eastern Region

transpiration rates of the vegetation are low (Chen and

Gerber 1990) (fig. 14c). The Ouchita Mountains of western

Arkansas and eastern Oklahoma also have two distinct wet

periods, with rainfall peaking around May and November

(fig. 14d). Farther east, rainfall generally becomes more

evenly distributed throughout the year (fig. 14b).

Forests of the Subtropical region are dominated by

overstory pines and scrub oaks closer to the coast, bottomland hardwoods along waterways, and a mixture of pines

and upland hardwoods farther inland from the coast. The

longleaf pine (Pinus palustris Mill.) forest once covered

the majority of the Coastal Plain, but has been reduced

owing to a variety of past land management activities

(Frost 1993). Shortleaf pine (Pinus echinata Mill.) was

at one time the most widespread pine species across the

East, occupying a variety of soil types and environmental

conditions, but has been reduced because of fire exclusion

(Komarek 1968), logging, and replanting with other species. Slash pine (Pinus elliottii Engelm.) and loblolly pine

(Pinus taeda L.) were historically restricted to wetter areas

but are now found throughout the pine zone (Komarek

1968). Forest composition in the pine-oak forests has

shifted to a greater percentage of mesophytic hardwoods

and planted species such as loblolly pine (Nowacki and

Abrams 2008).

Historical fire regime—

Based on fire scars in the tree ring record, it is believed

that prior to Euro-American settlement, many forests in

the south had an average fire-return interval of less than

15 years (Henderson 2006, Huffman et al. 2004, Huffman

2006, Wade et al. 2000) (table 4). The mean fire-return

interval increased along a moisture gradient with more

mesic sites having less frequent fire. The southernmost

longleaf pine forests grow in areas with longer dry periods,

and these forests burned with the highest frequency. Oakpine forests in the highlands of eastern Okahoma burned

every 2 to 12 years (Masters et al. 1995). Fires ranged from

The Eastern region includes everything east of the Central

grasslands to the Atlantic coast. Much of this area has the

potential to support forest vegetation. Forests have been

cleared and converted to farmland in many areas, with

major disruptions of the historical fire regime. Precipitation

is generally higher here than in the Central grasslands to

the west. Like the grasslands, much of the Eastern region is

characterized by fuels that respond quickly to fluctuations

in moisture (Wade et al. 2000).

Climate, Vegetation, and Fire

Past landform shifts, latitudinal temperature and precipitation gradients, as well as the presence of disturbances

such as fire have all shaped forest cover types of the

Eastern United States. For this synthesis, we divide the

Eastern region into two zones with distinct fire regimes.

The Subtropical zone in the south consists of forests

historically dominated by a mix of overstory pine species

along the Coastal Plain grading into pine-oak forests in

other areas (fig. 14). The Hot Continental/Warm Continental zone is dominated primarily by hardwoods in the

central Eastern United States, and a mix of coniferous and

deciduous species farther north.

Subtropical

The Subtropical region averages approximately 50 in of

precipitation a year. Much of the rainfall occurs with the

passing of maritime tropical air masses that arise from the

Gulf of Mexico, as well as continental polar airmasses.

These same air masses bring warm temperatures and high

humidity in the summer months and cold temperatures in

the winter. In south Florida, rainfall peaks in the summer,

with drier weather in the fall, winter, and spring (Bailey

1980, Beckage et al. 2003) (fig. 14e). Along the gulf coast,

two distinct wet periods occur: one during the summer

lightning storms (June-August) and a second during the

winter (January-March) with the arrival of cold fronts when
44

GENERAL TECHNICAL REPORT PSW-GTR-224

Figure 14—Climographs (monthly average temperature and precipitation) and the average time of the year of peak historical

and prescribed fire seasons from five representative locations within the Eastern region: (a) Chillicothe, Ohio; (b) Clemson,

South Carolina; (c) Ocala, Florida; (d) Fort Smith, Arkansas; and (e) Everglades National Park, Florida. Note that because the

timing of anthropogenic fire is poorly understood, the historical fire season reflects mainly lightning-ignited fires. Historical

anthropogenic fires were likely ignited during the dormant season (early spring and fall) as well.
45

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

Table 4—Estimated fire-return interval and approximate season of fire in the Eastern and Southern United States,

determined from fire scars

Firereturn Season

Location Forest type Dates interval of firea Author(s)

Years Percent in

category

Garrett County, Oak forest 1615–1958 7.6 D (>90) (Shumway et al. 2001)

Maryland

Jefferson Southern 1694–2004 3.3 D (80) (DeWeese 2007)

National Forest, Appalachian G (20)

Virginia pine forest

Southeastern Ohio Mixed oak 1871–1997 5.4 D (69) (Sutherland 1997)

S (25)

Pope County, Oak— 1680–1910 2.8-11.2 D (Guyette and Spetich 2003)

Arkansas shortleaf pine

Choccolocco Montane 1589–2006 3.2-11.5 D (Bale et al. 2008)

Mt., NE Alabama longleaf pine

Big Thicket, Longleaf pine 1668–1984 10.6 D (75) (Henderson 2006)

Texas forest E (9)

A (11)

Sandy Island, Longleaf pine 1580–2004 15.6 D (72) (Henderson 2006)

South Carolina forest E (9)

M (12)

A (2)

Eglin Air Force Longleaf pine 1517–2004 6.4 D (28) (Henderson 2006)

Base, Florida forest E (25)

M (9)

A (30)

Gulf County, Mainland 1592–1883 2–3 D (5) (Huffman 2006)

Florida pine savanna LS(10)

M (80)

Little Saint Coastal slash 1864–2000 4–9 D (14) (Huffman et al. 2004)

George Island, pine M (67)

Franklin

County, Florida

a

 D = dormant (September—March), G = growing season (March—October), E = early spring (March—April), S = spring (March—May), LS = late spring/summer

(April—May), M = mid-season (May—August), A = late season (August—September).
46

GENERAL TECHNICAL REPORT PSW-GTR-224

low-severity surface fires in longleaf pine forests to

mixed-severity fires in oak-pine forests (Wade et al. 2000).

Drought conditions sometimes led to higher fire intensity

and greater damage to trees, even in areas that typically

experienced low-severity fire.

The Coastal Plain pine zone is characterized by flammable understory fuels (both live and dead) that respond

quickly to fluctuations in moisture, and will burn nearly

year round. Moisture content of live vegetation is often

lowest from April through June, prior to the onset of the

summer rains and as temperatures are warming (Hough

1973, Hough and Albini 1978). Understory shrubs such as

saw palmetto (Serenoa repens Bartram (Small)), gallberry

(Ilex glabra (L.) A. Gray), and wax myrtle (Morella cerifera

(L.) Small) that contain volatile oils are common (Van Lear

et al. 2005). These oils allow combustion at high moisture

levels, and fire is therefore not dependent upon longer

periods of drying. Burns can be conducted just days after

rainfall (Schroeder and Buck 1970).

Location of fire scars within annual tree rings suggest

that slash and longleaf pine communities adjacent to the

Gulf of Mexico were most likely to burn in the middle of

the growing season (April—August) (Henderson 2006,

Huffman 2006, Huffman et al. 2004) (table 4). In contrast,

fire scars from other parts of the longleaf pine range showed

evidence of a higher proportion of dormant season fire

(September—March) (table 4).

The Southeast has the highest lightning strike frequency in the United States. Ten to 21 strikes per square

mile per year are common, with a peak of 22 to over 41

strikes per square mile per year found throughout much of

Florida (National Oceanic and Atmospheric Administration

2009). Most convective activity occurs between May and

September, with the number of lightning-ignited fires peaking in July (Myers 2000) (fig. 15). Lighting-ignited fires

burn more acres in May and June, at the transition between

the dry season and the onset of the summer rains (Barden

and Woods 1973, Komarek 1964, Myers and White 1987,

Outcalt 2008, Petersen and Drewa 2006). Thus, the peak

fire season occurs prior to the peak convective season

(fig. 15). Precipitation increases with the summer convective storms (Komarek 1964, 1968), and although fires do

occur during the summer, they are typically less intense

and smaller as a result of higher moisture levels. During

drought years, fire season tends to expand into the warm

summer months, and fires are often larger and more intense

(Glitzenstein et al. 1995b). In the Ouachita highlands of

eastern Oklahoma, the majority of lightning fires occur in

August and September, with drying following an early

summer peak in thunderstorm activity and rainfall (Foti

and Glenn 1991).

There is some question as to the temporal and spatial

extent of burning by Native Americans and its effects on

native species in the East, leading to controversy as to

what constitutes the historical fire regime (Henderson

2006, Myers 2000, Robbins and Myers 1992). Native

Americans used fire for a number of reasons, including

propagating native plants, hunting, clearing of land, defense, and communication (Fowler and Konopik 2007). It

is thought that Native Americans would not have restricted

their burning to a particular season, but rather used fire in

a variety of seasons to meet their needs (Johnson 1992,

Stewart 2002). Ignitions in the fall or late winter to early

spring would have coincided with hunting season and

preparation of agricultural fields, respectively (Fowler

and Konopik 2007, Henderson 2006). The prevalence of

Figure 15—Percentage of lightning-ignited fires and acreage

burned by month within the St. Marks National Wildlife Refuge,

Florida, between 1980 and 2006 (compiled from incident records in

FAMWEB).
47

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

dormant season scars, despite the relative lack of lightning

during this time suggests a contribution of Native American ignitions to the fire regime (Henderson 2006) (table 4).

Prescribed fire regime—

Active prescribed fire management was preceded by a

period of fire exclusion that began in the 1920s (fig. 16).

However, the practice of fire suppression was not widely

embraced in the East. Passage of the Weeks and ClarkeMcNary Act, which established significant fire control

organizations, sparked a debate concerning the importance of fire to the Eastern ecosystems (Chapman 1932).

Managers quickly realized the importance of fire to this

region and continued to use prescribed burning to maintain forested ecosystems. A substantial proportion of the

prescribed burning in the southeastern coastal plain is

conducted during the fall, winter, and spring (October—

April) (fig. 17). Burning at this time of year is/was based

on the belief that such burns would be less likely to

impact nesting birds or harm growing trees (Cox and

Widener 2008). Operational issues also favor burning at

this time of year, with lower temperatures and more

predictable winds making prescribed burns easier to

conduct (Wade and Lunsford 1989). Gusty and unpredictable winds associated with thunderstorms are common

during the late spring and summer.

The main prescribed burning season occurs during

the dormant phase of many plant species and at a time

of reduced biological activity, whereas the historical fire

regime consisted of a higher proportion of spring and

summer fire, when vegetation was actively growing and

birds were nesting. Because these dormant-season burns

Figure 16—Summary of ignition source, fire type, and season of

fire attributed to natural and human causes in the Eastern region

during different time periods (based on Komarek 1964, Robbins

and Myers 1992, Frost 1998, and Fowler and Konopik 2007).

Figure 17— Percentage of prescribed burns by month at (a) St. Marks National Wildlife Refuge, Florida for the period from 1980 to 2006

(compiled from incident records in FAMWEB), and (b) Everglades National Park, Florida for the period from 1948 to 1997 (data from

Myers 2000).
48

GENERAL TECHNICAL REPORT PSW-GTR-224

are outside of the typical historical period of lightningignited fire, there is some concern that repeated burning

at this time of year may result in undesirable ecological

changes.

On lands where the objective of prescribed burning

is to restore historical processes, such as Everglades National Park, considerable burning is also often done during the growing season (May—August), the peak of the

lightning season (fig. 17b). In Everglades National Park,

South Florida slash pine (Pinus elliottii Engalm. var. densa

Little & Dorman) has been burned every 2 to 3 years

during the months of May and June since 1989 (Slocum et

al. 2003). However, such burns are considered risky

because of the chance for escape. Specific resource conflicts, such as game bird management and concern about

nest destruction, can also make managers reluctant to use

growing-season prescribed fire (Cox and Widener 2008).

Two excellent reviews and syntheses by Robbins and

Myers (1992) and Streng et al. (1993) cover the issues of

prescribed burning in southern pine forests at times of the

year different from the historical lightning fire regime, and

we draw heavily on their conclusions for this synthesis.

Hot Continental and Warm Continental

The fire climate in the Hot/Warm Continental regions of

the Central, Great Lakes, and North Atlantic States is generally driven by airmasses bringing moist humid tropical

air in the spring and summer and polar continental air in

the late fall and winter. The annual precipitation averages

between 20 to 45 inches in the Central States and about 30

inches in the Great Lakes States (Bailey 1980). Although

precipitation is fairly well distributed throughout the year,

somewhat more rainfall occurs during the summer, coinciding with the highest temperatures of the year (fig. 14a).

Forests of the warmer areas from the middle-Atlantic

States through the Appalachian Mountains and into the

Northeast are mostly dominated by oaks and hickories. The

main carrier of fire is leaf litter, which differs in flammability with site and time of year (Wade et al. 2000). Recurring

fire has maintained oak dominance in these forests at the

expense of mesophytic tree species (Abrams 1992, Elliott

et al. 2004). On xeric sites, lack of fire has increased the

presence of ericaceous shrubs such as rhododendron

(Rhododendron L.) and mountain laurel (Kalmia latifolia

L.). When fire does occur, these shrubs can burn at high

intensity, potentially leading to stand replacement (Wade

et al. 2000).

Farther north, in the Great Lakes region, spruce (Picea

A. dietr.), fir (Abies Mill.), eastern white pine (Pinus strobus

L.), red pine (Pinus resinosa Aiton), jack pine (Pinus

banksiana Lamb.), and aspen (Populus L.) dominate as a

fire-maintained stage or as a climax forest (Duchesne and

Hawkes 2000). Fires in pure aspen stands are typically of

low intensity, but fire can still cause significant mortality

of aboveground stems (Duchesne and Hawkes 2000). Areas

with higher fuel loads in mature aspen stands may sometimes experience high-intensity stand-replacing fires (Jones

and DeByle 1985). Red and eastern white pines occupied

pure stands or were found in association with aspen, jack

pine, and a variety of hardwoods, and experienced similar

mixed-severity fire regimes as aspen stands (Duchesne and

Hawkes 2000, Johnson 1992).

Historical fire regime—

The fire regime of the mixed-oak forests consisted of

primarily low-severity events occurring approximately

every 3 to 13 years (Guyette et al. 2006, Shumway et al.

2001) (table 4). Farther north, in the Great Lakes and

Atlantic regions, typical fire-return intervals ranged from

35 to over 200 years (Duchesne and Hawkes 2000).

Location of fire scars within growth rings suggests

that most fires occurred during the dormant season (Bale

et al. 2008, DeWeese 2007, Guyette and Spetich 2003,

Shumway et al. 2001) (table 4). The dormant season lasts

from approximately late September to April. When hardwoods are not in leaf, litter is exposed to sunlight, making

it susceptible to fast drying. Aboveground portions of

understory herbaceous vegetation are also mostly dead at

these times of the year, increasing the likelihood of fire

spread (Wade et al. 2000). When leaves are on the trees, the

microclimate on the forest floor is often shady and moist,
49

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

which generally results in poor conditions for burning. The

exception occurs during periods of drought. Convective

activity is still high during this time (Petersen and Drewa

2006), and drought overrides the typically moist understory microclimate.

Lightning is fairly common and mostly associated with

summer convective storms that also typically include precipitation. However, because of the lack of dry fuels, the

peak fire season is frequently driven more by the surface

fuel conditions than the times of peak lightning frequency

(Barden and Woods 1973, Johnson 1992, Petersen and

Drewa 2006). The relative lack of lightning during times

of the year when leaves are on the ground and most

flammable, and tree ring data indicating predominantly

dormant-season burns, suggests the role of another ignition

source in many areas. Paleoecological evidence that fireresistant tree species were more abundant during periods

of Native American settlement also hints at a link between

forest composition, fire, and human activity (Henderson

2006, Shumway et al. 2001).

Prescribed fire regime—

Most prescribed burning in eastern hardwood forests

is conducted during the dormant season, prior to leaf

emergence in the spring or after leaf drop in the fall

(Johnson 1992). The period of time that the litter fuel

bed is receptive to fire depends on the latitude and yearto-year weather variation. In southern hardwood forests,

prescribed burning may be conducted any time the litter

layer is dry, whereas farther north, persistent snow cover

limits application of fire to a narrower period in the spring

and fall.

The prescribed fire season appears not to differ greatly

from the historical fire season, at least for the period of

Native American settlement. However, in areas where late

summer burning was historically part of the fire regime, a

higher proportion of the landscape is now possibly being

treated in the dormant season (fig. 18). Because vegetation

is dormant and wildlife species are less likely to be active

during the dormant season, concerns about direct fire

effects are minimized. The extent to which prescribed fire

effects differ from historical fire effects may be due primarily to differences in fire intensity, if any. One concern may

be the lack of heterogeneity in the fire regime when a

strictly dormant-season prescribed burning program is

employed.

Figure 18—Most prescribed burns in oak and mixed

oak and pine forests are conducted in the dormant

season after leaf fall and prior to leaf expansion in the

spring. During this time, exposure of the litter to

sunlight hastens drying and ignition. Stephen Hudson
50

GENERAL TECHNICAL REPORT PSW-GTR-224

Fuel Consumption and Fire Intensity

Fuel consumption and fire intensity do not appear to differ

consistently with season in eastern forests. In eastern pine

forests west of the Mississippi River, dormant-season prescribed burns are sometimes described as more intense than

growing-season burns (Sparks et al. 1998). Sparks et al.

(2002) reported dormant-season (March—April) fuel beds

in Arkansas shortleaf pine stands to consist of a reduced

proportion of live fuels, more fine (1 hour) fuels, and a

greater total fuel load than fuel beds in the growing season

(September—October), all of which led to greater fire-line

intensity and total fuel consumption when burned. In different fuel types of southern pine forests, growing-season

burns were described as more intense than dormant-season

burns (Komarek 1965, Liu and Menges 2005). Boyer

(1993) measured greater tree crown scorch with summer

burns than spring or winter burns, most likely as a result

of higher air temperatures. Within the growing season,

early fires (May) have been reported as generally more

intense than late fires (September) (Slocum et al. 2003).

Still other studies have not found differences in fire

intensity among seasons. Boring et al. (2004) noted

approximately the same peak temperature and flaming

duration with dormant-season (March—April) burns and

early growing-season (June) burns. Glitzenstein et al.

(1995a) found that burns conducted at eight different times

of the year consumed approximately the same amount of

fuel (43 to 62 percent), but differed in fire-line intensity by

a factor of four owing to variable rates of spread. This variation was not consistently associated with any time of the

year. In another study of fuel moisture and consumption,

burns in February, March, April, and September under

conditions that started out as wet and progressed to very

dry, consumed approximately the same amount of litter,

but burns in May and September (the two driest times)

resulted in greater duff consumption (Ferguson et al. 2002).

Growing-season burns in southeastern pine forests can

be patchier than dormant-season burns because of higher

moisture conditions and spatial variation in rate of green

up (Slocum et al. 2003, Wade et al. 2000). However,

Glitzenstein et al. (1995a) found that burns conducted at

eight different times of the year all burned across much of

the forest floor (96 to 100 percent).

In hardwood forests, prescribed burns typically reduce

the fine woody material, but leave the duff layer intact. A

fall (November) burn in southern Ohio was found to be

hotter than a spring (March) burn, presumably because the

leaf litter had freshly fallen and was less degraded/decomposed (Schwemlein and Williams 2007). The opposite,

with spring (April) prescribed burns having greater intensity than winter (February) or summer (August) burns has

also been reported (Brose and Van Lear 1998, Brose et al.

1999). In the latter study, relative humidity was the lowest

during the spring burns.

Differences among studies and among season-ofburning treatments within the same study may be due

to weather at the time of burning, timing relative to recent

precipitation events, or just year-to-year climatic variability. Variation among studies is also possible because of

differences in vegetation type and fuel bed composition.

One common trend is that air temperatures are often higher

at the time of growing-season burns, and higher initial

vegetation temperature means that less heating may be

necessary to reach the threshold for tissue damage (Byram

1948). In the Southeastern United States, greater intensity

often reported with growing-season burns may also be due

to the tendency to use heading fires (burning with the wind

or upslope) at this time of year. Higher relative humidity

and more fine fuel moisture can make backing fires (burning into the wind or downslope) slow moving or ineffective, leading to a greater use of heading fire to speed the

rate of burnout (Robbins and Myers 1992).

Ecological Effects of Burning Season

Trees

Pines—

Fire is a vital tool in the management of pine forests in

the Southeastern United States. Early research guided

managers who were focused on timber production to avoid
51

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

burning during the growing season (May—August)

because of concern that damage to the tree crop would

reduce profits (Bruce 1954).

Tree mortality can result from excessive root loss, damage to bole cambium, or crown scorch/photosynthetic material loss, some or all of which can differ among seasons.

Duff consumption, which is considerably greater when

moisture levels are low, has been linked to increased longleaf pine mortality in long unburned forests where substantial duff had accumulated (Varner et al. 2007). Lowest duff

moisture levels are often found from late in the dormant

season to early in the growing season (March—April)

(Varner et al. 2007). Mechanisms of tree mortality were

not determined, but could be the result of root death or

cambium damage. However, fine roots are less likely to

grow into the duff layer when it is dry. Thus greater duff

consumption may not necessarily translate into greater fine

root mortality. There is some evidence that root growth

may be reduced more following summer (July) burns than

dormant- (March) and early-growing-season burns (May)

(Sword Sayer et al. 2006). However, this later study occurred during a drought year and results may be most applicable to these drought conditions.

Early research showed that pine crowns were more

severely scorched by spring or summer (May—August)

burns than by fall and winter (October—March) burns,

owing to higher ambient temperature, which reduced the

time to reach lethal heating and cause foliage mortality

(Byram 1948). It was assumed that greater crown loss

would mean higher mortality or slower growth. To test the

effect of crown loss in different seasons, Weise et al. (1989)

experimentally defoliated loblolly and slash pines to

varying levels in January, April, July, and October, and

found substantial mortality only after October defoliation.

Southern pines form new buds and flush multiple times

during the growing season and can therefore recover better

from defoliation if it occurs prior to the last flush of the

season. In the fall, when no additional growth is expected

until the following spring, the time between tissue loss and

regrowth of photosynthetic structures is greater, which

apparently causes more stress on the tree.

In one of the most robust long-term studies of burning season (St. Marks study, St. Marks National Wildlife

Refuge, Florida), mortality of mature longleaf pines with

burns conducted at eight different times of the year did

not vary in any predictable way (Glitzenstein et al. 1995a).

Much of the other literature on postfire mortality or growth

rates of pines has also reported no effect or mixed results of

burning season (Boyer 1987, 1993, 2000; Waldrop et al.

1987, 1992). Magnitude of any burning-season effect may

be a function of tree age, with the seedling stage of some

species being more sensitive. Greater mortality was noted

in longleaf pine saplings following summer (August) than

following winter (January) or spring (May) burns (Boyer

1982, 1987) (fig. 19). Seedlings appear to benefit more

from growing season (May) burns, presumably at least in

part because fire at this time of year reduces the incidence

of fungal infections such as brown spot needle blight

(Scirrhia acicola (Dearn.) Siggers) (Bruce and Bickford

1950; Glitzenstein et al. 1995a; Grelen 1978, 1983; Wade

and Johansen 1986).

Fire intensity and season are confounded in many

burning studies. In south Florida, slash pine was found to

experience higher mortality with fall burns (September—

November) compared to burns in other seasons (Menges

and Deyrup 2001). However, char heights were also

greatest for the fall burns. Measures associated with fire

intensity (i.e., percentage of green canopy and bark char)

were most strongly associated with tree mortality within

burns in the same season as well. Glitzenstein et al. (1995a)

reported fall (October) burns to be the most intense in one

treatment cycle, and found that pine mortality in the 2 to

3.5 in diameter at breast height (dbh) size class appeared

to be correlated with amount of fuel consumed (and total

heat released), whereas mortality of pines greater than 3.5

in dbh was correlated with fireline intensity. Menges and

Deyrup (2001) noted that mortality was less when lowintensity backing fires were used, rather than heading fires.
52

GENERAL TECHNICAL REPORT PSW-GTR-224

Reviewing the literature, Robbins and Myers (1992)

concluded that burning season appears to have little effect

on the growth rate of mature pines, but mortality is often

somewhat more following fires late in the growing season,

which corroborates the previously mentioned defoliation

study of Weise et al. (1989). However, considerable variation in results have been reported, which probably has to

do with differences in fuel consumption and resulting fire

intensity among seasons, as well as the time of year burns

were conducted relative to tree phenology (Glitzenstein et

al. 1995a, Robbins and Myers 1992) (fig. 20). Glitzenstein

et al. (1995a) suggested that fire intensity may explain

much more of the variation in effects to longleaf pine than

either the ambient temperature at the time of burning or the

phenology/burning season.

Overstory hardwoods—

A frequent goal in restoring fire to eastern hardwood

forests is to improve growing conditions for a variety

of oak (Quercus L.) species. In the absence of fire, hardwoods such as maples (Acer L. sp.) and yellow poplar

(Liriodendron tulipifera L.) that would normally be

restricted to wetter sites have gradually moved into the

uplands, outcompeting and eventually replacing the oaks

(Nowacki and Abrams 2008). Because the majority of

prescribed burning in this forest type is done in the dormant season when litter is the most flammable (similar to

the main historical fire season), season of burning is less

of an issue. As a result, few burning-season studies have

been done in hardwood stands.

Of fires conducted in February, April, and August, the

April fires were, on average, the most intense and did the

most to favor oaks over yellow poplar (Brose and Van Lear

1998, 1999; Brose et al. 1999). Variation within burns was

exploited to investigate regeneration differences with burn

intensity and burn season. High-intensity fire was the most

effective at reducing yellow poplar and favoring oak in

each burn season. Although summer burns were predominantly low intensity owing to shading and higher relative

humidity, moderate- and high-intensity patches within the

fires at this time of year produced the strongest differences

in regeneration success between species. It therefore appears that both fire intensity and phenology play a role.

Figure 19—Comparison of the effects of prescribed burns conducted in a 6-year-old longleaf pine plantation established on old agricultural

fields at the Monroeville Experiment Station, Alabama burned in (a) February 2007 and (b) August 2007. Photos were taken in March of

2008. Fire intensity was greater in the August burns, causing more crown scorch. However, little mortality was noted with fires in either

season. Trees burned in the February fire dropped their scorched needles during the summer, but trees burned in the August fire retained

scorched needles throughout the winter.

A B both photos John McGuire
53

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

Understory Vegetation

Shrubs and hardwoods—

Maintaining adequate regeneration of overstory trees is

a common goal in the management of pine forests. To reduce competition for light, fire is used to selectively topkill the hardwood midstory and shrub understory, while

minimizing the impact to overstory pines. Numerous research results have suggested that burning during the

peak of the historical fire season (May) reduces stem

density of understory hardwoods more so than burns at

other times of the year (Boyer 1993, Drewa et al. 2006,

Glitzenstein et al. 1995b, Streng et al. 1993, Waldrop

et al. 1987, White et al. 1991). This is particularly true if

burns are repeated at annual or biennial intervals. A single

burn in any season will not kill enough plants to control

hardwood resprouting (Boyer 1990). However, following

43 years of burning on the Santee Experimental Forest in

A B

C D

Figure 20—Longleaf pine stands on the Escambia Experimental Forest, near Brewton, Alabama, that have been managed with prescribed

fire for multiple decades, showing both the phenology of vegetation at different times of the year and vegetation changes owing to timing of

the fire treatments. The top two photos are from a unit managed with a mix of growing- and dormant-season burns, with photos taken in (a)

March—dormant season and (b) July—growing season. The bottom two photos are from a unit managed with primarily dormant-season

burns, with photos taken in (c) March—dormant season and (d) July—growing season. Note that the growing season burns appear to have

more effectively reduced shrubby understory vegetation. The mechanisms of burning-season effects are often difficult to isolate because fire

frequency, fuel loading, and canopy cover can confound the role season may play in determining the outcome. all photos by Becky Estes
54

GENERAL TECHNICAL REPORT PSW-GTR-224

South Carolina, fewer hardwood sprouts survived with

a fire regime of annual late spring/early summer (June)

burns than annual winter (December) burns (Waldrop and

Lloyd 1991). With late spring/early summer burns, the

woody vegetation was gradually replaced by an understory dominated by forbs and grasses (Waldrop et al. 1987)

(fig. 20). In contrast, more oaks and other hardwoods were

maintained with repeated burns during the fall/winter dormant season (October—January) (Jacqmain et al. 1999). In

another study in shortleaf pine—grassland ecosystems of

Arkansas, late-growing-season (September—October)

burns were found to be less effective for reducing understory hardwoods than late-dormant-season (March—April)

burns (Sparks et al. 1999).

It is important to note that the regime of annual

growing-season burning that most successfully reduced

competing hardwoods and shrubs in two of the most

widely cited studies (Santee Experimental Forest study

(Waldrop et al. 1987, Waldrop and Lloyd 1991), and St.

Mark’s National Wildlife Refuge study (Glitzenstein et al.

1995a, 1995b; Streng et al. 1993) was considerably more

frequent and invariant than was likely the case historically

and therefore may not be the most beneficial for other components of the ecosystem. Although fires as frequent as 1 to

2 years apart have been recorded in the tree ring record of

eastern pine forests, the overall historical fire-return interval averaged 3 to 7 years (Henderson 2006). Annual prescribed burning may not even be possible in some stands,

if fuel accumulation rates are slower. For pines to regenerate naturally, longer fire-free periods may be necessary so

that seedlings can establish and grow above the zone of

lethal heat. Depending on the management objective, a

prescribed burning regime of variable frequency and seasonality (within the historical ranges) may be preferred.

There are several explanations relating both to the

physiological status of the plant and to fire intensity for

the difference in midstory hardwood and shrub mortality

following burns in different seasons. Physiological status

appears to play a role in the greater shrub and hardwood

reductions noted with growing-season burns in many

studies. During the dormant season, shrubs store more of

their carbohydrates underground, and these carbohydrates

enable resprouting when the aboveground portion is killed

by fire (Drewa et al. 2002). During the growing season,

more of the carbohydrates are allocated aboveground, and

are lost with topkill, leaving fewer reserves for resprouting.

Drewa et al. (2002) evaluated shrub response to fires of

different temperatures in different seasons and found

changes were not associated with fire intensity, suggesting

that the physiological status of the shrub at the time of

burning may be playing a greater role. Studies of defoliation of evergreen shrub species in different seasons also

point to a physiological influence, with one study reporting complete kill following leaf removal in October, but a

much reduced effect in April (Kramer and Wetmore 1943).

Other studies suggest an effect of fire intensity differences

among burn seasons (Drewa et al. 2006, Glitzenstein et al.

2003). Shorter statured vegetation, such as midstory hardwoods and shrubs, is more likely to be affected by seasonal

differences in scorch height (Robbins and Myers 1992). In

the Sparks et al. (1999) study, the dormant-season burns

(March—April) were both more intense and more effective

at thinning the midstory hardwoods than the late-growingseason burns (September—October), suggesting that

differential intensity may have overwhelmed effects of

phenology. Indeed, some burning-season studies have

reported fire intensity to be just as important as phenology

in shaping the outcome (Glitzenstein et al. 1995b, Sparks

and Masters 1996).

Less is known about effects of burning season on

understory shrubs in areas north of the southeastern pine

zone. In one of the few studies on the topic, Schwartz and

Heim (1996) noted that 95 percent of small saplings and

shrubs in an Illinois forest were top-killed by either a

single dormant-season (March) burn or a single growingseason (May) burn. In another study of understory response

in mixed-hardwood and pine forests of Minnesota, both

spring dormant-season and summer growing-season burns

completely top killed hazel (Corylus L.), but resprouting

was enhanced by repeated spring burning and reduced

by repeated summer burning (Buckman 1964). Because

humus was combustible during dry summer conditions,
55

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

fires at this time of year were more likely to kill the roots.

Carbohydrate reserves were also more likely to be exhausted following repeated summer burning.

Herbaceous understory—

Burning during the historical fire season has been hypothesized as important because organisms are presumably

best adapted to disturbance at this time of year. Studies

show that this may indeed be the case for some understory

plant species of southern pine forests. At the St. Marks

National Wildlife Refuge in Florida, greater increases in

shoot number and flowering of narrowleaf goldenaster

(Pityopsis graminifolia (Michx.) Nutt.) were observed

following burns in May than burns in January or August

(Brewer and Platt 1994a, 1994b; Brewer et al. 1996). The

increases in shoot numbers did not lead to long-term increases in stem densities, however, suggesting that there

may be some cost to using resources for reproduction

(Brewer 2006). Although the flowering response indicates

an adaptation to and dependence on growing-season fire,

Brewer (2006) hypothesized that this species would likely

benefit from “modest variability in fire frequency and fire

season.” Numerous grass species, including the commonly

studied wiregrass (Aristida beyrichiana Trin. & Rupr.),

also flower more vigorously after growing-season burns

(Main and Barry 2002, Outcalt 1994, Streng et al. 1993).

Saw palmetto produced more flowers and fruits with

periodic growing-season (April—July) burns than with

dormant-season (November—February) burns (Carrington

and Mullahey 2006). Growing-season burns have also

been shown to increase flowering synchrony of forbs and

shrubs by decreasing the flowering duration (Platt et al.

1988). Flowering synchrony may lead to a higher probability of cross pollination. In another study, no difference in the density of reproductive American chaffseed

(Schwalbea americana L.) plants was noted between

burning-season treatments (Kirkman et al. 1998). Numbers

increased following burning in either the growing or

dormant season. However, burning season did influence

the timing of flowering, with plants flowering earlier after

dormant-season burns than after growing-season burns.

The positive response of some species to growingseason burning provides evidence for fire at this time of

year being an important part of the natural disturbance

regime. However, what is best for one species may not be

for all, with some species also responding more strongly to

dormant-season burns (Hiers et al. 2000, Liu and Menges

2005, Sparks et al. 1998). Many species do not appear to

be influenced by burning season at all. For example, of the

more than 150 plant species evaluated for response to late

growing-season (September—October) and late dormantseason (March—April) burns in a shortleaf pine-grassland

community in Arkansas, fewer than 10 percent were differentially affected by burning season (Sparks et al. 1998).

The variable response of understory species to fire season

suggests that a heterogeneous fire regime (including variation in the seasonal timing of fire) may help conserve

biodiversity (Hiers et al. 2000, Liu et al. 2005).

For species with growth or flowering influenced by

burning season, response has sometimes been shown to

differ at fine temporal scales—i.e., for fires within the same

growing season (Negron-Ortiz and Gorchov 2000, Rideout

et al. 2003), or among plant growth stages (Spier and

Snyder 1998). Negron-Ortiz and Gorchov (2000) reported

that early wet-season (May—June) fires were beneficial and

late wet season (July—September) fires detrimental to the

cycad species Zamia pumila L. The variation in response of

herbs and woody plants observed among burns within the

growing season by Rideout et al. (2003) was attributed

mainly to climatic differences. Liu and Menges (2005)

noted that slight differences in burn timing within the wet

(growing) season had substantial effects on survival and

growth of big pine partridge pea (Chamaecrista lineata

(Sw.) var. keyensis (Pennell) Irwin & Barneby), and concluded that comparing fires by seasons may be too broad

and not useful to managers.

In a study of response of multiple growth stages,

small plants of the forest herb pineland Jacquemontia

(Jacquemontia curtisii Peter ex Hallier f.) suffered greater

mortality with growing-season (June) prescribed burns than

dormant-season (January) burns, even though the latter
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GENERAL TECHNICAL REPORT PSW-GTR-224

burns were hotter (Spier and Snyder 1998). However,

the plants surviving the growing-season burns produced

more flowers. Therefore, different parts of the plant lifecycle were variably affected by burning season. Similar

findings have been reported for wiregrass, where growingseason burns promote flowering, but also cause higher

mortality of established seedlings than dormant-season

burns (Mulligan et al. 2002, Streng et al. 1993), and big

pine partridge pea, where stem growth was greater but plant

survival lower following growing-season (summer) burns

(Liu and Menges 2005). These results all highlight the

importance of variability in the fire regime.

At the plant community level, repeated growingseason burning generally increases the cover of grasses

and diversity of herbaceous species (Drewa et al. 2002,

2006; Lewis and Harshbarger 1976; Waldrop et al. 1987;

White et al. 1991). This shift is likely because of release

from shrub competition (shrubs are selected against by

growing-season burns) and removal of the litter layer

(Lewis et al. 1982). The robustness of the understory herbaceous layer is not only important for biodiversity conservation, but also for grazing animals. Studies focused on

livestock management have reported grass productivity

gains with early growing-season burns (Grelen and Epps

1967, Lewis and Harshbarger 1976), which is likely also

tied to reduced shrub competition. However, overall productivity (herbs and shrubs) was found to be greater

following fall burns than spring burns (Schneider 1988).

Other studies that have followed productivity over several

years have been unable to document any increase in biomass and cover of grasses and forbs with burns in different

seasons (Streng et al. 1993).

Sparks et al. (1998) suggested that understory composition was, in part, influenced by fire intensity through its

effect on litter consumption and woody shrub removal.

However, fire intensity did not appear to play much of a

role in another study. Hierro and Menges (2002) burned

plots containing between 2.6 and 7.1 tons/ac of surface

fuel with and without 54.0 tons/ac of additional fuel, and

found little effect on understory shrub species richness or

density although the fuel addition treatment significantly

increased fire temperatures and soil heating. The authors

suggested that species are well adapted to variation in fire

intensity. In another study, plant mortality did not differ

with fuel consumption differences, suggesting that seasonal timing may be more important than fire intensity

(Liu and Menges 2005). However, a big picture view suggests that the effect of season of burning is less critical to

maintaining understory biodiversity in the longleaf pine

system, than frequency of burning (Palik et al. 2002).

One cautionary note on repeated burning: despite the

many benefits of growing-season burning that have been

reported in the literature, a recent publication from the St.

Marks study in Florida indicates that growing-season

burns, if applied annually, may over time actually reduce

the cover of plants such as wiregrass that are stimulated

to flower by fire (Glitzenstein et al. 2008). These results

warrant closer scrutiny to determine whether invariable

and frequent growing-season burns cause the grasses to

invest an excessive amount of carbohydrate reserves into

reproduction, thereby reducing the plants’ ability to grow

vegetatively over the long term (Brewer et al. 2009).

Much less literature is available for the understory

of eastern hardwood ecosystems than southern pinedominated ecosystems. Perennial herbs in oak forests

generally emerge from rhizomes and are dormant during

the typical spring and fall burning periods. Because heat

penetration into the soil with the burning of leaf litter is

generally minimal, resprouting from dormant rhizomes is

likely little affected by burning at either time. Any change

in the understory as a result of burning season is expected

to result more from indirect effects, such as reduced competition with top kill of midstory shrubs, or consumption

of the litter layer (Keyser et al. 2004). Keyser et al. (2004)

found that plant cover and species richness in an oakdominated forest increased following fire regardless of

whether burning occurred in February, April, or August,

but the more intense spring and summer burns led to a

shift toward herbaceous species, whereas the winter burn

resulted in dominance by shrubs. In a degraded Illinois

woodland, growing-season (May) burns were more effective than dormant-season (March) burns at controlling an
57

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

exotic species (Schwartz and Heim 1996). However, May

burns also caused different and longer lasting effects to the

native herbaceous understory than March burns, with

composition in the March (dormant season) burn plots

appearing more similar to the unburned control.

Soils

Consumption of surface and live fuels releases nutrients,

some of which may be leached from the system unless they

are taken back up by micro-organisms or growing vegetation. It is therefore believed that prescribed fire close to the

onset of growth or during the active season when growing

tissue is accumulating nutrients might lead to less leaching

from the system (Robbins and Myers 1992). Another possibility is that more nutrients could be volatilized when

actively growing tissues are burned than when tissues are

burned during the dormant season. By the time of the

dormant season, at least some of the nutrients from aboveground structures have already been translocated to underground storage structures and therefore escape being

volatilized (Robbins and Myers 1992). However, as

Robbins and Myers (1992) noted, very little data are

available to back up either the leaching or volatilization

theories. In longleaf pine forests, Boring et al. (2004)

documented greater nitrogen loss with growing-season

(June) burns than dormant-season (March—April) burns,

presumably as a result of live fuels being volatilized.

However, nitrogen fixation and atmospheric deposition

were believed sufficient to compensate for this loss if

the fire regime is not exclusively growing season—i.e.,

including a mix of seasons. There was no difference in

phosphorous with burning-season treatments (Boring et al.

2004). Temperatures were apparently not high enough for

any of the burns to volatilize this nutrient. Another recent

study reported very little effect of burning season on soil

variables in an oak-pine forest in Massachusetts (Neill et

al. 2007). The organic horizon (duff layer) was reduced

more by summer burns than by spring burns, and replacement with mineral soil caused the bulk density to also

be higher. All other variables including pH, acidity, base

saturation, total exchangeable cations, carbon, and

nitrogen did not differ between burn seasons.

Some other potential impacts of fire in different seasons on soils are likely associated with variation in fire

intensity or extent of soil exposure. Soil is exposed for

a longer period after burns in fall and winter (dormant

season), and this could alter the rate of erosion. In their

literature review, Robbins and Myers (1992) found only

a single study addressing erosion and season of burning.

Dobrowolski et al. (1987) reported greater sediment yields

after winter burns than spring and summer burns, attributing this to direct exposure of the soil to raindrops for a

longer period with winter burns. Summer burning produced the least erosion, possibly because these burns were

patchier. The lack of studies on erosion with prescribed fire

in the Eastern region may be due, in part, to the relative

lack of topography in many areas with active prescribed

burning programs.

Wildlife

Early forest managers generally avoided burning southeastern pine forests during the late spring and early summer, because of concerns about harming wildlife species.

However, with this time of year being the peak historical

fire season, others concluded that wildlife must have

evolved means to survive (Komarek 1965). Direct effects

to wildlife are perhaps less of a concern in the eastern

hardwood forest ecosystems because especially in the

north, fire historically occurred primarily during the

dormant season when many species are less active.

Birds—

Timing of early prescribed burning in the Southeastern

United States was strongly influenced by concerns about

game birds and other ground-nesting species (Stoddard

1931). Late winter to early spring burning became popular

because this period occurred after the end of hunting season but prior to nesting season for quail and other species

(Brennan et al. 1998). To reduce the feared catastrophic

effect on clutch success of ground-nesting species, burning at this time of year became “ingrained in the culture
58

GENERAL TECHNICAL REPORT PSW-GTR-224

of the Southeast” (Brennan et al. 1998). However, the

majority of studies have since shown few strong effects of

burn season on direct mortality, breeding success, or survival of birds (Cox and Widener 2008; Engstrom et al.

1996; Tucker et al. 2004, 2006). In fact, overwinter survival of Henslow’s sparrow (Ammodramus henslowii)

was found to be greater in areas previously burned in the

growing season than in areas previously burned in the

dormant season (Thatcher et al. 2006). In another study,

abundance of wintering bird communities did not differ

one year after burns conducted in the growing season

(April—August) or the dormant season (January—March)

(King et al. 1998).

Many bird species prefer to nest in stands that have

been burned within 1 or 2 years (Cox and Widener

2008). For example, the majority of Bachman’s sparrow

(Aimophila aestivalis) nests (>85 percent) were found in

areas that were recently burned during the growing season,

and the majority of wild turkey nests (62 percent) were

found in forest that had experienced a growing-season burn

within the past 2 years (Cox and Widener 2008, Sisson et

al. 1990). Management using a regime of growing-season

burns 3 or more years apart, but within a patchy landscape

with units varying in time since last fire, would therefore

likely impact relatively few ground nests.

Changes in vegetation brought about by burning in

different seasons can indirectly influence bird populations

(Engstrom 1993). Dormant-season burning in longleaf pine

forests can impact the structure and composition preferred

by different bird species by promoting hardwoods over

grasses and forbs (Abrahamson and Hartnett 1990). Redcockaded woodpecker (Picoides borealis) and other bird

species are generally less abundant in forests where understory hardwoods have encroached (Provencher et al. 2002,

Sparks et al. 1999). Although lengthening of fire intervals

is believed to be the main cause of red-cockaded woodpecker decline, growing-season burns have been shown to

more effectively suppress midstory hardwoods and promote

a ground cover composition favorable for arthropod food

sources for these birds (James et al. 1997). In a study of bird

community response to fire, Fitzgerald and Tanner (1992)

found that neither January nor June prescribed burns in a

dry prairie in south Florida altered bird species richness,

compared with the unburned control. Both of the burningseason treatments reduced shrub cover.

Ground cover is beneficial to some overwintering

migratory birds such as Henslow’s sparrow—burns in the

winter (February—March) eliminate this ground cover, and

research shows that growing-season burns improve survival

over dormant-season winter burns (Thatcher et al. 2006).

Spatial patchiness is another characteristic of fires potentially important for birds and other wildlife (Sparks et al.

1999), and spatial patchiness can differ among burning

seasons because of variation in fuel moisture.

Overall, reviews of the limited literature show few if

any effects of burning season on bird populations. Although growing-season burns may cause some direct

mortality by destroying nests and killing young birds,

many bird species renest, and the indirect benefits of

habitat alteration are usually far more important and likely

compensate or more than compensate for losses (Engstrom

et al. 2005, Robbins and Myers 1992).

Small mammals—

The effect of different prescribed burning seasons on

small mammal populations remains poorly studied. Both

historical fires and prescribed burns in eastern forests may

be of sufficiently low intensity and patchy enough that

the variable needs of small mammal populations are met,

regardless of burn season (Keyser and Ford 2006). A study

in oak stands in Virginia that compared effects of winter,

spring, and summer prescribed burns reported no detectable short-term losses of ground-dwelling species such

as shrews (Sorex and Blarina) and white-footed mice

(Peromyscus leucopus) (Keyser et al. 1996, 2001). Longer

term habitat changes, such as differences in hardwood

midstory cover or ground exposure owing to variation in

the burning season, could potentially affect small mammal

populations. Fires conducted in March or August annually

for 3 years in Florida longleaf pine sandhill forests resulted in no difference in pocket gopher (Geomys pinetis)
59

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

mounding or body size (Gates and Tanner 1988). Although a minor increase in herbaceous biomass was

noted following March burns, this apparently did not influence gopher behavior (Gates and Tanner 1988). Overall,

consistent trends in small mammal response to habitat

changes with burning season have not emerged (Ahlgren

1966, Brose and Van Lear 1999, Kirkland et al. 1996).

Amphibians and reptiles—

Direct effects of fire are not expected to be strong for

amphibian species occupying moist habitats that are less

flammable than the surrounding landscape. Prescribed

burns during cool weather in the winter, or any time of the

year when moisture is high, have a lower probability of

passing through and consuming fuels in wetter areas.

However, burns at these times of year may also coincide

with winter mating migrations when individuals can be

more vulnerable. Varying the burn season to include

growing-season burns as well as dormant-season burns has

been suggested as one means of reducing the potential

impact of fire (Schurbon and Fauth 2003).

It is possible that amphibian and reptile species are

indirectly influenced by burning season through differential effects on habitat structure. For example, certain

species of longleaf pine forests require bare sandy habitats

and are thus benefited most by relatively intense and

spatially variable burns (Russell et al. 1999), which are

more likely in some seasons than others. Growing-season

burns have been reported to be more intense and more

likely to clear overgrown vegetation surrounding wetlands;

some amphibians apparently prefer the higher light levels

and warmer temperatures that result (Bishop and Haas

2005). In a study by Yager et al. (2007), reduction of

midstory cover of longleaf pine forests through application

of a mixture of dormant-season and growing-season burns

increased habitat usage by gopher tortoises (Gopherus

polyphemus). Burn seasons most effective at reducing the

height of understory vegetation are likely to favor not only

the gopher tortoise but other species that utilize gopher

tortoise burrows such as the Florida pine snake (Pituophis

melanoleucus mugatis) and Florida gopher frog (Rana

capito aesopus).

Several studies have compared amphibian and reptile

populations after dormant- and growing-season prescribed

burns and none have found a significant difference in

numbers (Floyd et al. 2002, Keyser et al. 2004). The lack

of an effect can be attributed to incomplete consumption

of coarse woody debris and duff, the existence of moist

environments such as tunnels and cracks in the soil or

under rocks that escape heating, mobility of the organisms, relatively few changes to the overstory canopy, and

generally quick regrowth of understory vegetation (Renken

2006).

Arthropods—

Hall and Schweitzer 1992 (cited in Hermann et al. 1998)

hypothesized that burning during the growing season may

have fewer detrimental effects on arthropods than burning

during the dormant season because a greater number of

individuals have wings and are mobile at this time of year.

Arthropod abundance was found to be equal or greater

following growing-season burns than following dormantseason burns (Hermann et al. 1998). However, a fall survey

in Florida oak scrub found that garden orbweaver spider

(Argiope sp.) numbers were not affected by burns in

February, but were substantially reduced by burns in July

and August (Carrel 2008). Spiderlings disperse in April

and May through ballooning, so the low numbers immediately following summer fires may simply be due to lack of

dispersal opportunities between the time of the fire and the

time of sampling. In a hardwood stand in Kentucky, a

single March prescribed fire reduced the invertebrate mass

by 36 percent, with the majority of this loss occurring

among species associated with the forest floor (Kalisz and

Powell 2000).

Neither burns in July nor November altered the population size of the Karner Blue butterfly (Lycaeides melissa

samuelis) 1 to 3 years later, compared with unburned controls (King 2003). The July burns were during the period of

the second flight of the summer, whereas the November

burns occurred after activity had ceased for the year. Burns

in both seasons were described as “cool” (i.e., not at times

of the year when flame lengths are greatest), which may

have allowed some of the eggs on vegetation in this oak

savanna system to survive.
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GENERAL TECHNICAL REPORT PSW-GTR-224

Overall, it is apparent that how fire affects arthropods

will differ greatly by species and functional group, with

burning potentially most detrimental if the timing coincides with a particularly vulnerable life history stage

(Robbins and Myers 1992). Several authors have recommended that prescribed burning be done in such a way as

to maximize patchiness so that invertebrates are able to

survive in refugia and recolonize the burned areas (Kalisz

and Powell 2000, Knight and Holt 2005).

Implications for Managers

The majority of studies on burning-season effects in

eastern forest ecosystems have been conducted in pineand pine-oak-dominated forests of the Southeast. In this

forest type, the literature provides compelling evidence

that growing-season fire can lead to shifts in the plant community, relative to a regime of dormant-season fire. Repeated burns during the growing season (especially in

May, early in the growing season) curtail resprouting and

eventually suppress the less fire-resistant midstory hardwood vegetation more so than burns at other times of the

year. On the other hand, the pine overstory appears to be

minimally affected by burns in any season. This is particularly true for longleaf pine, a strongly fire-adapted species.

The end result is that repeated growing-season burning

leads to greater grass and herbaceous species abundance

and diversity under the pine canopy, whereas more shrubs

may be maintained with a regime of dormant-season

burning.

Key Points—Eastern Region

• There is little evidence that mortality or growth of southern pines differs after growing- or dormant-season

prescribed burns.

• Phenology does influence the response of midstory hardwoods in pine forests, with early-growing-season

(May) burns (coupled with short fire-return intervals) more likely to control or kill these species than

dormant-season burns. The result of early-growing-season burns is often an understory with greater cover of

grasses and forbs.

• Burning season has little effect on growth and mortality of overstory oak species, but higher intensity fire

(in whatever season fuels are sufficiently dry to burn at higher intensity) likely favors oaks over the long

term, by killing competing mesophytic species such as yellow poplar or maple.

• Although some understory plant species respond positively to fire in the growing season and others respond

positively to fire in the dormant season, the majority do not appear to be significantly affected by burning

season.

• Few strong direct impacts to wildlife from prescribed fire in any season have been documented; effects, both

positive and negative, appear to be mostly indirect, and primarily the result of fire-season-specific habitat

changes.

• Whether the ecosystem is burned or not (fire frequency) appears to play a stronger role in the response of

most species than the relatively minor effect caused by different burning seasons.

• Differences in fire effects among species suggests that a variable fire regime, including a mix of growingand dormant-season burns and different burn intensities may maximize biodiversity.
61

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

Early prescribed burning was often done during the

dormant season to avoid conflicts with wildlife reproduction, including bird nesting. However, recent research

generally does not show that burns in the growing season

affect bird populations more than burns at other times of

the year. In the few cases where differences in animal communities with varying burning seasons have been reported,

the mechanism is usually indirect, involving some alteration of understory structure. Understory vegetation of the

Southern United States grows so rapidly in the absence of

fire that the effect of burning or not (fire frequency) is generally much greater than the effect of burning season.

Both phenology and fire intensity appear to play a role

in determining fire effects in forests of the Eastern region,

with the outcome depending on the species and the differences in intensity between burn seasons. As with fire in

the Western and Central regions, phenology and intensity

are often confounded, making their relative contributions a

challenge to determine. Several of the more robust studies

of burning season concluded that for many species, fire

intensity plays a significant role in determining the outcome. Differences in intensity, if any, are often due to

higher ambient temperatures and greater use of heading

fires during the growing season. However, lower fuel moisture levels can also sometimes result in dormant-season

burns being more intense, particularly in hardwood forests

that lack the pyrogenic vegetation of the southeastern pine

zone.

The importance of phenology relative to fire intensity

in the Eastern region appears to be intermediate between

the Western region and the Central grasslands; this goes

along with apparent differences in the amount of fuel consumed between seasons, which are in most cases less than

differences among burning seasons in the Western region,

but greater than differences among burning seasons in the

Central grasslands. For some species, fire intensity may

override the effects of phenology at the time of the burn,

especially if the difference in fire intensity among seasons

is substantial.

Data from the many long-term burning studies conducted in the Southeastern United States indicate that

substantial changes likely require many burn cycles to

achieve. A single burn in any season generally does little

to alter plant or animal communities. Therefore one burn

or a few burns outside of normal season is/are unlikely

to have a major impact. In addition, the importance of

burning generally outweighs any effect of season of burning. Because prescribed burns are usually easier to conduct

during the dormant season than during the growing season/

lightning season, more acres may ultimately be treated by

employing a regime of both dormant- and growing-season

burns.

One key point mentioned repeatedly in the literature is

that a frequent yet heterogeneous fire regime, including a

range of fire seasons, may be necessary to sustain species

diversity, or even to maximally benefit individual species

where different parts of the life cycle are variably affected

by burning season. To mimic the variability inherent in the

historical fire regime, Robbins and Myers (1992) created a

table of random fire frequencies and seasons (within

specified ranges) for xeric to mesic longleaf pine habitats,

with a weighting so that two growing-season (May-June)

burns are conducted for each dormant-season burn. The

objective of such a table is to ensure that rigid burning

schedules, which would tend to favor some species over

others, are avoided. Also, occasional longer (8 to 10 years)

rest periods are incorporated that would allow seedlings of

certain species to become established (Robbins and Myers

1992).

Acknowledgments

This synthesis was funded by the Joint Fire Science

Program (07-S-08), and was inspired by observations

and results from the Fire and Fire Surrogate study site in

Sequoia National Park, where the ecological differences

between early summer and fall prescribed burns were

evaluated, and through conversations with many who

have been involved with season-of-burning research.
62

GENERAL TECHNICAL REPORT PSW-GTR-224

Steven Apfelbaum, Laura Calabrese, Walker

Chancellor, Todd Hamilton, Stephen Hudson, Quinn Long,

Ron Masters, and John McGuire kindly provided photographs. The authors thank Mark Daniels, Jennifer Gibson,

Jeff Glitzenstein, Jon Keeley, Deb Kennard, Quinn Long,

Ron Masters, Malcolm North, Dale Shippelhoute, Judith

Springer, Timothy Swedberg, and Gene Towne who

provided comments on a previous version of the manuscript (or sections of it), improving its clarity and application to managers and pointed us to publications we were

not aware of.

Metric Equivalents

When you know: Multiply by: To find:

Inches (in) 2.54 Centimeters

Square miles (mi2) 2.59 Square kilometers

Tons per acre 2.24 Tonnes or megagrams

 per hectare

Degrees Fahrenheit 0.55 Degrees Celsius

(F-32)

Literature Cited

Abrahamson, C.R.; Hartnett, D.C. 1990. Pine ecosystems.

In: Myers, R.L.; Ewel, J.J., eds. Ecosystems of Florida.

Orlando, FL: University of Central Florida Press:

103–149.

Abrams, M.D. 1992. Fire and the development of oak

forest. Bioscience. 42(5): 346–353.

Agee, J.K. 1993. Fire ecology of Pacific Northwest forests.

Covelo, CA: Island Press. 505 p.

Ahlgren, C.E. 1966. Small mammals and reforestation

following prescribed burning. Journal of Forestry. 64:

614–618.

Anderson, K.L.; Smith, E.F.; Owensby, C.L. 1970.

Burning bluestem range. Journal of Range Management. 23: 81–92.

Anderson, R.C. 1990. The historic role of fire in the North

American grassland. In: Collins, S.L.; Wallace, L.L.,

eds. Fire in North American tallgrass prairies. Norman,

OK: University of Oklahoma Press: 8–18.

Ansley, R.J.; Castellano, M.J. 2007a. Texas wintergrass

and buffalograss response to seasonal fires and clipping. Rangeland Ecology and Management. 60(2):

154–164.

Ansley, R.J.; Castellano, M.J. 2007b. Prickly pear cactus

responses to summer and winter fires. Rangeland

Ecology and Management. 60(3): 244–252.

Axelrod, D.I. 1985. Rise of the grassland biome, central

North America. Botanical Review. 51(2): 163–201.

Bailey, A.D.; Mickler, R.; Frost, C.C. 2007. Presettlement

fire regime and vegetation mapping in southeastern

coastal plain forest ecosystems. In: Butler, B.W.; Cook,

W., eds. The fire environment—innovations, management, and policy: proceedings of a conference. Fort

Collins, CO: U.S. Department of Agriculture, Forest

Service, Rocky Mountain Research Station: 275–286.

Bailey, R.G. 1980. Description of the ecoregions of the

United States. Misc. Publ. 1391. Washington, DC: U.S.

Department of Agriculture, Forest Service. 77 p.

Bailey, R.G. 1983. Delineation of ecosystem regions.

Environmental Management. 7(4): 365–373.

Baisan, C.H.; Swetnam, T.W. 1990. Fire history on a desert

mountain range: Rincon Mountain Wilderness,

Arizona, U.S.A. Canadian Journal of Forest Research.

20: 1559–1569.

Baker, R.G.; Bettis, E.A.I.; Schwert, D.P.; Horton, D.G.;

Chumbley, C.A.; Gonzalez, L.A.; Reagan, M.K. 1996.

Holocene paleoenvironments of northeast Iowa.

Holocene. 66(2): 203–234.

Bale, A.M.; Guyette, R.P.; Stambaugh, M.C. 2008.

Fire regime of a montane longleaf pine ecosystem,

Alabama. In: Kush, J.S.; Hermann, S.M., eds. Proceedings of the third montane longleaf conference. Auburn,

AL: Longleaf Alliance: 54–59.
63

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

Barden, L.S.; Woods, F.W. 1973. Characteristics of

lightning fires in Southern Appalachian forests. In:

Proceedings, 13th Tall Timbers fire ecology conference. Tallahassee, FL: Tall Timbers Research Station:

345–361.

Beckage, B.; Platt, W.J.; Slocum, M.G.; Panko, B. 2003.

Influence of the El Niño southern oscillation on fire

regimes in the Florida Everglades. Ecology. 84(12):

3124–3130.

Bekker, M.F.; Taylor, A.H. 2001. Gradient analysis of fire

regimes in montane forests of the southern Cascade

Range, Thousand Lakes Wilderness, California, USA.

Plant Ecology. 155: 15–28.

Benning, T.L.; Bragg, T.B. 1993. Response of big bluestem (Andropogon gerardii Vitman) to timing of spring

burns. American Midland Naturalist. 130(1): 127–132.

Beyers, J.L.; Wakeman, C.D. 2000. Season of burn effects

in southern California chaparral. In: Second interface

between ecology and land development in California.

Open-File Report 00-62. Sacramento, CA: U.S. Department of the Interior, Geological Survey: 45–55.

Bidwell, T.G.; Engle, D.M.; Claypool, P.L. 1990. Effects

of spring headfires and backfires on tallgrass prairie.

Journal of Range Management. 43(3): 209–212.

Biondini, M.E.; Steuter, A.A.; Grygiel, C.E. 1989. Seasonal fire effects on the diversity patterns, spatial

distribution, and community structure of forbs in

the northern mixed prairie. Vegetatio. 85: 21–31.

Bishop, D.C.; Haas, C.A. 2005. Burning trends and

potential negative effects of suppressing wetland

fires on flatwoods salamanders. Natural Areas Journal.

25(3): 290–294.

Biswell, H.H.; Taber, R.D.; Hedrick, D.W.; Schultz, A.M.

1952. Management of chamise brushlands for game in

the north coast region of California. California Fish

and Game. 38(4): 453–484.

Bond, W.J.; van Wilgen, B.W. 1996. Fire and plants.

London: Chapman and Hall. 272 p.

Boring, L.R.; Hedrick, J.J.; Wilson, C.A.; Mitchell, R.J.

2004. Season of burn and nutrient losses in a longleaf

pine ecosystem. International Journal of Wildand Fire.

13: 443–453.

Boyer, W.D. 1982. Growth of young longleaf pine as

affected by biennial burns plus chemical or mechanical

treatment of competition control. In: Proceedings, 2nd

biennial southern silvicultural research conference.

Gen. Tech. Rep. SE-24. Atlanta, GA: U.S. Department

of Agriculture, Forest Service: 62–65.

Boyer, W.D. 1987. Volume growth loss: a hidden cost of

periodic prescribed burning in longleaf pine? Southern

Journal of Applied Forestry. 11(3): 154–157.

Boyer, W.D. 1990. Growing season burns for control of

hardwoods in longleaf pine stands. Res. Pap. SO-256.

New Orleans, LA: U.S. Department of Agriculture, Forest Service, Southern Forest Experiment Station. 7 p.

Boyer, W.D. 1993. Eighteen years of seasonal burning in

longleaf pine: effects on overstory growth. In: Fire,

meteorology and the landscape: 12th conference on

fire and forest meteorology. Bethesda, MD: Society

of American Foresters: 602–610.

Boyer, W.D. 2000. Long-term effects of biennial prescribed

fires on the growth of longleaf pine. In: Moser, W.K.;

Moser, C.F., eds. Proceedings, 21st Tall Timbers fire ecology conference. Fire and forest ecology: innovative

silviculture and vegetation management. Tallahassee,

FL: Tall Timbers Research Station: 18–21.

Bradley, T.; Tueller, P. 2001. Effects of fire on bark beetle

presence on Jeffrey pine in the Lake Tahoe Basin. Forest Ecology and Management. 142(1-3): 205–214.

Bragg, T.B. 1982. Seasonal variations in fuel and fuel

consumption by fires in a bluestem prairie. Ecology.

63(1): 7–11.
64

GENERAL TECHNICAL REPORT PSW-GTR-224

Brennan, L.A.; Engstrom, R.T.; Palmer, W.E.; Hermann,

S.M.; Hurst, G.A.; Burger, L.W.; Hardy, C.L. 1998.

Whither wildlife without fire? In: Transactions of the

63rd North American wildland and natural resources

conference. Washington, DC: Wildlife Management

Institute: 402–414.

Brewer, J.S. 2006. Long-term population changes of

a fire-adapted plant subjected to different fire seasons.

Natural Areas Journal. 26(3): 267–273.

Brewer, J.S.; Cunningham, A.L.; Moore, T.P.; Brooks,

R.M.; Waldrup, J.L. 2009. A six-year study of firerelated flowering cues and coexistence of two perennial grasses in a wet longleaf pine (Pinus palustris)

savanna. Plant Ecology. 200: 141–154.

Brewer, J.S.; Platt, W.J. 1994a. Effects of fire season

and herbivory on reproductive success in a clonal

forb, Pityopsis graminifolia. Journal of Ecology. 82:

665–675.

Brewer, J.S.; Platt, W.J. 1994b. Effects of fire season and

soil fertility on clonal growth in a pyrophilic forb,

Pityopsis graminifolia (Asteraceae). American Journal

of Botany. 81(7): 805–814.

Brewer, J.S.; Platt, W.J.; Glitzenstein, J.S.; Streng, D.R.

1996. Effects of fire-generated gaps on growth and

reproduction of golden aster (Pityopsis graminifolia).

Bulletin of the Torrey Botanical Club. 123(4):

295–303.

Brockway, D.G.; Gatewood, R.G.; Paris, R.B. 2002.

Restoring fire and an ecological process in shortgrass

prairie ecosystems: initial effects of prescribed burning

during the dormant and growing seasons. Journal of

Environmental Management. 65: 135–152.

Brose, P.H.; Van Lear, D.H. 1998. Responses of hardwood

advance regeneration to seasonal prescribed fires in

oak-dominated shelterwood stands. Canadian Journal

of Forest Research. 28: 331–339.

Brose, P.; Van Lear, D. 1999. Effects of seasonal prescribed

fires on residual overstory trees in oak-dominated

shelterwood stands. Southern Journal of Applied

Forestry. 23(2): 88–93.

Brose, P.; Van Lear, D.; Cooper, R. 1999. Using

shelterwood harvests and prescribed fire to regenerate

oak stands on productive upland sites. Forest Ecology

and Management. 113: 125–141.

Brown, J.K. 2000. Introduction and fire regimes. In:

Brown, J.K.; Kapler Smith, J., eds. Wildland fire in ecosystems: effects of fire on flora. Gen. Tech. Rep. RMRSGTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research

Station: 1–7.

Bruce, D. 1954. Mortality of longleaf pine seedlings after

a winter fire. Journal of Forestry. 52(6): 442–443.

Bruce, D.; Bickford, C.A. 1950. Use of fire in natural regeneration of longleaf pine. Journal of Forestry. 48(2):

114–117.

Buckman, R.E. 1964. Effects of prescribed burning on

hazel in Minnesota. Ecology. 45(3): 626–629.

Busse, M.D.; Hubbert, K.R.; Fiddler, G.O.; Shestak, C.J.;

Powers, R.F. 2005. Lethal temperatures during burning

of masticated forest residues. International Journal of

Wildland Fire. 14: 267–276.

Byram, G.M. 1948. Vegetation temperature and fire

damage in the southern pines. Fire Control Notes. 9:

34–36.

Campbell, G.S.; Jungbauer, J.D.J.; Bristow, K.L.;

Hungerford, R.D. 1995. Soil temperature and water

content beneath a surface fire. Soil Science. 159(6):

363–374.

Carrel, J.E. 2008. The effect of season of fire on density

of female garden orbweavers (Araneae: Araneidae:

Argiopei) Florida scrub. Florida Entomologist. 91(2):

332–334.
65

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

Carrington, M.E.; Mullahey, J.J. 2006. Effects of burning

season and frequency on saw palmetto (Serenoa

repens) flowering and fruiting. Forest Ecology and

Management. 230: 69–78.

Chamrad, A.D.; Dodd, J.D. 1973. Prescribed burning

and grazing for prairie chicken habitat manipulation

in the Texas coastal prairie. Proceedings of the 12th

Tall Timbers fire ecology conference. Tallahassee, FL:

Tall Timbers Research Station: 257–276.

Chapman, H.H. 1932. Some further relations of fire to

longleaf pine. Journal of Forestry. 30: 602–604.

Chen, E.; Gerber, J.F. 1990. Climate. In: Myers, R.L.;

Ewel, J.J., eds. Ecosystems of Florida. Gainesville, FL:

University of Florida Press: 11–34.

Collins, S.L.; Wallace, L.L. 1990. Fire in North American

tallgrass prairies. Norman, OK: University of Oklahoma

Press. 176 p.

Copeland, T.E.; Sluis, W.; Howe, H.F. 2002. Fire season

and dominance in an Illinois tallgrass prairie restoration. Restoration Ecology. 10(2): 315–323.

Covington, W.W.; Sackett, S.S. 1992. Soil mineral nitrogen

changes following prescribed burning in ponderosa

pine. Forest Ecology and Management. 54: 175–191.

Cox, J.; Widener, B. 2008. Lightning-season burning:

friend or foe of breeding birds? Misc. Publ. 17.

Tallahassee, FL: Tall Timbers Research Station. 16 p.

D’Antonio, C.M. 2000. Fire, plant invasions, and global

changes. In: Mooney, H.A.; Hobbs, R.J., eds. Invasive

species in a changing world. Washington, DC: Island

Press: 65–93.

de Groot, W.J.; Wein, R.W. 2004. Effects of fire severity

and season of burn on Betula glandulosa growth

dynamics. International Journal of Wildland Fire. 13:

287–295.

DeBano, L.F.; Neary, D.G.; Ffolliott, P.F. 1998. Fire’s

effects on ecosystems. New York, NY: John Wiley &

Sons. 333 p.

DeWeese, G. 2007. Past fire regimes of table mountain pine

(Pinus pungens Lamb.) stands in the central Appalachian Mountains, Virginia, U.S.A. Knoxville, TN:

University of Tennessee. 308 p. Ph.D. dissertation.

DiTomaso, J.M.; Brooks, M.L.; Allen, E.B.; Minnich, R.;

Rice, P.M.; Kyser, G.B. 2006. Control of invasive

weeds with prescribed burning. Weed Technology.

20(2): 535–548.

DiTomaso, J.M.; Kyser, G.B.; Hastings, M.S. 1999.

Prescribed burning for control of yellow starthistle

(Centaurea solstitialis) and enhanced native plant

diversity. Weed Science. 47(2): 233–242.

Dobrowolski, J.P.; Blackburn, W.H.; Grelen, H.E. 1987.

Sediment production from long-term burning of a

longleaf pine-bluestem association. In: Pearson, H.A.;

Smeins, F.E.; Thill, R.E., eds. Ecological, physical, and

socioeconomic relationships within southern national

forests. Proceedings, southern evaluation project workshop. Gen. Tech. Rep. SO-68. New Orleans, LA: U.S.

Department of Agriculture, Forest Service, Southern

Experiment Station: 251–260.

Drewa, P.B. 2003. Effects of fire season and intensity on

Prosopis glandulosa Torr. var. glandulosa. International Journal of Wildland Fire. 12: 147–157.

Drewa, P.B.; Platt, W.J.; Moser, E.B. 2002. Fire effects on

resprouting of shrubs in headwaters of southeastern

longleaf pine savannas. Ecology. 83(3): 755–767.

Drewa, P.B.; Thaxton, J.M.; Platt, W.J. 2006. Responses

of root-crown bearing shrubs to differences in fire

regimes in Pinus palustris (longleaf pine) savannas:

exploring old-growth questions in second-growth

systems. Applied Vegetation Science. 9(1): 27–36.

Duchesne, L.C.; Hawkes, B.C. 2000. Fire in northern

ecosystems. In: Brown, J.K.; Smith, J.K., eds. Wildland

fire in ecosystems: effects of fire on flora. Gen. Tech.

Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain

Research Station: 35–51.
66

GENERAL TECHNICAL REPORT PSW-GTR-224

Ehrenreich, J.H.; Aikman, J.M. 1963. An ecological

study of the effect of certain management practices on

native prairie in Iowa. Ecological Monographs. 33(2):

113–130.

Elliott, K.J.; Vose, J.M.; Clinton, B.D.; Knoepp, J.D. 2004.

Effects of understory burning in a mixed mesic-oak

forest of the Southern Appalachians. In: Engstrom,

R.T.; Galley, K.M.; de Groot, W.J., eds. Fire in temperate, boreal, and montane ecosystems: Proceedings, 22nd

Tall Timbers fire ecology conference. Tallahassee, FL:

Tall Timbers Research Station: 272–283.

Emery, S.M.; Gross, K.L. 2005. Effects of timing of prescribed fire on the demography of an invasive plant,

spotted knapweed Centaurea maculosa. Journal of

Applied Ecology. 42(1): 60–69.

Engle, D.M.; Bidwell, T.G. 2001. Viewpoint: the response

of central North American prairies to seasonal fire.

Journal of Range Management. 54: 2–10.

Engle, D.M.; Bultsma, P.M. 1984. Burning of northern

mixed prairie during drought. Journal of Range

Management. 37: 398–401.

Engstrom, R.T. 1993. Characteristic mammals and birds

of longleaf pine forests. In: Hermann, S.M., ed. The

longleaf pine ecosystem ecology, restoration, and

management. Proceedings of the 18th Tall Timbers fire

ecology conference. Tallahassee, FL: Tall Timbers

Research Station: 127–138.

Engstrom, R.T.; McNair, D.B.; Brennan, L.A.; Hardy,

C.L.; Burger, L.W. 1996. Influence on birds of dormant versus lightning-season prescribed fire in longleaf pine forests: experimental design and preliminary

results. In: Wadsworth, K.G.; McCabe, R.E., eds.

Transactions of the 61st North American wildlife and

natural resources conference. Washington, DC: Wildlife Management Institute: 200–207.

Engstrom, R.T.; Vickery, P.D.; Perkins, D.W.; Shriver,

W.G. 2005. Effects of fire regime on birds in southeastern pine savannas and native prairies. Studies in Avian

Biology. 30: 147–160.

Erwin, W.J.; Stasiak, R.H. 1979. Vertebrate mortality

during the burning of reestablished prairie in Nebraska.

American Midland Naturalist. 101(1): 247–249.

Everett, R.G. 2008. Dendrochronology-based fire history

of mixed-conifer forests in the San Jacinto Mountains,

California. Forest Ecology and Management. 256(11):

1805–1814.

Ferguson, S.A.; Ruthford, J.E.; McKay, S.J.; Wright, D.;

Wright, C.; Ottmar, R. 2002. Measuring moisture

dynamics to predict fire severity in longleaf pine

forests. International Journal of Wildland Fire. 11(4):

267–279.

Ferrenberg, S.M.; Schwilk, D.W.; Knapp, E.E.; Groth, E.;

Keeley, J.E. 2006. Fire decreases arthropod abundance

but increases diversity: early and late season prescribed fire effects in a Sierra Nevada mixed-conifer

forest. Fire Ecology. 2(2): 79–102.

Fettig, C.; Shea, P.; Borys, R. 2004. Seasonal flight

patterns of four bark beetle species (Coleoptera:

Scolytidae) along a latitudinal gradient in California.

Pan-Pacific Entomologist. 80(1-4): 4–17.

Filip, G.M.; Yang-Erve, L. 1997. Effects of prescribed

burning on the viability of Armillaria ostoyae in

mixed-conifer forest soils in the Blue Mountains

of Oregon. Northwest Science. 71(2): 137–144.

Finch, D.M.; Ganey, J.L.; Yong, W.; Kimball, R.T.;

Sallabanks, R. 1997. Effects and interactions of fire,

logging, and grazing. In: Block, W.M.; Finch, D.M.,

tech. eds. Songbird ecology in southwestern ponderosa

pine forests: a literature review. Gen. Tech. Rep. RMGTR-292. Fort Collins, CO: U.S. Department of

Agriculture, Forest Service, Rocky Mountain Forest

and Range Experiment Station: 103–136.

Fitzgerald, S.M.; Tanner, G.W. 1992. Avian community

response to fire and mechanical shrub control in

south Florida. Journal of Range Management. 45(4):

396–400.
67

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

Flinn, M.A.; Wein, R.W. 1977. Depth of underground plant

organs and theoretical survival during fire. Canadian

Journal of Botany. 55: 2550–2554.

Floyd, T.M.; Russell, K.R.; Moorman, C.E.; Van Lear,

D.H.; Guynn, D.C., Jr.; Lanham, J.D. 2002. Effects of

prescribed fire on herpetofauna within hardwood forests of the upper piedmont of South Carolina: a preliminary analysis. In: Outcalt, K.W., ed. Proceedings,

11th biennial southern silvicultural research conference.

Gen. Tech. Rep. SRS-48. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southern Research

Station: 123–127.

Ford, P.L.; Johnson, G.V. 2006. Effects of dormant- vs.

growing-season fire in shortgrass steppe: biological

soil crust and perennial grass responses. Journal of Arid

Environments. 67: 1–14.

Foti, T.L.; Glenn, S.M. 1991. The Ouachita Mountain

landscape at the time of settlement. In: Henderson,

D.; Hedrick, L.D., eds. Proceedings, Conference on

restoring old-growth forests in the interior highlands

of Arkansas and Oklahoma. Morrilton, AR: Winrock

International Institute: 49–65.

Fowells, H.A. 1941. The period of seasonal growth of

ponderosa pine and associated species. Journal of

Forestry. 39: 601–608.

Fowler, C.; Konopik, E. 2007. The history of fire in the

Southern United States. Human Ecology Review.

14(2): 165–176.

Frandsen, W.F.; Ryan, K.C. 1986. Soil moisture reduces

belowground heat flux and soil temperatures under a

burning fuel pile. Canadian Journal of Forest Research.

16: 244–248.

Frost, C. 1993. Four centuries of changing landscape patterns in the longleaf pine ecosystem. In: Hermann,

S.M., ed. Longleaf pine ecosystem ecology, restoration, and management. Proceedings of the 18th Tall

Timbers fire ecology conference. Tallahassee, FL: Tall

Timbers Research Station: 17–44.

Frost, C.C. 1998. Presettleement fire frequency regimes of

the United States: a first approximation. In: Pruden,

T.L.; Brennan, L.A., eds. Fire in ecosystem management: shifting the paradigm from suppression to

prescription. Proceedings of the 20th Tall Timbers fire

ecology conference. Tallahassee, FL: Tall Timbers

Research Station: 70–81.

Fry, D.L.; Stephens, S.L. 2006. Influence of humans and

climate on the fire history of a ponderosa pine-mixed

conifer forest in the southeastern Klamath Mountains,

California. Forest Ecology and Management. 223:

428–438.

Fuhlendorf, S.D.; Harrell, W.C.; Engle, D.M.; Hamilton,

R.G.; Davis, C.A.; Leslie, D.M., Jr. 2006. Should

heterogeneity be the basis for conservation? Grassland

bird response to fire and grazing. Ecological Applications. 16(5): 1706–1716.

Fulé, P.Z.; Covington, W.W.; Moore, M.M. 1997. Determining reference conditions for ecosystem management of southwestern ponderosa pine forests. Ecological Applications. 7(3): 895–908.

Fulé, P.Z.; Denton, C.; Springer, J.D.; Kalies, E.L.; Egan,

D. 2007. Prescribed and wildland use fires in the Southwest: Do frequency and timing matter? Working Papers

in southwestern ponderosa pine forest restoration.

Flagstaff, AZ: Ecological Restoration Institute, Northern Arizona University. 8 p.

Fulé, P.Z.; Heinlein, T.A.; Covington, W.W.; Moore, M.M.

2003. Assessing fire regimes on Grand Canyon landscapes with fire-scar and fire-record data. International

Journal of Wildland Fire. 12(2): 129–145.

Garrison, G.A. 1972. Carbohydrate reserves and response

to use. In: McKell, C.M.; Blaisdell, J.P.; Goodin, J.R.,

eds. Wildland shrubs—their biology and utilization.

Proceedings of a symposium. Gen. Tech. Rep. GTRINT-1. Ogden, UT: U.S. Department of Agriculture,

Forest Service, Intermountain Forest and Range

Experiment Station: 271–278.
68

GENERAL TECHNICAL REPORT PSW-GTR-224

Gates, C.A.; Tanner, G.W. 1988. Effects of prescribed

burning on herbaceous vegetation and pocket gophers

(Geomys pinetis) in a sandhill community. Florida

Scientist. 51(3): 129–139.

Gibbens, R.P.; Schultz, A.M. 1963. Brush manipulation on

a deer winter range. California Fish and Game. 49(2):

95–118.

Glitzenstein, J.S.; Platt, W.J.; Streng, D.R. 1995a. Effects

of fire regime and habitat on tree dynamics in north

Florida longleaf pine savannas. Ecological Monographs. 65(4): 441–476.

Glitzenstein, J.S.; Streng, D.R.; Masters, R.E.; Platt, W.J.

2008. Clarifying long-term impacts of fire frequency

and fire season in southeastern Coastal Plain pine

savannas. In: Managing an ecosystem on the edge:

Proceedings, 6th eastern native grass symposium.

Columbia, SC: Clemson University. 16 p.

Glitzenstein, J.S.; Streng, D.R.; Platt, W.J. 1995b. Evaluating the effects of season of burn on vegetation in longleaf pine savannas. Nongame Wildlife Project Report.

Tallahassee, FL: Florida Game and Fresh Water Fish

Commission. 118 p.

Glitzenstein, J.S.; Streng, D.R.; Wade, D.D. 2003. Fire

frequency effects on longleaf pine vegetation in South

Carolina and Northeast Florida, USA. Natural Areas

Journal. 23(1): 22–37.

Grelen, H.E. 1978. May burns stimulate growth in longleaf pine seedlings. Res. Note SO-234. Asheville, NC:

U.S. Department of Agriculture, Forest Service, Southern Research Station. 5 p.

Grelen, H.E. 1983. May burning favors survival and early

height growth of longleaf pine Pinus palustris seedlings. Southern Journal of Applied Forestry. 7(1):

16–20.

Grelen, H.E.; Epps, E.A. 1967. Herbage responses to fire

and litter removal on southern blue-stem range. Journal

of Range Management. 20: 403–404.

Griffiths, A.D.; Christian, K.A. 1996. The effect of fire

on the frillneck lizard (Chlamydosaurus kingii) in

northern Australia. Australian Journal of Ecology.

21: 386–398.

Grissino-Mayer, H.D.; Swetnam, T.W. 2000. Centuryscale climate forcing of fire regimes in the American

Southwest. The Holocene. 10(2): 213–220.

Guyette, R.P.; Dey, D.C.; Stambaugh, M.C.; Muzika, R.

2006. Fire scars reveal variability and dynamics of

eastern fire regimes. In: Dickinson, M.B., ed. Fire in

eastern oak forests: delivering science to land managers. Gen. Tech. Rep. GTR-NRS-P-1. Columbus, OH:

U.S. Department of Agriculture, Forest Service, Northern Research Station: 20–39.

Guyette, R.P.; Spetich, M.A. 2003. Fire history of oak-pine

forests in the Lower Boston Mountains, Arkansas, USA.

Forest Ecology and Management. 180: 463–474.

Hamman, S.T.; Burke, I.C.; Knapp, E.E. 2008. Soil

nutrients and microbial activity after early and late

season prescribed burns in a Sierra Nevada mixed

conifer forest. Forest Ecology and Management.

256(3): 367–374.

Harper, M.G.; Dietrich, C.H.; Larimore, R.L.; Tessene,

P.A. 2000. Effects of prescribed fire on prairie

arthropods: an enclosure study. Natural Areas Journal.

20: 325–335.

Harrington, M.G. 1985. The effects of spring, summer, and

fall burning on gambel oak in a southwestern ponderosa pine stand. Forest Science. 31(1): 156–163.

Harrington, M.G. 1987. Ponderosa pine mortality from

spring, summer, and fall crown scorching. Western

Journal of Applied Forestry. 2(1): 14–16.

Harrington, M.G. 1989. Gambel oak root carbohydrate

response to spring, summer, and fall prescribed burning. Journal of Range Management. 42(6): 504–507.
69

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

Harrington, M.G. 1993. Predicting Pinus ponderosa

mortality from dormant season and growing season

fire injury. International Journal of Wildland Fire. 3(2):

65–72.

Harrod, R.J.; Halpern, C.B. 2009. Effects of experimental

burning on individual performance and population

structure of two rare plants of northcentral Washington.

Restoration Ecology 17(2): 215–225.

Hartford, R.A.; Frandsen, W.F. 1992. When it’s hot, it’s

hot... or maybe it’s not! (Surface flaming may not

portend extensive soil heating). International Journal

of Wildland Fire. 2(3): 139–144.

Hatten, J.A.; Zabowski, D.; Ogden, A.; Thies, W. 2008.

Soil organic matter in a ponderosa pine forest with

varying seasons and intervals of prescribed burn. Forest

Ecology and Management. 255: 2555–2565.

Henderson, J.P. 2006. Dendroclimatological analysis and

fire history of longleaf pine (Pinus palustris Mill.) in

the Atlantic and Gulf Coastal Plain. Knoxville, TN:

University of Tennessee. 197 p. Ph.D. dissertation.

Hermann, S.M.; Van Hook, T.; Flowers, R.W.; Brennan,

L.A.; Glitzenstein, J.S.; Streng, D.R.; Walker, J.L.;

Myers, R.L. 1998. Fire and biodiversity: studies of

vegetation and arthopods. In: Transactions of the 63rd

North American wildland and natural resources conference. Washington, DC: Wildlife Management

Institute: 384–401.

Heyerdahl, E.K.; Brubaker, L.B.; Agee, J.K. 2001. Spatial

controls of historical fire regimes: a multiscale example

from the interior West, USA. Ecology. 82(3): 660–678.

Heyerdahl, E.K.; Miller, R.F.; Parsons, R.A. 2006. History

of fire and Douglas-fir establishment in a savanna and

sagebrush-grassland mosaic, southwestern Montana,

USA. Forest Ecology and Management. 230: 107–118.

Hierro, J.L.; Menges, E.S. 2002. Fire intensity and shrub

regeneration in palmetto-dominated flatwoods of

central Florida. Florida Scientist. 65(1): 51–61.

Hiers, J.K.; Wyatt, R.; Mitchell, R.J. 2000. The effects of

fire regime on legume reproduction in longleaf pine

savannas: Is a season selective? Oecologia. 125(4):

521–530.

Higgins, K.F. 1984. Lightning fires in North Dakota

grasslands and in pine-savanna lands of South Dakota

and Montana. Journal of Range Management. 37(2):

100–103.

Higgins, K.F. 1986a. A comparison of burn season effects

on nesting birds in North Dakota mixed-grass prairie.

Prairie Naturalist. 18(4): 219–228.

Higgins, K.F. 1986b. Interpretation and compendium of

historical fire accounts in the northern Great Plains.

Resource Publ. 161. Washington, DC: U.S. Department

of the Interior, Fish and Wildlife Service. 39 p.

Hough, W.A. 1968. Carbohydrate reserves of saw-palmetto:

seasonal variation and effects of burning. Forest

Science. 14(4): 399–405.

Hough, W.A. 1973. Fuel and weather influence wildfires in

sand pine forests. Res. Pap. SE-106. Asheville, NC: U.S.

Department of Agriculture, Forest Service, Southeast

Forest Experiment Station. 9 p.

Hough, W.A.; Albini, F.A. 1978. Predicting fire behavior

in palmetto-gallberry fuel complexes. Res. Pap. SE174. Asheville, NC: U.S. Department of Agriculture,

Forest Service, Southeastern Forest Experimental

Station. 44 p.

Howe, H.F. 1994a. Managing species diversity in tallgrass

prairie: assumptions and implications. Conservation

Biology. 8(3): 691–704.

Howe, H.F. 1994b. Response of early- and late-flowering

plants to fire season in experimental prairies. Ecological Applications. 4(1): 121–133.

Howe, H.F. 1995. Succession and fire season in experimental prairie plantings. Ecology. 76(6): 1917–1925.
70

GENERAL TECHNICAL REPORT PSW-GTR-224

Howe, H.F. 1999. Response of Zizia aurea to seasonal

mowing and fire in a restored prairie. American Midland Naturalist. 141: 373–380.

Howe, H.F. 2000. Grass response to seasonal burns in

experimental plantings. Journal of Range Management. 53: 437–441.

Huff, M.H.; Smith, J.K. 2000. Fire effects on animal

communities. In: Smith, J.K., ed. Wildland fire in

ecosystems: effects of fire on fauna. Gen. Tech. Rep.

RMRS-GTR-42-vol. 1. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain

Research Station: 35–42.

Huffman, J. 2006. Historical fire regimes in southeastern

pine savannas. Baton Rouge, LA: Louisiana State

University. 77 p. Ph.D. dissertation.

Huffman, J.M.; Platt, W.J.; Grissino-Mayer, H.; Boyce,

C.J. 2004. Fire history of a barrier island slash pine

(Pinus elliottii) savanna. Natural Areas Journal. 24(3):

258–268.

Hulbert, L.C. 1969. Fire and litter effects in undisturbed

bluestem prairie in Kansas. Ecology. 50(5): 874–877.

Hulbert, L.C. 1988. Causes of fire effects in tallgrass

prairie. Ecology. 69(1): 46–58.

Jacqmain, E.I.; Jones, R.H.; Mitchell, R.J. 1999. Influences of frequent cool-season burning across a soil

moisture gradient on oak community structure in

longleaf pine ecosystems. American Midland Naturalist. 141(1): 85–100.

James, F.C.; Hess, C.A.; Kufrin, D. 1997. Species-centered

environmental analysis: indirect effects of fire history

on red-cockaded woodpeckers. Ecological Applications. 7(1): 118–129.

Johansen, M.P.; Hakonson, T.E.; Breshears, D.D. 2001.

Post-fire runoff and erosion from rainfall simulations:

contrasting forests with shrublands and grasslands.

Hydrological Processes. 15: 2953–2965.

Johnson, E.A. 1992. Fire and vegetation dynamics: studies

from the North American boreal forest. Cambridge,

United Kingdom: Cambridge University Press. 144 p.

Johnson, S.D.; Horn, K.C.; Savage, A.M.; Windhager,

S.; Simmons, M.T.; Rudgers, J.T. 2008. Timing of

prescribed burns affects abundance and composition

of arthropods in the Texas Hill Country. The Southwestern Naturalist. 53(2): 137–145.

Jones, J.R.; DeByle, N.V. 1985. Fire. In: DeByle, N.;

Winokur, R.P., eds. Aspen: Ecology and management

in the Western United States. Gen. Tech. Rep. RM-119.

Fort Collins, CO: U.S. Department of Agriculture,

Forest Service, Rocky Mountain Forest and Range

Experiment Station: 77–81.

Jones, M.B.; Laude, H.M. 1960. Relationship between

sprouting in chamise and the physiological condition of the plant. Journal of Range Management. 13:

210–214.

Kalisz, P.J.; Powell, J.E. 2000. Effects of prescribed fire on

soil invertebrates in upland forests on the Cumberland

Plateau of Kentucky, USA. Natural Areas Journal.

20(1): 336–341.

Kansas Natural Heritage Inventory. 2007. Native prairie

hay meadows: a landowner’s management guide.

Lawrence, KS: University of Kansas. 32 p.

Kauffman, J.B. 1990. Ecological relationships of vegetation and fire in Pacific Northwest forests. In: Walsted,

J.D.; Radosevich, S.R.; Sandberg, D.V., eds. Natural and

prescribed fire in Pacific Northwest forests. Corvallis,

OR: Oregon State University Press: 39–52.

Kauffman, J.B.; Martin, R.E. 1989. Fire behavior, fuel

consumption, and forest-floor changes following

prescribed understory fires in Sierra Nevada mixed

conifer forests. Canadian Journal of Forest Research.

19: 455–462.
71

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

Kauffman, J.B.; Martin, R.E. 1990. Sprouting shrub

response to different seasons and fuel consumption

levels of prescribed fire in Sierra Nevada mixed conifer

ecosystems. Forest Science. 36(3): 748–764.

Kauffman, J.B.; Martin, R.E. 1991. Factors influencing

the scarification and germination of three montane

Sierra Nevada shrubs. Northwest Science. 65(4):

180–187.

Kaufman, D.W.; Finck, E.J.; Kaufman, G.A. 1990. Small

mammals and grassland fires. In: Collins, S.L.; Wallace,

L.L., eds. Fire in North American tallgrass prairies.

Norman, OK: University of Oklahoma Press: 46–80.

Kaufman, D.W.; Gurtz, S.K.; Kaufman, G.A. 1988. Movements of deer mouse in response to prairie fire. Prairie

Naturalist. 20(4): 225–229.

Keeley, J.E. 1987. Role of fire in seed germination of

woody taxa in California chaparral. Ecology. 68(2):

434–443.

Keeley, J.E. 2002. Fire management of California

shrubland landscapes. Environmental Management.

29(3): 395–408.

Keeley, J.E. 2006. South coast bioregion. In: Sugihara,

N.G.; van Wagtendonk, J.W.; Shaffer, K.E.; FitesKaufman, J.; Thode, A.E., eds. Fire in California’s

ecosystems. Berkeley, CA: University of California

Press: 350–414.

Keeley, J.E.; Fotheringham, C.J. 2001. History and management of crown-fire ecosystems: a summary and

response. Conservation Biology. 15(6): 1561–1567.

Kerns, B.K.; Thies, W.G.; Niwa, C.G. 2006. Season and

severity of prescribed burn in ponderosa pine forests:

implications for understory native and exotic plants.

Ecoscience. 13(1): 44–55.

Keyser, P.D.; Brose, P.H.; Van Lear, D.H. 1996. Enhancing oak regeneration with fire in shelterwood stands:

preliminary trials. In: Wadsworth, K.G.; McCabe, R.E.,

eds. Transactions of the 61st North American wildlife

and natural resources conference. Washington, DC:

Wildlife Management Institute: 215–219.

Keyser, P.D.; Ford, W.M. 2006. Influence of fire on mammals in eastern oak forests. In: Dickinson, M.B., ed.

Fire in eastern oak forests: delivering science to land

managers. Gen. Tech. Rep. GTR-NRS-P-1. Newton

Square, PA: U.S. Department of Agriculture, Forest

Service, Northern Research Station: 180–190.

Keyser, P.D.; Sausville, D.J.; Ford, W.M.; Mengak, M.J.;

Brose, P.H.; Van Lear, D.H. 2001. Fire impacts to

small mammals in Piedmont oak shelterwoods. In:

Proceedings of the annual conference of the Southeastern Association of Fish and Wildlife Agencies.

55: 375–381.

Keyser, P.D.; Sausville, D.J.; Ford, W.M.; Schwab, D.J.;

Brose, P. 2004. Prescribed fire impacts to amphibians

and reptiles in shelterwood-harvested oak-dominated

forests. Virginia Journal of Science. 55(4): 159–168.

King, R.S. 2003. Habitat management for the Karner blue

butterfly (Lycaeides melissa samuelis): evaluating the

short-term consequences. Ecological Restoration. 21:

101–106.

King, T.G.; Howell, M.A.; Chapman, B.R.; Miller, K.V.;

Schorr, R.A. 1998. Comparisons of wintering bird

communities in mature pine stands managed by

prescribed burning. The Wilson Bulletin. 110(4):

570–574.

Kirkland, G.L.; Snoddy, H.W.; Amsler, T.L. 1996. Impact

of fire on small mammals and amphibians in a central

Appalachian deciduous forest. American Midland

Naturalist. 135: 253–260.

Kirkman, L.K.; Drew, M.B.; Edwards, D. 1998. Effects of

experimental fire regimes on the population dynamics

of Schwalbea americana L. Plant Ecology. 137(1):

115–137.
72

GENERAL TECHNICAL REPORT PSW-GTR-224

Kirkpatrick, C.; Conway, C.J.; Jones, P.B. 2006. Distribution and relative abundance of forest birds in relation

to burn severity in southeastern Arizona. Journal of

Wildlife Management. 70(4): 1005–1012.

Klebenow, D.A.; Bruner, A.D. 1977. Determining factors

necessary for prescribed burning. In: Busby, F.E.;

Storey, E., eds. Use of prescribed burning in western

woodland and range ecosystems: a symposium. Logan,

UT: Utah Agricultural Experiment Station, Utah State

University: 69–74.

Klopatek, C.C.; DeBano, L.F.; Klopatek, J.M. 1988.

Effects of simulated fire on vesicular-arbuscular

mycorrhizae in pinyon-juniper woodland soil. Plant

and Soil. 109: 245–249.

Klopatek, J.M.; Klopatek, C.C.; DeBano, L.F. 1990.

Potential variation of nitrogen transformations in

pinyon-juniper ecosystems resulting from burning.

Biology and Fertility of Soils. 10: 35–44.

Knapp, A.K. 1984. Post-burn differences in solar radiation,

leaf temperature, and water stress influencing production in a lowland prairie. American Journal of Botany.

71(2): 220–227.

Knapp, E.E.; Keeley, J.E. 2006. Heterogeneity in fire

severity within early season and late season prescribed

burns in a mixed-conifer forest. International Journal of

Wildland Fire. 15: 37–45.

Knapp, E.E.; Keeley, J.E.; Ballenger, E.A.; Brennan, T.J.

2005. Fuel reduction and coarse woody debris dynamics with early season and late season prescribed fires in

a Sierra Nevada mixed conifer forest. Forest Ecology

and Management. 208: 383–397.

Knapp, E.E.; Schwilk, D.W.; Kane, J.M.; Keeley, J.E.

2007. Role of burning season on initial understory

vegetation response to prescribed fire in a mixed conifer forest. Canadian Journal of Forest Research. 37:

11–22.

Knight, T.M.; Holt, R.D. 2005. Fire generates spatial

gradients in herbivory: an example from a Florida

sandhill ecosystem. Ecology. 86(3): 587–593.

Knutson, H.; Campbell, J.B. 1976. Relationships of

grasshoppers (Acrididae) to burning, grazing, and

range sites on native tallgrass prairie in Kansas. In:

Proceedings of the Tall Timbers conference of ecology

animal control by habitat management. Tallahassee,

FL: Tall Timbers Research Station. 6: 107–120.

Komarek, E.V. 1964. The natural history of lightning. In:

Komarek, E.V., ed. Proceedings, 3rd Tall Timbers fire

ecology conference. Tallahassee, FL: Tall Timbers

Research Station: 139–183.

Komarek, E.V. 1965. Fire ecology—grasslands and man.

In: Komarek, E.V., ed. Proceedings, 4th Tall Timbers fire

ecology conference. Tallahassee, FL: Tall Timbers

Research Station: 169–220.

Komarek, E.V. 1967. The nature of lightning fire. In:

Komarek, E.V., ed. Proceedings, 7th Tall Timbers fire

ecology conference. Tallahassee, FL: Tall Timbers

Research Station: 5–41.

Komarek, E.V. 1968. Lightning and lightning fires as

ecological forces. In: Komarek, E.V., ed. Proceedings,

8

th Tall Timbers fire ecology conference. Tallahassee,

FL: Tall Timbers Research Station: 169–197.

Kramer, P.; Wetmore, T. 1943. Effects of defoliation on

cold resistance and diameter growth of broad-leaved

evergreens. American Journal of Botany. 30: 428–431.

Laughlin, D.C.; Bakker, J.D.; Daniels, M.L.; Moore,

M.M.; Casey, C.A.; Springer, J.D. 2008. Restoring

plant species diversity and community composition

in a ponderosa pine-bunchgrass ecosystem. Plant

Ecology. 197(1): 139–151.

Le Fer, D.; Parker, V.T. 2005. The effect of seasonality of

burn on seed germination in chaparral: the role of soil

moisture. Madroño. 52(3): 166–174.
73

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

Lesica, P. 1999. Effects of fire on the demography of the

endangered, geophytic herb Silene spaldingii

(Caryophyllaceae). American Journal of Botany. 86(7):

996–1002.

Lesica, P.; Martin, B. 2003. Effects of prescribed fire and

season of burn on recruitment of the invasive exotic

plant, Potentilla recta, in a semiarid grassland. Restoration Ecology. 11(4): 516–523.

Lewis, C.E.; Grelen, H.E.; Probasco, G.E. 1982. Prescribed burning in southern forest and rangeland

improves forage and its use. Southern Journal of

Applied Forestry. 6: 19–25.

Lewis, C.E.; Harshbarger, T.J. 1976. Shrub and herbaceous vegetation after 20 years of prescribed burning

in the South Carolina Coastal Plain. Journal of Range

Management. 29: 13–18.

Lewis, H.T. 1973. Patterns of Indian burning in California:

ecology and enthnohistory. Anthropological Paper

No. 1. Ramona, CA: Ballena Press. 101 p.

Liu, H.; Menges, E.S. 2005. Winter fires promote greater

vital rates in the Florida Keys than summer fires.

Ecology. 86(6): 1483–1495.

Liu, H.; Menges, E.S.; Quintana-Ascencio, P.F. 2005.

Population viability analysis of Chamaecrista

keyensis: effects of fire season and frequency. Ecological Applications 15(1): 210–221.

Lyon, L.J.; Huff, M.H.; Telfer, E.S.; Schreiner, D.S.;

Smith, J.K. 2000a. Fire effects on animal populations.

In: Smith, J.K., ed. Wildland fire in ecosystems: effects

of fire on fauna. Gen. Tech. Rep. RMRS-GTR-42-vol. 1.

Ogden, UT: U.S. Department of Agriculture, Forest

Service, Rocky Mountain Research Station: 25–34

Lyon, L.J.; Telfer, E.S.; Schreiner, D.S. 2000b. Direct

effects of fire and animal responses. In: Smith, J.K., ed.

Wildland fire in ecosystems: effects of fire on fauna.

Gen. Tech. Rep. RMRS-GTR-42-vol. 1. Ogden, UT:

U.S. Department of Agriculture, Forest Service, Rocky

Mountain Research Station: 17–23.

MacDonald, N.W.; Scull, B.T.; Abella, S.R. 2007. Midspring burning reduces spotted knapweed and increases native grasses during a Michigan experimental

grassland establishment. Restoration Ecology. 15(1):

118–128.

Main, M.B.; Barry, M.J. 2002. Influence of season of fire

on flowering of wet prairie grasses in south Florida,

USA. Wetlands. 22(2): 430–434.

Martin, R.E.; Sapsis, D.B. 1992. Fires as agents of biodiversity: pyrodiversity promotes biodiversity. In:

Kerner, H.M., ed. Proceedings, Symposium on biodiversity in northwestern California. Report 29. Santa

Rosa, CA: Wildland Resources Center, University of

California: 150–157.

Masters, R.E.; Skeen, J.E.; Whitehead, J. 1995. Preliminary fire history of McCurtain County Wilderness Area

and implications for red-cockaded woodpecker management. In: Kulhavy, D.L.; Hooper, R.G.; Costa, R.,

eds. Red cockaded woodpecker species recovery,

ecology, and management. Nagadoches, TX: School

of Forestry, Center for Applied Studies, Stephen F.

Austin State University: 240–232.

McCandliss, D.S. 2002. Prescribed burning in the Kings

River Ecosystem Project Area: lessons learned. In:

Verner, J., ed. Proceedings of a symposium on the

Kings River sustainable forest ecosystems project:

progress and current status. Gen. Tech. Rep. PSW-GTR183. Albany, CA: U.S. Department of Agriculture,

Forest Service, Pacific Southwest Research Station:

37–45.

McClain, W.E.; Elzinga, S.L. 1994. The occurrence of

prairie and forest fires in Illinois and other Midwestern

States 1679-1854. Eriginia. 13: 79–90.

McHugh, C.W.; Kolb, T.E. 2003. Ponderosa pine mortality

following fire in northern Arizona. International

Journal of Wildland Fire. 12: 7–22.
74

GENERAL TECHNICAL REPORT PSW-GTR-224

McHugh, C.W.; Kolb, T.E.; Wilson, J.L. 2003. Bark beetle

attacks on ponderosa pine following fire in northern

Arizona. Environmental Entomology. 32(3): 510–522.

Menges, E.S.; Deyrup, M.A. 2001. Postfire survival in

south Florida slash pine: interacting effects of fire

intensity, fire season, vegetation, burn size, and bark

beetles. International Journal of Wildland Fire. 10:

53–63.

Meyer, M.D.; Schiffman, P.M. 1999. Fire season and

mulch reduction in a California grassland: a comparison of restoration strategies. Madroño. 46(1): 25–37.

Monroe, M.E.; Converse, S.J. 2006. The effects of early

season and late season prescribed fires on small mammals in a Sierra Nevada mixed conifer forest. Forest

Ecology and Management. 236: 229–240.

Monsanto, P.G.; Agee, J.K. 2008. Long-term post-wildfire

dynamics of coarse woody debris after salvage logging

and implications for soil heating in dry forests of the

eastern Cascades, Washington. Forest Ecology and

Management. 255: 3952–3961.

Moody, T.J.; Fites-Kaufmann, J.; Stephens, S.L. 2006.

Fire history and climate influences from forests in

the northern Sierra Nevada, USA. Fire Ecology. 2(1):

115–141.

Mulligan, M.K.; Kirkman, L.K.; Mitchell, R.J. 2002.

Aristida beyrichiana (wiregrass) establishment and

recruitment: implications for restoration. Restoration

Ecology. 10(1): 68–76.

Myers, R.L. 1990. Scrub and high pine. In: Myers, R.L.;

Ewel, J.J., eds. Ecosystems of Florida. Orlando, FL:

University of Central Florida Press: 150–193.

Myers, R.L. 2000. Fire in tropical and subtropical ecosystems. In: Brown, J.K.; Smith, J.K., eds. Wildland fire in

ecosystems: effects of fire on flora. Gen. Tech. Rep.

RMRS-GTR-42-vol. 2. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain

Research Station: 161–173.

Myers, R.L.; White, D.L. 1987. Landscape history and

changes in sandhill vegetation in north-central and

south-central Florida. Bulletin of the Torrey Botanical

Club. 114(1): 21–32.

National Oceanic and Atmospheric Administration.

2009. Data from the U.S. National Lightning Detection

Network 1996–2000. http://www.crh.noaa.gov/Image/

pub/ltg2/usa_ltg_fdm.gif. (June 10, 2009).

Neary, D.G.; Klopatek, C.C.; DeBano, L.F.; Ffolliott, P.F.

1999. Fire effects on belowground sustainability: a

review and synthesis. Forest Ecology and Management. 122: 51–71.

Negron-Ortiz, V.; Gorchov, D.L. 2000. Effects of fire

season and post-fire herbivory on the cycad Zamia

pumila (Zamiaceae) in slash pine savanna, Everglades

National Park, Florida. International Journal of Plant

Science. 161: 659–669.

Neill, C.; Patterson, W.A., III; Crary, D.W., Jr. 2007.

Responses of soil carbon, nitrogen and cations to the

frequency and seasonality of prescribed burning in a

Cape Cod oak-pine forest. Forest Ecology and Management. 250: 234–243.

Niwa, C.G.; Peck, R.W. 2002. Influence of prescribed fire

on carabid beetle (Carabidae) and spider (Araneae)

assemblages in forest litter in southwestern Oregon.

Environmental Entomology. 31(5): 785–796.

Norman, S.P.; Taylor, A.H. 2003. Tropical and north

Pacific teleconnections influence fire regimes in pinedominated forests of north-eastern California, USA.

Journal of Biogeography. 30(7): 1081–1092.

Nowacki, G.; Abrams, M.D. 2008. The demise of fire and

“mesophication” of forests in the Eastern United

States. Bioscience. 58(2): 123–138.

Odion, D.C. 2000. Seed banks of long-unburned stands

of maritime chaparral: composition, germination

behavior, and survival without fire. Madroño. 47(3):

195–203.
75

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

Ortloff, W. 1996. Wood-anatomical evidence of fire

seasonality. In: Dean, J.S.; Meko, D.M.; Swetnam, T.M.,

eds. Tree rings, environment, and humanity. Radiocarbon. Tucson, AZ: University of Arizona: 89–93.

Outcalt, K.W. 1994. Seed production of wiregrass in

central Florida following growing season prescribed

burns. International Journal of Wildland Fire. 4(1):

123–125.

Outcalt, K.W. 2008. Lightning, fire and longleaf pine:

using natural disturbance to guide management. Forest

Ecology and Management. 255: 3351–3359.

Owens, M.K.; Mackley, J.W.; Carroll, C.J. 2002. Vegetation dynamics following seasonal fires in mixed

mesquite/acacia savannas. Journal of Range Management. 55: 509–516.

Palik, B.J.; Mitchell, R.J.; Hiers, J.K. 2002. Modeling

silviculture after natural disturbance to sustain

biodiversity in the longleaf pine (Pinus palustris)

ecosystem: balancing complexity and implementation.

Forest Ecology and Management. 155(1/3): 347–356.

Parker, V.T. 1987a. Can native flora survive prescribed

burns? Fremontia. 15(2): 3–6.

Parker, V.T. 1987b. Effects of wet-season management

burns on chaparral vegetation: implications for rare

species. In: Proceedings of a California conference on

the conservation and management of rare and endangered plants. Sacramento, CA: California Native Plant

Society: 233–237.

Parsons, D.J.; Stohlgren, T.J. 1989. Effects of varying fire

regimes on annual grasslands in the southern Sierra

Nevada of California. Madroño. 36(3): 154–168.

Perrakis, D.D.B.; Agee, J.K. 2006. Seasonal fire effects on

mixed-conifer forest structure and ponderosa pine resin

properties. Canadian Journal of Forest Research. 36:

238–254.

Petersen, S.M.; Drewa, P.B. 2006. Did lightning-initiated

growing season fires characterize oak dominated

ecosystems of southern Ohio? The Journal of the

Torrey Botanical Society. 133(2): 217–224.

Pilliod, D.S.; Bury, R.B.; Hyde, E.J.; Pearl, C.A.; Corn,

P.S. 2003. Fire and amphibians in North America.

Forest Ecology and Management. 178: 163–181.

Platt, W.J.; Evans, G.W.; Davis, M.M. 1988. Effects of fire

season on flowering of forbs and shrubs in longleaf

pine forests. Oecologia. 76: 353–363.

Platt, W.J.; Glitzenstein, J.S.; Streng, D.R. 1991. Evaluating pyrogenicity and its effects on vegetation in

longleaf pine savannas. In: Hermann, S.M., ed. Highintensity fire in wildlands: management challenges

and options. Proceedings, 17th Tall Timbers Fire

Ecology Conference. Tallahassee, FL: Tall Timbers

Research Station: 143–162.

Pollak, O.; Kan, T. 1998. The use of prescribed fire to

control invasive exotic weeds at Jepson Prairie Preserve. In: Witham, C.W.; Bauder, E.T.; Belk, D.; Ferren,

W.R., Jr.; Ornduff, R., eds. Ecology, conservation, and

management of vernal pool ecosystems. Sacramento,

CA: California Native Plant Society: 241–249.

Provencher, L.; Gobris, N.M.; Brennan, L.A. 2002.

Effects of hardwood reduction on winter birds in northwest Florida longleaf pine sandhill forests. The Auk.

119(1): 71–87.

Radosevich, S.R.; Conard, S.G. 1980. Physiological

control of chamise shoot growth after fire. American

Journal of Botany. 67(10): 1442–1447.

Reinking, D.L. 2005. Fire regimes and avian responses in

the central tallgrass prairie. Studies in Avian Biology.

30: 116–126.
76

GENERAL TECHNICAL REPORT PSW-GTR-224

Renken, R. 2006. Does fire affect amphibians and reptiles

in Eastern U.S. oak forests? In: Dickinson, M.B., ed.

Proceedings, Conference on fire in eastern oak forests:

delivering science to land managers. Gen. Tech. Rep.

NRS-P-1. Columbus, OH: U.S. Department of Agriculture, Forest Service, Northern Research Station:

158–166.

Rideout, S.; Rickard, J.K.; Wade, D.D. 2003. Preliminary

response of herbaceous plants to biennial cycles

applied at different dates during the growing season.

Natural Areas Journal. 23(1): 38–42.

Robbins, L.E.; Myers, R.L. 1992. Seasonal effects of

prescribed burning in Florida: a review. Misc. Publ. 8.

Tallahassee, FL: Tall Timbers Research Station. 97 p.

Rundel, P.W.; Baker, G.A.; Parsons, D.J. 1981. Productivity and nutritional responses of Chamaebatia foliolosa

(Rosaceae) to seasonal burning. In: Margaris, N.S.;

Mooney, H.A., eds. Components of productivity of

mediterranean-climate regions—basic and applied

aspects. The Hague/Boston/London: Dr. W. Junk:

191–196.

Russell, K.R.; Van Lear, D.H.; Guynn, D.C., Jr. 1999.

Prescribed fire effects on herpetofauna: review and

management implications. Wildlife Society Bulletin.

27(2): 374–384.

Ruthven, D.C.; Braden, A.W.; Knutson, H.J.; Gallagher,

J.F.; Synatzske, D.R. 2003. Woody vegetation response

to various burning regimes in South Texas. Journal of

Range Management. 56: 159–166.

Ruthven, D.C., III; Kazmaier, R.T.; Janis, M.W. 2008.

Short-term response of herpetofauna to various burning

regimes in the south Texas plains. The Southwestern

Naturalist. 53(4): 480–487.

Ryan, K.C.; Peterson, D.L.; Reinhardt, E.D. 1988.

Modeling long-term fire-caused mortality of Douglasfir. Forest Science. 34(1): 190–199.

Sackett, S.S.; Haase, S.M. 1998. Two case histories for

using prescribed fire to restore ponderosa pine ecosystems in northern Arizona. In: Pruden, T.L,; Brennan,

L.A., eds. Fire in ecosystem management: shifting

the paradigm from suppression to prescription. Proceedings of the 20th Tall Timbers fire ecology conference, Tallahassee, FL: Tall Timbers Research Station:

380–389.

Sackett, S.S.; Haase, S.M.; Harrington, M.G. 1996.

Prescribed burning in southwestern ponderosa pine. In:

Ffolliott, P.F.; DeBano, L.F.; Baker, M.B., Jr.; Gottfried,

G.J.; Solis-Garza, B.; Edminster, C.B.; Neary, D.G.;

Allen, L.S.; Hamre, R.H., eds. Effects of fire on Madrean

Province ecosystems. Gen. Tech. Rep. RM-289. Fort

Collins, CO: U.S. Department of Agriculture, Forest

Service, Rocky Mountain Forest and Range Experiment Station: 178–186.

Sakulich, J.; Taylor, A.H. 2007. Fire regime and forest

structure in a sky island mixed conifer forest,

Guadalupe Mountains National Park, Texas, USA.

Forest Ecology and Management. 241: 62–73.

Sala, A.; Peters, G.D.; McIntyre, L.R.; Harrington, M.G.

2005. Physiological responses of ponderosa pine in

western Montana to thinning, prescribed fire and

burning season. Tree Physiology. 25(3): 339–348.

Schneider, R.E. 1988. The effect of variation in season

of burning on a pine-wiregrass savanna in the Green

Swamp, North Carolina. Durham, NC: Duke University.

141 p. Ph.D. dissertation.

Schroeder, M.J.; Buck, C.C. 1970. Fire weather: a guide

for application of meteorological information to forest

fire control operations. Agric. Handb. 360. Washington,

DC: U.S. Department of Agriculture, Forest Service.

229 p.

Schurbon, J.M.; Fauth, J.E. 2003. Effects of prescribed

burning on amphibian diversity in a Southeastern

U.S. National Forest. Conservation Biology. 17(5):

1338–1349.
77

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

Schwartz, M.W.; Heim, J.R. 1996. Effects of a prescribed

fire on degraded forest vegetation. Natural Areas

Journal. 16(3): 184–191

Schwemlein, D.J.; Williams, R.A. 2007. Effects of landscape position and season of burn on fire temperature

in southern Ohio’s mixed oak forests. In: Buckley, D.S.;

Clatterbuck, W.L., eds. Proceedings, 15th central hardwood forest conference. Gen. Tech. Rep. SRS-101.

Asheville, NC: U.S. Department of Agriculture, Forest

Service, Southern Research Station: 250–257.

Schwilk, D.W.; Knapp, E.E.; Ferrenberg, S.M.; Keeley,

J.E.; Caprio, A.C. 2006. Tree mortality from fire and

bark beetles following early and late season prescribed

fires in a Sierra Nevada mixed-conifer forest. Forest

Ecology and Management. 232: 36–45.

Seastedt, T.R.; Ramundo, R.A. 1990. The influence of

fire on belowground processes of tallgrass prairie.

In: Collins, S.L.; Wallace, L.L., eds. Fire in North

American Tallgrass Prairies. Norman, OK: University

of Oklahoma Press: 99–117.

Shumway, D.; Abrams, M.D.; Ruffner, C.M. 2001. A

400-year history of fire and oak recruitment in an

old-growth oak forest in western Maryland, USA.

Canadian Journal of Forest Research. 31(8):

1437–1443.

Simmons, M.T.; Windhager, S.; Power, P.; Lott, J.; Lyons,

R.K.; Schwope, C. 2007. Selective and non-selective

control of invasive plants: the short-term effects of

growing-season prescribed fire, herbicide, and mowing in two Texas prairies. Restoration Ecology. 15(4):

662–669.

Sisson, D.C.; Speake, D.W.; Landers, J.L.; Buckner, J.L.

1990. Effects of prescribed burning on wild turkey

habitat preference and nest site selection in south

Georgia. National Wild Turkey Symposium. 6: 44–50.

Skinner, C.N.; Chang, C.-R. 1996. Fire regimes, past and

present. Sierra Nevada Ecosystem Project: final report

to Congress. Assessments and scientific basis for

management options. Davis, CA: Centers for Water and

Wildland Resources, University of California Davis:

1041–1069. Vol. 2.

Slocum, M.G.; Platt, W.J.; Cooley, H.C. 2003. Effects of

differences in prescribed fire regimes on patchiness and

intensity of fires in subtropical savannas of Everglades

National Park, Florida. Restoration Ecology. 11(1):

91–102.

Smith, J.E.; McKay, D.; Niwa, C.G.; Thies, W.G.;

Brenner, G.; Spatafora, J.W. 2004. Short-term effects

of seasonal prescribed burning on ectomycorrhizal

fungal community and root biomass in ponderosa pine

stands in the Blue Mountains of Oregon. Canadian

Journal of Forest Research. 34: 2477–2491.

Smith, M.D.; Knapp, A.K. 1999. Exotic plant species in a

C4-dominated grassland: invasibility, disturbance and

community structure. Oecologia. 120: 605–612.

Sparks, J.C.; Masters, R.E. 1996. Fire seasonality effects

on vegetation in mixed-, tall-and southeastern pinegrassland communities. In: Wadsworth, K.G.; McCabe,

R.E., eds. Transactions of the 61st North American wildlife and natural resources conference, Washington, DC:

Wildlife Management Institute: 246–255.

Sparks, J.C.; Masters, R.E.; Engle, D.M.; Bukenhofer,

G.A. 2002. Season of burn influences fire behavior and

fuel consumption in restored shortleaf pine-grassland

communities. Restoration Ecology. 10(4): 714–722.

Sparks, J.C.; Masters, R.E.; Engle, D.M.; Palmer, M.W.;

Bukenhofer, G.A. 1998. Effects of late growing-season

and late dormant-season prescribed fire on herbaceous

vegetation in restored pine-grassland communities.

Journal of Vegetation Science. 9: 133–142.
78

GENERAL TECHNICAL REPORT PSW-GTR-224

Sparks, J.C.; Masters, R.E.; Engle, D.M.; Payton, M.E.;

Bukenhofer, G.A. 1999. Influence of fire season and

fire behavior on woody plants in red-cockaded woodpecker clusters. Wildlife Society Bulletin. 27(1):

124–133.

Sperry, J.H.; George, T.L.; Zack, S. 2008. Ecological

factors affecting response of dark-eyed juncos to prescribed burning. The Wilson Journal of Ornithology.

120(1): 131–138.

Spier, L.P.; Snyder, J.R. 1998. Effects of wet- and dryseason fires on Jacquemontia curtisii, a south Florida

pine forest endemic. Natural Areas Journal. 18(4):

350–357.

Steele, R.W.; Beaufait, W.R. 1969. Spring and autumn

broadcast burning of interior Douglas-fir slash. Bulletin 36. Missoula, MT: University of Montana, School

of Forestry. 12 p.

Steuter, A.A. 1987. C3

/C4 production shift on seasonal

burns—northern mixed prairie. Journal of Range Management. 40: 27–31.

Steuter, A.A.; McPherson, G.R. 1995. Fire as a physical

stress. In: Bedunah, D.J.; Sosebee, R.E., eds. Wildland

plants: physiological ecology and developmental

morphology. Denver, CO: Society for Range Management: 550–579.

Stewart, O.C. 2002. Forgotten fires: Native Americans and

the transient wilderness. Norman, OK: University of

Oklahoma Press. 364 p.

Stoddard, H.L. 1931. The Bobwhite quail: its habits,

preservation, and increase. New York, NY: Charles

Scribner’s Sons. 559 p.

Streng, D.R.; Glitzenstein, J.S.; Platt, B. 1993. Evaluating

effects of season of burn in longleaf pine forests: a

critical literature review and some results from an

ongoing long-term study. In: Hermann, S.M., ed. The

longleaf pine ecosystem ecology, restoration, and management. Proceedings of the 18th Tall Timbers fire

ecology conference. Tallahassee, FL: Tall Timbers

Research Station: 227–264.

Sutherland, E.K. 1997. History of fire in a southern Ohio

second-growth mixed-oak forest. In: Pallardy, S.G.;

Cecich, R.A.; Garrett, E.H.; Johnson, P.S., eds. Proceedings, 11th central hardwood forest conference. Gen.

Tech. Rep. NC-188. St. Paul, MN: U.S. Department of

Agriculture, Forest Service, North Central Forest

Experiment Station: 172–183.

Svejcar, T.J. 1990. Response of Andropogon gerardii to

fire in the tallgrass prairie. In: Collins, S.L.; Wallace,

L.L., eds. Fire in North American tallgrass prairies.

Norman, OK: University of Oklahoma Press: 19–27.

Swetnam, T.W.; Baisan, C.H.; Kaib, J.M. [In press]. Forest

fire histories of La Frontera: fire scar reconstructions of

fire regimes in the United States/Mexico borderlands.

In: Webster, G.L.; Bahre, C.J., eds. Vegetation and flora

of La Frontera: historic vegetation change along the

United States/ Mexico boundary. Albuquerque, NM:

University of New Mexico Press.

Swezy, D.M.; Agee, J.K. 1991. Prescribed fire effects

on fine-root and tree mortality in old-growth ponderosa pine. Canadian Journal of Forest Research. 21:

626–634.

Sword Sayer, M.A.; Goelz, J.C.G.; Haywood, J.D. 2006.

Effects of prescribed fire on production of foliage by

sapling longleaf pine. In: Connor, K.F., ed. Proceedings, 13th biennial southern silvicultural research conference. Gen. Tech. Rep. SRS-92. Asheville, NC: U.S.

Department of Agriculture, Forest Service, Southern

Research Station: 478–485.
79

Ecological Effects of Prescribed Fire Season: A Literature Review and Synthesis for Managers

Taylor, A.H. 2000. Fire regimes and forest changes in mid

and upper montane forests of the southern Cascades,

Lassen Volcanic National Park, California, U.S.A.

Journal of Biogeography. 27: 87–104.

Taylor, A.H.; Skinner, C.N. 2003. Spatial patterns and

controls on historical fire regimes and forest structure

in the Klamath Mountains. Ecological Applications.

13(3): 704–719.

Thatcher, B.S.; Krementz, D.G.; Woodrey, M.S. 2006.

Henslow’s sparrow winter-survival estimates and

response to prescribed burning. Journal of Wildlife

Management. 70(1): 198–206.

Thies, W.G.; Westlind, D.J.; Loewen, M. 2005. Season of

prescribed burn in ponderosa pine forests of eastern

Oregon: impact on pine mortality. International

Journal of Wildland Fire. 14(3): 223–231.

Thies, W.G.; Westlind, D.J.; Loewen, M.; Brenner, G.

2008. A field guide to predict delayed mortality of firedamaged ponderosa pine: application and validation

of the Malheur model. Gen. Tech. Rep. PNW-GTR-769.

Portland, OR: U.S. Department of Agriculture, Forest

Service, Pacific Northwest Research Station. 16 p.

Thompson, D.J.; Shay, J.M. 1989. First-year response of a

Phragmites marsh community to seasonal burning.

Canadian Journal of Botany. 67: 1448–1455.

Towne, E.G.; Kemp, K.E. 2003. Vegetation dynamics from

annually burning tallgrass prairie in different seasons.

Journal of Range Management. 56: 185–192.

Towne, E.G.; Kemp, K.E. 2008. Long-term response

patterns of tallgrass prairie to frequent summer burning. Rangeland Ecology and Management. 61(5):

509–520.

Towne, G.; Owensby, C. 1984. Long-term effects of annual

burning at different dates in ungrazed Kansas tallgrass.

Journal of Range Management. 37: 392–397.

Tucker, J.W., Jr.; Robinson, W.D.; Grand, J.B. 2004.

Influence of fire on Bachman’s sparrow, an endemic

North American songbird. Journal of Wildlife Management. 68(4): 1114–1123.

Tucker, J.W., Jr.; Robinson, W.D.; Grand, J.B. 2006.

Breeding productivity of Bachman’s sparrows in firemanaged longleaf pine forests. The Wilson Journal of

Ornithology. 18(2): 131–280.

Van Lear, D.H.; Carroll, W.D.; Kapeluck, P.R.; Johnson,

R. 2005. History and restoration of the longleaf pinegrassland ecosystem: implications for species at risk.

Forest Ecology and Management. 211(1-2): 150–165.

Varner, J.M.; Hiers, J.K.; Ottmar, R.; Gordon, D.R.; Putz,

F.E.; Wade, D. 2007. Overstory tree mortality resulting

from reintroducing fire to long-unburned longleaf pine

forests: the importance of duff moisture. Canadian

Journal of Forest Research. 37: 1349–1358.

Vermeire, L.T.; Mitchell, R.B.; Fuhlendorf, S.D.; Wester,

D.B. 2004. Selective control of rangeland grasshoppers

with prescribed fire. Journal of Range Management.

57: 29–33.

Vogl, R.J. 1979. Some basic principles of grassland fire

management. Environmental Management. 3(1):

51–57.

Volland, L.A.; Dell, J.D. 1981. Fire effects on Pacific

Northwest forest and range vegetation. Portland, OR:

U.S. Department of Agriculture, Forest Service, Pacific

Northwest Region. 23 p.

Wade, D.D.; Brock, B.L.; Brose, P.; Grace, J.B.; Hoch,

G.A.; Patterson, W.A. 2000. Fire in eastern ecosystems. In: Brown, J.K.; Smith, J.K., eds. Wildland fire in

ecosystems: effects of fire on flora. Gen. Tech. Rep.

RMRS-GTR-42-vol. 2. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain

Research Station: 53–96.
80

GENERAL TECHNICAL REPORT PSW-GTR-224

Wade, D.D.; Johansen, R.W. 1986. Relating wildland fire

to defoliation and mortality in pine. In: Proceedings,

4th biennial southern silvicultural research conference.

Gen. Tech. Rep. SE-42. Asheville, NC: U.S. Department

of Agriculture, Forest Service, Southeastern Forest

Experiment Station: 107–110.

Wade, D.D.; Lunsford, J.D. 1989. A guide for prescribed

fire in southern forests. Tech. Publ. R8-TP11. Atlanta,

GA: U.S. Department of Agriculture, Forest Service,

Southern Region. 56 p.

Waldrop, T.A.; Lloyd, F.T. 1991. Forty years of prescribed

burning on the Santee fire plots: effects on overstory

and midstory vegetation. In: Nodvin, S.C.; Waldrop,

T.A., eds. Fire and the environment: ecological and

cultural perspectives, Proceedings of an international

symposium. Gen. Tech. Rep. SE-69. Knoxville, TN:

U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station: 45–50.

Waldrop, T.A.; Van Lear, D.H.; Lloyd, F.T.; Harms, W.R.

1987. Long-term studies of prescribed burning in

loblolly pine forests of the southeastern Coastal Plain.

Gen. Tech. Rep. SE-45. Asheville, NC: U.S. Department

of Agriculture, Forest Service Southern Research

Station. 23 p.

Waldrop, T.A.; White, D.L.; Jones, S.M. 1992. Fire

regimes for pine-grassland communities in the Southeastern United States. Forest Ecology and Management. 47: 195–210.

Warren, S.D.; Scifres, C.J.; Teel, P.D. 1987. Response of

grassland arthopods to burning: a review. Agriculture,

Ecosystems and Environment. 19: 105–130.

Weatherspoon, C.P. 1988. Preharvest prescribed burning

for vegetation management: effects on Ceanothus

velutinus seeds in duff and soil. In: Proceedings, 9th

annual vegetation management conference. Redding,

CA: Forest Vegetation Management Conference:

125–141.

Weatherspoon, C.P.; Skinner, C.N.; Simpson, C.S. 1991.

Reducing bearclover by repeated growing-season

prescribed burns: preliminary test results. In: Proceedings, 12th annual forest vegetation management conference. Redding, CA: Shasta County Opportunity

Center: 1–9.

Weise, D.R.; Wade, D.D.; Johansen, R.W. 1989. Survival

and growth of young southern pine after simulated

crown scorch. In: MacIver, D.C.; Auld, H.; Whitewood,

R., eds. Proceedings, 10th conference on fire and forest

meteorology. Bethesda, MD: Society of American

Foresters: 161–168.

Westmeier, R.L. 1973. Prescribed burning in grassland

management for prairie chickens in Illinois. In:

Komarek, E.V., ed. Proceedings, 12th Tall Timbers fire

ecology conference, Tallahassee, FL: Tall Timbers

Research Station: 317–338.

White, D.L.; Waldrop, T.A.; Jones, S.M. 1991. Forty years

of prescribed burning on the Santee fire plots: effects

on understory vegetation. In: Nodvin, S.C.; Waldrop,

T.A., eds. Fire and the environment: ecological and

cultural perspectives: Proceedings of an international

symposium. Gen. Tech. Rep. SE-69. Knoxville, TN:

U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station: 51–59.

White, R.S.; Currie, P.O. 1983. Prescribed burning in the

northern Great Plains: yield and cover responses of

three forage species in a mixed grass prairie. Journal

of Range Management. 36: 179–183.

Wright, C.; Agee, J.K. 2004. Fire and vegetation history

in the eastern Cascade Mountains, Washington.

Ecological Applications. 14(2): 443–459.

Wright, H.A.; Klemmedson, J.O. 1965. Effect of fire on

bunchgrasses of the sagebrush-grass region in southern

Idaho. Ecology. 46(5): 680–688.

Yager, L.Y.; Hinderliter, M.G.; Heise, C.D.; Epperson,

D.M. 2007. Gopher tortoise response to habitat management by prescribed burning. Journal of Wildlife

Management. 71(2): 428–434.
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