United States

Department

of Agriculture

Forest Service

Rocky Mountain

Research Station

General Technical

Report RMRS-GTR-42-

volume 1

January 2000

Wildland Fire in

Ecosystems

Effects of Fire on Fauna
Abstract _____________________________________

Smith, Jane Kapler, ed. 2000. 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. 83 p.

Fires affect animals mainly through effects on their habitat. Fires often cause short-term increases in

wildlife foods that contribute to increases in populations of some animals. These increases are

moderated by the animals’ ability to thrive in the altered, often simplified, structure of the postfire

environment. The extent of fire effects on animal communities generally depends on the extent of change

in habitat structure and species composition caused by fire. Stand-replacement fires usually cause

greater changes in the faunal communities of forests than in those of grasslands. Within forests, standreplacement fires usually alter the animal community more dramatically than understory fires. Animal

species are adapted to survive the pattern of fire frequency, season, size, severity, and uniformity that

characterized their habitat in presettlement times. When fire frequency increases or decreases

substantially or fire severity changes from presettlement patterns, habitat for many animal species

declines.

Keywords: fire effects, fire management, fire regime, habitat, succession, wildlife

The volumes in “The Rainbow Series” will be published during the year 2000. To order, check the box or boxes below, fill in the

address form, and send to the mailing address listed below. Or send your order and your address in mailing label form to one of the

other listed media. Your order(s) will be filled over the months of 2000 as the volumes are published.

RMRS-GTR-42-vol. 1. Wildland fire in ecosystems: effects of fire on fauna.

RMRS-GTR-42-vol. 2. Wildland fire in ecosystems: effects of fire on flora.

RMRS-GTR-42-vol. 3. Wildland fire in ecosystems: effects of fire on cultural resources and archeology.

RMRS-GTR-42-vol. 4. Wildland fire in ecosystems: effects of fire on soil and water.

RMRS-GTR-42-vol. 5. Wildland fire in ecosystems: effects of fire on air.

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Editor

Jane Kapler Smith, Rocky Mountain Research Station, U.S. Department of Agriculture, Forest Service, Missoula, MT 59807.

Authors

L. Jack Lyon, Research Biologist (Emeritus) and Project Leader for the

Northern Rockies Forest Wildlife Habitat Research Work Unit, Intermountain (now Rocky Mountain) Research Station, U.S. Department

of Agriculture, Forest Service, Missoula, MT 59807.

Mark H. Huff, Ecologist, Pacific Northwest Research Station, U.S.

Department of Agriculture, Forest Service, Portland, OR 97208.

Robert G. Hooper, Research Wildlife Biologist, Southern Research

Station, U.S. Department of Agriculture, Forest Service, Charleston,

SC 29414.

Edmund S. Telfer, Scientist (Emeritus), Canadian Wildlife Service,

Edmonton, Alberta, Canada T6B 2X3.

David Scott Schreiner, Silvicultural Forester (retired), Los Padres

National Forest, U.S. Department of Agriculture, Forest Service, Goleta,

CA 93117.

Jane Kapler Smith, Ecologist, Fire Effects Research Work Unit, Rocky

Mountain Research Station, U.S. Department of Agriculture, Forest

Service, Missoula, MT 59807.

Cover photo—Male black-backed woodpecker on fire-killed lodgepole pine. Photo by Milo Burcham.

Wildland Fire in Ecosystems

Effects of Fire on Fauna
Preface _____________________________________

In 1978, a national workshop on fire effects in Denver, Colorado, provided the impetus

for the “Effects of Wildland Fire on Ecosystems” series. Recognizing that knowledge of

fire was needed for land management planning, state-of-the-knowledge reviews were

produced that became known as the “Rainbow Series.” The series consisted of six

publications, each with a different colored cover, describing the effects of fire on soil,

water, air, flora, fauna, and fuels.

The Rainbow Series proved popular in providing fire effects information for professionals,

students, and others. Printed supplies eventually ran out, but knowledge of fire effects

continued to grow. To meet the continuing demand for summaries of fire effects knowledge,

the interagency National Wildfire Coordinating Group asked Forest Service research leaders

to update and revise the series. To fulfill this request, a meeting for organizing the revision was

held January 4-6, 1993, in Scottsdale, Arizona. The series name was then changed to “The

Rainbow Series.” The five-volume series covers air, soil and water, fauna, flora and fuels, and

cultural resources.

The Rainbow Series emphasizes principles and processes rather than serving as a

summary of all that is known. The five volumes, taken together, provide a wealth of information

and examples to advance understanding of basic concepts regarding fire effects in the United

States and Canada. As conceptual background, they provide technical support to fire and

resource managers for carrying out interdisciplinary planning, which is essential to managing

wildlands in an ecosystem context. Planners and managers will find the series helpful in many

aspects of ecosystem-based management, but they will also need to seek out and synthesize

more detailed information to resolve specific management questions.

–– The Authors

January 2000

Acknowledgments____________________________

The Rainbow Series was completed under the sponsorship of the Joint Fire Sciences

Program, a cooperative fire science effort of the U.S. Department of Agriculture, Forest Service

and the U.S. Department of the Interior, Bureau of Indian Affairs, Bureau of Land Management,

Fish and Wildlife Service, and National Park Service. We thank Marcia Patton-Mallory and

Louise Kingsbury for persistence and support.

The authors are grateful for reviews of the manuscript from James K. Brown, Luc C.

Duchesne, R. Todd Engstrom, Bill Leenhouts, Kevin C. Ryan, and Neil Sugihara; the

reviews were insightful and helpful. Reviews of special topics were provided by David R.

Breininger, John A. Crawford, Steve Corn, and Kevin R. Russell; their help strengthened

many sections of the manuscript. We are thankful to Nancy McMurray for editing; Dennis

Simmerman for assistance with graphics; Bob Altman for literature reviews of special topics;

Loren Anderson, Steve Arno, Milo Burcham, Robert Carr, Chris Clampitt, Betty Cotrille, Kerry

Foresman, Jeff Henry, Catherine Papp Herms, Robert Hooper, Dick Hutto, Bob Keane, Larry

Landers, Melanie Miller, Jim Peaco, Dean Pearson, Rick McIntyre, Dale Wade, and Vita

Wright for providing photographs or helping us locate them.

ii
Summary ........................................................................... iv

Chapter 1: Introduction .................................................... 1

by L. Jack Lyon, James K. Brown, Mark H. Huff, and

Jane Kapler Smith

Historic Perspective ............................................................ 2

Fire Regimes ....................................................................... 3

Changes in Vegetation Structure ........................................ 4

Understory Fire Regimes ............................................ 4

Stand-Replacement Fire Regimes ............................. 5

Mixed-Severity Fire Regimes ..................................... 6

Snags and Dead Wood ....................................................... 7

Chapter 2: Regional Variation in Fire Regimes .............. 9

by Edmund S. Telfer

Northern Ecosystems .......................................................... 9

Boreal Forest .............................................................. 9

Laurentian Forest ..................................................... 10

Eastern Ecosystems and the Great Plains ....................... 10

Eastern Deciduous Forest ........................................ 10

Southeastern Forests ............................................... 11

Prairie Grassland ...................................................... 11

Western Forests ................................................................ 12

Rocky Mountain Forest ............................................. 12

Sierra Forest ............................................................. 12

Pacific Coast Maritime Forest ................................... 13

Western Woodlands, Shrublands, and Grasslands .......... 13

Pinyon-Juniper .......................................................... 13

Chaparral and Western Oak Woodlands .................. 13

Sagebrush and Sagebrush Grasslands .................... 14

Deserts ..................................................................... 14

Subtropical Ecosystems .................................................... 15

Florida Wetlands ....................................................... 15

Chapter 3: Direct Effects of Fire and Animal

Responses .................................................... 17

by L. Jack Lyon, Edmund S. Telfer, and

David Scott Schreiner

Injury and Mortality ............................................................ 17

Birds.......................................................................... 18

Mammals .................................................................. 18

Reptiles and Amphibians .......................................... 20

Invertebrates ............................................................. 20

Escape and Emigration ..................................................... 20

Birds.......................................................................... 20

Mammals .................................................................. 21

Immigration ....................................................................... 21

Birds.......................................................................... 21

Mammals .................................................................. 22

Reptiles and Amphibians .......................................... 23

Chapter 4: Fire Effects on Animal Populations ............ 25

by L. Jack Lyon, Mark H. Huff, Edmund S. Telfer,

David Scott Schreiner, and Jane Kapler Smith

Changes in Animal Populations ........................................ 25

Birds.......................................................................... 25

Mammals .................................................................. 28

Reptiles and Amphibians .......................................... 29

Invertebrates ............................................................. 30

Effects of Altered Fire Regimes ........................................ 31

Understory Fire Regimes .......................................... 31

Stand-Replacement Fire Regimes ........................... 32

Mixed-Severity Fire Regimes ................................... 33

Animal Influences on Postfire Habitat ............................... 33

Chapter 5: Fire Effects on Animal Communities ......... 35

by Mark H. Huff and Jane Kapler Smith

Understory Fire Regimes .................................................. 36

Frequent Understory Fires ........................................ 36

Infrequent Understory Fires ...................................... 37

Stand-Replacement Fire Regimes .................................... 37

Grasslands................................................................ 37

Shrub-Grasslands ..................................................... 38

Shrublands ............................................................... 39

Forests and Woodlands ............................................ 40

Mixed-Severity Fire Regimes ............................................ 42

Chapter 6: Fire Effects on Fauna at Landscape

Scales ............................................................ 43

by L. Jack Lyon, Mark H. Huff, and Jane Kapler Smith

Effects of Altered Fire Regimes ........................................ 48

Chapter 7: Fire Effects on Wildlife Foods ..................... 51

by L. Jack Lyon, Robert G. Hooper, Edmund S. Telfer, and

David Scott Schreiner

Northern Ecosystems ........................................................ 52

Boreal Forest ............................................................ 52

Laurentian Forest ..................................................... 52

Eastern Ecosystems and the Great Plains ....................... 52

Eastern Deciduous Forests ...................................... 52

Southeastern Forests ............................................... 53

Prairie Grassland ...................................................... 55

Western Forests ................................................................ 56

Rocky Mountain Forest ............................................. 56

Sierra Forest ............................................................. 57

Pacific Coast Maritime Forest ................................... 57

Western Woodlands, Shrublands, and Grasslands .......... 57

Pinyon-Juniper .......................................................... 57

Chaparral and Western Oak Woodlands .................. 57

Sagebrush and Sagebrush Grasslands .................... 58

Deserts ..................................................................... 58

Subtropical Ecosystems .................................................... 58

Florida Wetlands ....................................................... 58

Chapter 8: Management and Research

Implications .................................................. 59

by L. Jack Lyon and Jane Kapler Smith

Management Implications ................................................. 59

One: Alternatives in Managing Fire .......................... 59

Two: Integrating Management Objectives ................ 60

Needs for Further Understanding ..................................... 60

Fauna-Habitat Relationships .................................... 61

Presettlement Fire Regimes ..................................... 61

Human Dimension ............................................................ 61

References ....................................................................... 63

Appendices ...................................................................... 75

Appendix A: Common and Scientific Names of Animal

Species ..................................................................... 75

Appendix B: Common and Scientific Names of Plant

Species ......................................................................... 79

Appendix C: Glossary ....................................................... 82

Contents________________________________________________

Page Page

iii
Summary

Fire regimes—that is, patterns of fire occurrence, size,

uniformity, and severity—have been a major force shaping landscape patterns and influencing productivity

throughout North America for thousands of years. Faunal

communities have evolved in the context of particular fire

regimes and show patterns of response to fire itself and

to the changes in vegetation composition and structure

that follow fire.

Animals’ immediate responses to fire are influenced by

fire season, intensity, severity, rate of spread, uniformity,

and size. Responses may include injury, mortality, immigration, or emigration. Animals with limited mobility, such

as young, are more vulnerable to injury and mortality than

mature animals.

The habitat changes caused by fire influence faunal

populations and communities much more profoundly

than fire itself. Fires often cause a short-term increase in

productivity, availability, or nutrient content of forage and

browse. These changes can contribute to substantial

increases in herbivore populations, but potential increases

are moderated by animals’ ability to thrive in the altered,

often simplified, structure of the postfire environment.

Fires generally favor raptors by reducing hiding cover

and exposing prey. Small carnivores respond to fire

effects on small mammal populations (either positive or

negative). Large carnivores and omnivores are opportunistic species with large home ranges. Their populations

change little in response to fire, but they tend to thrive in

areas where their preferred prey is most plentiful—often

in recent burns. In forests and woodlands, understory

fires generally alter habitat structure less than mixedseverity and stand-replacement fires, and their effects on

animal populations are correspondingly less dramatic.

Stand-replacing fires reduce habitat quality for species

that require dense cover and improve it for species that

prefer open sites. Population explosions of wood-boring

insects, an important food source for insect predators

and insect-eating birds, can be associated with fire-killed

trees. Woodpecker populations generally increase after

mixed-severity and stand-replacement fire if snags are

available for nesting. Secondary cavity nesters, both

birds and mammals, take advantage of the nest sites

prepared by primary excavators.

Many animal-fire studies depict a reorganization of

animal communities in response to fire, with increases in

some species accompanied by decreases in others.

Like fire effects on populations, fire effects on communities are related to the amount of structural change in

vegetation. For example, understory fires and standreplacement fires in grasslands often disrupt bird community composition and abundance patterns for only

1 to 2 years, but stand-replacement fires in shrublands and forests cause longer lasting effects, which

are initially positive for insect- and seed-eating species

and negative for species that require dense, closed

canopy. Bird abundance and diversity are likely to be

greatest early in succession. When shrub or tree canopy

closure occurs, species that prefer open sites and habitat

edges decline and species that prefer mature structures

increase.

Major changes to fire regimes alter landscape patterns, processes, and functional linkages. These

changes can affect animal habitat and often produce

major changes in the composition of faunal communities. In many Western ecosystems, landscape changes

due to fire exclusion have changed fuel quantities and

arrangement, increasing the likelihood of large or

severe fires, or both. Where fire exclusion has changed

species composition and fuel arrays over large areas,

subsequent fires without prior fuel modification are

unlikely to restore presettlement vegetation and habitat.

In many desert and semidesert habitats where fire

historically burned infrequently because of sparse

fuels, invasion of weedy species has changed the

vegetation so that burns occur much more frequently.

Many animals in these ecosystems are poorly adapted

to avoid fire or use resources in postfire communities.

In the past 10,000 years, fire in North American ecosystems has not operated in isolation from other disturbances, nor has it occurred independent of human

influence. In many areas, however, fire has been prevented or excluded for nearly 100 years, a level of

success that is not likely to continue. Collaboration

among managers, researchers, and the public is needed

to address tradeoffs in fire management, and fire management must be better integrated with overall land

management objectives to address the potential interactions of fire with other disturbances such as

grazing, flood, windthrow, and insect and fungus

infestations.

iv
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 1

L. Jack Lyon

James K. Brown

Mark H. Huff

Jane Kapler Smith

Chapter 1:

Introduction

Effects of wildland fire on fauna show almost infinite variety. Previous authors have limited discussion of this subject to only a few vertebrate groups

(Bendell 1974), specific biotic provinces (Fox 1983;

Stanton 1975), or general summaries (Lyon and others 1978). This report surveys the principles and

processes governing relationships between fire and

fauna. We recognize that this approach has limitations. We focus almost entirely on vertebrates, particularly terrestrial mammals and birds, because

the information on those groups is most complete

and the principles best documented. (Fire effects on

aquatic vertebrates are summarized in “Effects of Fire

on Soil and Water,” another volume in this Rainbow

Series.) We describe fire effects on specific faunal

populations and communities by way of example, not

as a survey of all that is known. Those seeking a

detailed description of fire effects on fauna are referred to books that discuss the subject in general,

such as Whelan (1995, chapter 6) and Wright and

Bailey (1982, chapter 4); reports about fire effects in

specific geographic regions (for example, McMahon

and deCalesta 1990; Viereck and Dyrness 1979); and

summaries of fire effects on specific faunal groups

(for example, Crowner and Barrett 1979; Lehman

and Allendorf 1989; Russell 1999). The Fire Effects

Information System, on the Internet at www.fs.fed.us/

database/feis, provides detailed descriptions of fire

effects on more than 100 North American animal

species and nearly 1,000 plant species.

Fires affect fauna mainly in the ways they affect

habitat. Repeated fires have been a major force

shaping landscapes and determining productivity

throughout North America for thousands of years,

with the possible exception of some portions of West

Coast rain forests. Climate, vegetation, Native Americans, and fire interacted in a relatively consistent

manner within each biotic region of North America

before the advent of disease and settlers from Europe

(Kay 1998). Prior to modern agriculture, fire suppression, and urbanization, vegetation patterns in each

region were shaped by fire regimes with characteristic

severity, size, and return interval (Frost 1998; Gill

1998; Heinselman 1981; Kilgore 1981).

The animal species native to areas with a centurieslong history of fire can obviously persist in habitat

shaped by fire; many species actually thrive because

of fire’s influence. How? Animals’ immediate response to fire may include mortality or movement. It

is influenced by fire intensity, severity, rate of

spread, uniformity, and size. Long-term faunal response to fire is determined by habitat change, which

influences feeding, movement, reproduction, and availability of shelter (fig. 1). Alteration of fire regimes

alters landscape patterns and the trajectory of change

on the landscape; these changes affect habitat and

often produce major changes in faunal communities.
2 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

Figure 1—Elk rest and graze in unburned meadow adjacent to area burned by crown fire, Yellowstone National Park. Fire and

fauna have coexisted in this ecosystem for thousands of years. Photo by Rick McIntyre, courtesy of National Park Service.

This volume is organized on the premises that fire

regimes strongly influence animal response to fire and

that fire affects animals at every level of ecosystem

organization. In this chapter, we describe the fundamental concepts of fire regimes and their effects on

vegetation structure. Chapter 2 describes the role of

fire in several North American vegetation communities prior to settlement by European Americans. Because the vegetation provides habitat for fauna, this

chapter provides background for understanding examples used in later chapters. The next four chapters

describe animal response to fire at four levels of

organization: individual, population, community, and

landscape. Chapter 7 surveys fire effects on wildlife

foods. Finally, chapter 8 discusses management

implications of fire-fauna relationships, particularly in light of past fire exclusion, and identifies

information gaps and research needs. Scientific

names of all animals described in this report are

listed in appendix A. Appendix B lists scientific

names of plants. Appendix C contains a glossary of

technical terms.

Historic Perspective _____________

Fire has influenced composition, structure, and

landscape patterns of animal habitat for millennia, so

it is reasonable to assume that animals have coexisted

and adapted to periodic perturbations from fire. Records

show that lightning starts more than 6,000 fires each

year in the United States; surely this force was just as

powerful and ubiquitous in past millennia as it is now

(Pyne 1982). Prior to the 1500s, millions of Native

Americans lived in North America. They used fire

regularly for many purposes (Kay 1998). Only recently, since the advent of fire exclusion policy and

other activities that strongly influence fire regimes,

has fire’s influence on fauna been intensely questioned and investigated (Kilgore 1976).

During the era of European settlement of North

America, fire came to be viewed in some geographic

areas as a hopelessly destructive event that could

not be stopped. Early legislation promoting fire

control responded primarily to the loss of lives and

settlements and vast mortality of harvestable trees
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 3

that occurred in large fires around the end of the

18th century (for example, the Peshtigo Fire of

1871). However, proclaimed reductions of elk, deer,

bison, and other wildlife populations associated with

these fire events were also important considerations

in establishing fire control legislation (Brown and

Davis 1973).

Resource protection and associated fire control began with the establishment of forest reserves throughout North America at the end of the 1900s. The

reserves were established mainly to protect the land

from abuse by timber and grazing interests, but early

reports from the reserves singled out fire as the

greatest threat to America’s grasslands, forests, and

wildlife (Komarek 1962). The primary fire-related

mission of land management agencies was reinforced

in the 1930s: to stop fires wherever possible, and to

prevent large fires from developing (Moore 1974).

From that time until the 1960s, most managers and

many of the public viewed fire as an unnatural event

and an environmental disaster. The land area under

vigorous fire protection grew every year, and the

resources assigned to fire suppression grew accordingly (Brown and Davis 1973).

Even in the early days of fire exclusion policy,

there were dissenting voices. In 1912, ecologist

H. H. Chapman recognized that longleaf pine in the

South was adapted to grow and mature in the presence of repeated fires (Chapman 1912). Subsequent

studies by other researchers found that controlled

burning improved quality of ungulate forage (Green

1931) and improved, restored, and maintained habitat for certain game species, especially the northern

bobwhite (Stoddard 1931, 1935, 1936). The scientific

community was beginning to view fire as a natural

process and a tool for wildlife habitat management,

but many public and private land managers strongly

resisted the concept (Schiff 1962).

During World War II, fire suppression capability

declined. The disastrous 1943 drought-related fires

in the Southeastern United States prompted major

shifts in government policies (Schiff 1962). By the

1950s, controlled burning to reduce fuels and enhance habitat for specific wildlife species had become

commonplace, but all other fires were vigorously controlled. Meanwhile, scientists began to report striking changes in plant community composition and

structure associated with fire exclusion. Important

functions of fire were described for ponderosa pine in

the Pacific Northwest (Weaver 1943), California chaparral and ponderosa pine (Biswell 1963), Arizona

ponderosa pine (Cooper 1960), Florida Everglades

(Loveless 1959), and interior Alaska (Lutz 1956).

With the publication of the Leopold Report (Leopold

and others 1963) on ecological conditions of National

Parks in the United States, managers and the public

began to see the benefits of fires in wildlands. The

Leopold Report established the concept that wildlife

habitat is not a stable entity that persists unchanged

in perpetuity, but rather a dynamic entity; suitable

habitat for many wildlife species and communities

must be renewed by fire. Policy began to shift away

from the assumption that all wildland fires are destructive (Pyne 1982). In 1968, the fire policy of the

USDI National Park Service changed drastically as

managers began to adopt the recommendations of the

Leopold Report. Policy officially recognized fire as a

natural process to be managed for maintaining ecosystems and improving wildlife habitat. Thus began the

current era of fire management in which fire is

recognized as an integral part of ecosystems, including those aspects relating to fauna (Habeck and Mutch

1973).

Fire Regimes ___________________

Knowledge of the ecological role of fire in past

centuries and descriptions of significant changes in

the role of fire over time are essential for communication among professionals and citizens interested in

resource management. Nearly every North American

ecosystem has been drastically changed from conditions of past millennia. Regardless of how fire might

be managed in the future in various ecosystems,

information about its past role is important. As

Morgan and others (1994) said, “Study of past ecosystem behavior can provide the framework for understanding the structure and behavior of contemporary

ecosystems, and is the basis for predicting future

conditions.”

Fire varies in its frequency, season, size, and prominent, immediate effects, but general patterns occur

over long periods. These patterns describe fire regimes. The practice of organizing biotic information

around fire regimes originated in North America

around 1980 (Heinselman 1978, 1981; Kilgore 1981;

Sando 1978). Descriptions of fire regimes are general

because of fire’s tremendous variability over time and

space (Whelan 1995). Nevertheless, the fire regime is

a useful concept because it brings a degree of order to

a complicated body of knowledge. The fire regimes

that have influenced North American ecosystems in

an evolutionary sense are those of pre-Columbian

times (prior to 1500), before diseases introduced by

European explorers began to decimate populations of

Native Americans (see Kay 1995). While knowledge of

pre-Columbian fire regimes would be useful for understanding ecosystem patterns and processes today,

little information is available from that era. Detailed

information available about past fire regimes is mostly

based on biophysical evidence, written records, and

oral reports that encompass the time from about 1500
4 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

to the mid- to late-1800s. This was a time before

extensive settlement by European Americans in most

parts of North America, before extensive conversion of

wildlands for agricultural and other purposes, and

before fire suppression effectively reduced fire frequency in many areas. In this volume, we refer to the

fire regimes of the past several centuries as “presettlement” fire regimes.

Fire frequency and severity form the basis for the

commonly referenced fire regime classifications described by Heinselman (1978) and Kilgore (1981). Two

concepts, fire return interval and fire rotation, describe the frequency with which fires occur on a landscape. Mean fire return interval is the average number

of years between fires at a given location. Fire rotation,

called by some authors the fire cycle, is the number of

years that would be required to completely burn over

a given area.

Fire severity describes the immediate effects of

fire, which result from the rate of heat release in the

fire’s flaming front and the total heat released during

burning. Fire severity determines in large part the

mortality of dominant vegetation and changes in the

aboveground structure of the plant community, so

Kilgore (1981) refers to severe fires in forests as

“stand-replacement” fires. The concept of stand replacement by fire applies to nonforest as well as forest

areas. Fires in vegetation types such as prairie, tundra, and savannah are essentially all stand-replacing

because the aboveground parts of dominant vegetation are killed (and often consumed) by fire. Most

shrubland ecosystems also have stand-replacement

fire regimes because fire usually kills the aboveground

parts of shrubs. In this report, we refer to the following

four kinds of fire regime:

1. Understory fire regime (applies to forest and

woodland vegetation types)—Fires are generally

not lethal to the dominant vegetation and do not

substantially change the structure of the dominant vegetation. Approximately 80 percent or

more of the aboveground dominant vegetation

survives fires.

2. Stand-replacement regime (applies to forests,

shrublands, and grasslands)—Fires kill or topkill aboveground parts of the dominant vegetation, changing the aboveground structure substantially. Approximately 80 percent or more of

the aboveground dominant vegetation is either

consumed or killed as a result of fires.

3. Mixed-severity regime (applies to forests and

woodlands)—Severity of fire either causes selective mortality in dominant vegetation, depending on different species’ susceptibility to

fire, or varies between understory and standreplacement.

4. Nonfire regime—Little or no occurrence of natural fire (not discussed further in this volume).

See “Effects of Fire on Flora” (also in the Rainbow

Series) for further discussion of fire regimes and comparison of this fire regime classification with others.

The literature demonstrates great local variation in

fire effects on habitat, even within small geographic

areas with a single fire regime. Fires theoretically

should spread in an elliptical pattern (Anderson 1983;

Van Wagner 1969), but the shape of burned areas and

the fire severity patterns within them are influenced

by fluctuations in weather during fires, diurnal changes

in burning conditions, and variation in topography,

fuels, and stand structure. Variable and broken topography and sparse fuels are likely to produce patchy

burns, while landscapes with little relief and homogeneous fuels may burn more uniformly. It is no wonder

then that fires shape a complex mosaic of size classes,

vegetation structure, and plant species occurrence

across the landscape, and this variety has a profound

influence on the animals that live there.

Changes in Vegetation

Structure ______________________

For animals, the vegetation structure spatially arranges the resources needed to live and reproduce,

including food, shelter and hiding cover. Some fires

alter the vegetation structure in relatively subtle

ways, for example, reducing litter and dead herbs

in variable-sized patches. Other fires change nearly

every aspect of vegetation structure: woody plants

may be stripped of foliage and killed; litter and duff

may be consumed, exposing mineral soil; underground

structures, such as roots and rhizomes, may be killed

(for example, in most coniferous trees) or rejuvenated (for example, in many grass and shrub species,

aspen, and oak). In this section, we summarize postfire structural changes according to the fire regimes

described above.

Understory Fire Regimes

Understory fires change the canopy in two ways: by

killing or top-killing a few of the most fire-susceptible

trees, and by killing or top-killing a cohort of tree

regeneration, also selectively according to fire resistance. Understory fires also reduce understory plant

biomass, sometimes in a patchy pattern. Although the

structural changes caused by any one understory fire

are not dramatic, repeated understory fires shape and

maintain a unique forest structure identified by O’Hara

and others (1996) as “old forest, single stratum.” It is

characterized by large, old trees, parklike conditions,

and few understory trees (fig. 2).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 5

Figure 2—Mature longleaf pine forest, typical of forest structure maintained by frequent understory fire, in

Francis Marion National Forest, South Carolina. This kind of habitat favors many fauna species, included

red-cockaded woodpecker, Bachman’s sparrow, northern bobwhite, fox squirrel, and flatwoods salamander.

Photo by Robert G. Hooper.

Stand-Replacement Fire Regimes

Grasslands—In grasslands, the prefire structure

of the vegetation reasserts itself quickly as a new

stand of grass springs up from surviving root systems. Standing dead stems and litter are reduced.

The proportion of forbs usually increases in the first

or second postfire year. In about 3 years the grassland structure is usually reestablished (Bock and

Bock 1990), and faunal populations are likely to

resemble those of the preburn community. Repeated

fires can convert some shrublands to grass, and fire

exclusion converts some grasslands to shrubland and

forest.

Shrublands—In shrub-dominated areas, including sagebrush, chaparral, and some oak woodlands,

stand-replacing fires top-kill or kill aboveground

vegetation. Canopy cover is severely reduced, but

initial regrowth usually increases cover of grasses and

forbs. Dead woody stems often remain standing and

serve as perch sites for songbirds, raptors, and even

lizards (fig. 3). Burning increases seed visibility and

availability for small mammals but also increases

their visibility to predators. Because cover for ungulates is reduced by fire, some species do not use the

abundant postfire forage. Shrubs regenerate from

underground parts and seed. The length of time required to reestablish the shrubland structure varies,

from 2 years in saw palmetto scrub (Hilmon and

Hughes 1965) to more than 50 years in big sagebrush

(Wright 1986).

Forests and Woodlands—In tree-dominated areas, stand-replacing fires change habitat structure

dramatically. When crown fire or severe surface fire

kills most of the trees in a stand, surface vegetation

is consumed over much of the area, and cover for

animals that use the tree canopy is reduced. Crown

fires eliminate most cover immediately; severe surface

fires kill the tree foliage, which falls within a few

months. Stand-replacing fires alter resources for herbivores and their predators. The habitat is not “destroyed,” but transformed: The fire-killed trees become food for millions of insect larvae and provide

perches for raptors. Trees infected by decay before the

fire provide nest sites for woodpeckers and then for

secondary cavity nesters (birds and mammals). As
6 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

Figure 3—Sagebrush 3 years after stand-replacing fire, east-central Idaho. Fire kills sagebrush but

leaves dead stems that birds and reptiles use as perches. The photo shows early successional

dominance by dense bluebunch wheatgrass. Photo by Loren Anderson.

these snags fall, other fire-killed trees decay and

provide habitat for cavity nesters. For 10 to 20 years

after stand-replacing fire, biomass is concentrated on

the forest floor, as grasses and forbs, shrubs and tree

saplings reoccupy the site. These provide forage and

dense cover for small mammals, nest sites for shrubland

birds, and a concentrated food source for grazing and

browsing ungulates. In 30 to 50 years after standreplacing fire, saplings become trees and suppress the

early successional shrub and herb layers. The forest

again provides hiding and thermal cover for ungulates

and nesting habitat for animals that use the forest

interior. The remaining fire-killed snags decay and

fall, reducing nest sites for cavity-nesting birds and

mammals but providing large pieces of dead wood on

the ground. This fallen wood serves as cover for small

mammals, salamanders, and ground-nesting birds.

The fungi and invertebrates living in dead wood provide food for birds and small mammals (for example,

see McCoy and Kaiser 1990).

In some northern and western coniferous forests,

the initial postfire stand is composed of broad-leafed,

deciduous trees such as aspen or birch. Conifer dominance follows later in succession. Some bird and

mammal species prefer the broad-leafed successional

stage to earlier and later stages of succession. As

succession continues, conifers dominate and broadleafed trees decay. This process creates snags and

adds to dead wood on the ground, enhancing habitat

for cavity nesters and small mammals. It also creates

openings that are invaded by shrubs and saplings.

Dense patches of shrubs and tree regeneration in

long-unburned forests provide excellent cover for

ungulates. Birds (for example, crossbills, nuthatches,

brown creeper, and woodpeckers), tree squirrels, and

American marten find food, cover, and nest sites

within the structure of the old-growth coniferous

forest.

In some Southeastern forests, the roles of pine and

hardwood tree species are reversed. Many Southeastern forests regenerate to pine immediately after standreplacing fire. In the absence of repeated understory

fires, these pine stands are invaded and eventually

dominated by broad-leafed deciduous species such as

American beech, hickory, and southern magnolia

(Engstrom and others 1984; Komarek 1968). As in the

hardwood-conifer sere of the Western States, each

structural stage supports a somewhat different assemblage of wildlife.

Mixed-Severity Fire Regimes

In mixed-severity fire regimes, fires either cause

selective mortality of fire-susceptible species in the

overstory or alternate between understory and standreplacement, with overlapping burn boundaries. The
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 7

net result is a fine grained pattern of stand ages and

structures across the landscape. This pattern is accentuated in areas where variable topography and microclimate influence fire spread. Through feedback of fuel

patterns into subsequent fire behavior, the variety in

fuels and stand structures resulting from mixedseverity fire perpetuate the complex mosaic of ages

and structures.

Snags and Dead Wood ___________

It would be difficult to overestimate the importance

of large trees, snags, and dead, down wood to North

American birds and small mammals. According to

Brown and Bright (1997), “The snag represents perhaps the most valuable category of tree-form diversity

in the forest landscape.” Fire and snags have a complex relationship. Fires convert live trees to snags, but

fires also burn into the heartwood of old, decayed

snags and cause them to fall. Fire may facilitate decay

in surviving trees by providing an entry point for

fungi, which increases the likelihood that the trees

will be used by cavity excavators. Fire may harden the

wood of trees killed during a burn, causing their outer

wood to decay more slowly than that of trees that die

from other causes. This “case-hardening” process reduces the immediate availability of fire-killed snags

for nest excavation but slows their decay after they fall.

It is difficult to identify fire-injured trees that are

likely to become snags, and it is also difficult to

determine which snags may have the greatest

“longevity,” that is, may stand the longest time before

falling. In ponderosa pine stands in Colorado, for

example, the trees most likely to become long-lasting

snags are underburned trees with moderate crown

scorch that remain alive for at least 2 years after fire,

a group that cannot be determined until 2 or 3 years

after fire (Harrington 1996). According to Smith (1999),

longevity of ponderosa pine snags is positively related

to tree age and size at death. Fire-scarred trees may

have greater longevity than trees never underburned

(Harrington 1996).

The usefulness of snags to fauna is enhanced or

reduced by the surrounding habitat, since cavity nesters vary in their needs for cover and food. Many cavity

excavators require broken-topped snags because partial decay makes them easier to excavate than sound

wood (Caton 1996). Some bird species nest only in

large, old snags, which are likely to stand longer than

small snags (Smith 1999). Pileated woodpeckers are

an example. Some excavators and secondary cavity

nesters prefer clumps of snags to individual snags, so

the spatial arrangement of dead and decaying trees in

an area influences their usefulness to wildlife (Saab

and Dudley 1998).

Dead wood on the ground is an essential habitat

component for many birds, small mammals (fig. 4),

and even large mammals, including bears (Bull and

Blumton 1999). Large dead logs harbor many invertebrates and are particularly productive of ants; they

also provide shelter and cover for small mammals,

amphibians, and reptiles. Fire both destroys and creates woody debris. While large, down logs are not

always abundant in early postfire years, fire-killed

trees eventually fall and become woody debris. Down

wood from fire-killed trees often decays more slowly

than wood of trees killed by other means (Graham and

others 1994).

Figure 4—Vagrant shrew travelling in shelter of dead log, Lolo National Forest,

western Montana. Large dead wood is an essential source of food and shelter for

many small mammals. Photo by Kerry R. Foresman.
8 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

Notes

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 9

To provide a context for discussion of fire effects on

animals and their habitat, this chapter describes the

vegetation, fire regimes, and postfire succession of

several plant communities referred to in subsequent

sections of this report. This description is not meant to

be a complete survey of fire regimes in North America;

such a survey is available in “Effects of Fire on Flora,”

also part of the Rainbow Series. Instead, it provides

examples of plant communities and fire regimes

throughout the continent, many of them described in

earlier reviews, including Wright and Bailey (1982).

These communities are divided according to the geographic regions used to describe fire effects on the

flora in this series: northern ecosystems; eastern ecosystems, including the Great Plains; western forests;

western woodlands, shrublands, and grasslands; and

subtropical ecosystems.

Northern Ecosystems ____________

Boreal Forest

Vegetation and Fire Regimes—The Boreal Forest

was characterized in presettlement times by standreplacement fire regimes, although understory fires

were common in some dry forest types (Heinselman

Edmund S. Telfer

Chapter 2:

Regional Variation in Fire

Regimes

1981; Johnson 1992). Most of the presettlement fire

rotations reported in the literature were relatively

short, ranging from 50 to 100 years (Heinselman 1981;

Payette and others 1989; Wein 1993). Johnson (1992)

determined that the fire rotation was between 40 and

60 years for Minnesota, Ontario, the Northwest Territories, and Alaska. Relatively short rotations probably

occurred in dry continental interior regions; for example, a 39-year rotation in northern Alberta (Murphy

1985). Longer rotations occurred in floodplains

(Heinselman 1981) and Eastern boreal forests (Viereck

1983). From 1980 to 1989, frequency of fires larger

than about 500 acres (200 ha) in the Canadian boreal

forest was greatest in the central part of the continent

and decreased toward the east and northwest. Fires

were more frequent during dry climatic periods than

during wet periods (Clark 1988; Swain 1973).

Due to frequent fires in the Boreal Forest, there

probably has been no time during the last 6,000 to

10,000 years when ancient or even old forest covered a high proportion of the region (Telfer 1993).

Van Wagner (1978) and Johnson (1992) found that

the distribution of forest area over age classes often

approximates a negative exponential distribution,

permitting prediction of the distribution of age class

areas under various fire rotations. Based on this
10 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

relationship, Johnson (1992) commented that a 40- to

60-year fire rotation, “...by definition, suggests that

most (63 percent) stands will never live much beyond

the age at which stand canopy closure occurs and very

few will reach anything resembling old age.”

Postfire Succession—Principal Boreal Forest

trees include black spruce, white spruce, jack pine,

and quaking aspen. All of these species regenerate

well on burned sites. Most of the understory plants

that occur in the Boreal Forest sprout from underground parts that can survive fire. Ahlgren (1974)

does not consider any boreal shrub species likely to

suffer substantial mortality due to burning.

Croskery and Lee (1981) examined plant regrowth

at burned and unburned sites on a large May and June

stand-replacing fire in northwestern Ontario. Existing trees, mostly black spruce and jack pine, were

killed by the fire, and aboveground parts of shrubs

and ground cover were mostly consumed. However,

regrowth began immediately. By mid-July, ground

cover in the burned area had rebounded to 50 percent

of that in the unburned area, with an average of 14

species present compared to 21 on unburned sites. In

the second growing season after fire, shrubs began to

appear on the burn. By the fifth growing season,

ground cover was 40 percent and mean height of

deciduous species was 5 feet (1.5 m). Browse biomass

was eliminated on severely burned areas for 2 years,

then became available in small amounts. By the fifth

year, browse was abundant.

Laurentian Forest

Vegetation and Fire Regimes—The Laurentian

Forest constitutes a broad ecotone between the Eastern Deciduous Forest and Boreal Forest. It contains

plant and animal species characteristic of both regions

and some species, like eastern white pine, red pine,

and red spruce, whose distributions are centered

here. The forest consists of extensive pine forest and

stands of northern hardwoods intermixed with eastern hemlock. Studies of charcoal and plant pollen in

lake sediments show that fire has influenced species

composition of the vegetation in the eastern portion

of the Laurentian Forest during much of the past

10,000 years (Green 1986).

Overall, the most common kinds of fire in the

Laurentian Forest were stand-replacement and

mixed-severity fire, although understory fires occurred

as well (Heinselman 1981). Stand-replacing fire predominated in jack pine, black spruce, and spruce-fir

forests, with fire rotations in the 50- to 100-year range

(Heinselman 1981). In red and white pine forests,

mixed-severity fires predominated. Presettlement fire

rotations in some coniferous forests were 150 to 300

years (Wein and Moore 1977). In Northern hardwood

forests, fire rotations may sometimes have exceeded

1,000 years. The proportion of early successional stand

area was small at any given time (Telfer 1993). Many

fires were large, estimated at 1,000 to 10,000 acres

(400 to 4,000 ha) (based on Heinselman 1981).

Postfire Succession—With so many species of

both boreal and southern affinities in the Laurentian

Forest, many combinations of species form in postfire

succession. Long fire rotations create extensive

stands of mature and old hardwoods (American beech,

birches, and maples). Stand-replacement fires are

followed by a flush of shrubs and saplings, including

red and sugar maple, paper and gray birch, alders,

quaking aspen and bigtooth aspen, and cherry and

shadbush species. White and red pines are also prominent, especially on sandy soils.

Early in succession, northern red oak and bur oak

often intermix with less shade-tolerant hardwoods

and pines. Pole-sized trees may be dense. Balsam fir

and red spruce invade and gradually increase in

dominance. On dry ridges, sugar maple, red maple,

American beech, and oaks eventually dominate. On

uplands, sugar maple, yellow birch, and American

beech dominate the usually long-lasting mature stage.

Eastern hemlock dominates on mesic sites with red

spruce, yellow birch, paper birch, and occasional eastern white pine. One particularly volatile combination

of species occurs in the northern Laurentian Forest

and the southeastern fringe of the Boreal Forest.

There balsam fir is a dominant species that supports

outbreaks of spruce budworm; budworm-killed forest

is highly flammable.

Eastern Ecosystems and the

Great Plains ____________________

Eastern Deciduous Forest

Vegetation and Fire Regimes—The Eastern Deciduous Forest had understory and stand-replacing

fire regimes in the centuries before settlement by

European Americans. Lightning-caused fires were

common in the mixed mesophytic hardwood forests of

the Appalachian uplands and the Mississippi Valley

(Komarek 1974). Because precipitation was plentiful

in most years, the fires usually burned small areas.

Some areas in this forest type burned frequently,

including those near the bluegrass grasslands of

Kentucky, which supported herds of bison (Komarek

1974). Historians and anthropologists now suggest

that a substantial proportion of this deciduous forest

was kept in early successional stages through shifting cultivation, firewood cutting, and extensive burning by agricultural tribes of Native Americans (Day

1953; MacCleery 1993). Annual burning in these areas
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 11

created parklike stands of large, open-grown trees, a

high proportion of which were fire-resistant oaks and

eastern white pines. These hardwood forests had little

understory and many openings.

Stand-replacement and mixed-severity fires shaped

most of the pine forests of Eastern North America,

particularly the extensive stands of eastern white and

red pine along the northern periphery of the Midwestern States and in southern Ontario (Szeicz and

MacDonald 1990; Vogl 1970) and the pitch pine and

eastern redcedar forests on the Atlantic Coastal Plain

(Wright and Bailey 1982).

Postfire Succession—Pines are common early successional species in the Eastern Deciduous Forest

(Komarek 1974). Hardwood species with vigorous

sprouting ability, especially oaks, also tend to dominate after fire. Increased prominence of oaks is one of

the most common results of disturbance in this kind of

forest (Williams 1989). Shade-intolerant species, including tuliptree and sweetgum, regenerate well on

burned land (Little 1974). Many herbaceous species

invade burned areas aggressively. In southern parts of

the region, repeated burning leads to a mixed ground

cover of grasses and legumes amid patches of trees

(Komarek 1974). Without repeated disturbance, hardwood trees reoccupy the land with oaks in the vanguard. Continued absence of fire permits Eastern

deciduous forests to be dominated by sugar maple, red

maple, eastern hemlock, and American beech.

Southeastern Forests

Vegetation and Fire Regimes—The Southeastern Pine Region extends in a great arc from eastern

Texas around the Appalachian uplands to Virginia.

The vegetation is characterized by the “southern

pines”—longleaf, slash, loblolly, shortleaf, and sand

pines (Komarek 1974; Wright and Bailey 1982). These

pine species tolerate and even depend upon fire to

different degrees, while most hardwood species in the

Southeast are suppressed by fire. Protection from fire

enables hardwood forests to develop.

The Southeastern Pine Region has a high incidence

of lightning strikes. Lightning and ignitions by aboriginal peoples caused understory fires in most

longleaf pine forests every 1 to 15 years during presettlement times (Christensen 1988; Myers 1990).

Since many of the grass and forb species associated

with these forests also depend upon frequent fires

(Frost and others 1986), cattlemen, farmers, and hunters continued burning the southern pine forests until

the widespread adoption of fire suppression practices

in the 1930s. By that time, intentional burning to

improve wildlife habitat was already recognized as a

management tool; by 1950 it was a common practice

(Riebold 1971). Longleaf pine dominated the Coastal

Plain forests (except wetlands) until the early 1900s.

Several factors, including alteration of the fire regime,

have since favored dominance by loblolly and slash

pines, which are somewhat less fire tolerant.

In eastern Oklahoma, shortleaf pine forests probably burned in large, low-severity understory fires at

intervals of about 2 to 5 years prior to fire exclusion

(Masters and others 1995).

The dominant vegetation in sand pine-scrub stands

was killed or top-killed by fire every 15 to 100 years.

One such fire burned 34,000 acres (14,000 ha) in 4 hours

(Myers 1990). Maintenance of sand pine-scrub vegetation requires these infrequent, severe fires; more

frequent fires can convert sand pine-scrub to longleaf

pine (Christensen 1988).

Postfire Succession—The overriding impact of

fire in the Southeastern Pine Region has been the

maintenance of pine forest at the expense of hardwood

forest. Relatively frequent understory fires shape a

pine forest of variable density and well developed

ground cover. Understory burning removes shrubs

and small trees as sources of mast for wildlife, but it

creates and maintains a vigorous understory of grasses,

forbs, and fire-resistant shrubs (Wright and Bailey

1982). In the absence of fire, hardwood species invade

the pine stands and deciduous forests develop. In

much of the region, these are dominated by a mixture

of oak and hickory in combination with many other

deciduous species (Eyre 1980).

Prairie Grassland

Vegetation and Fire Regimes—The primeval prairie grasslands of North America stretched from the

Gulf Coast in Texas north to central Alberta and from

Illinois to western Montana. Precipitation increases

from west to east, creating three north-south belts of

vegetation—shortgrass prairie in the West, mixed

prairie in the North and East, and tallgrass prairie in

the Central and Eastern regions. Of all natural regions of North America, the Prairie Grassland has

been most heavily impacted by human use. Tallgrass

Prairie has been almost totally converted to agriculture. Development is somewhat less in westerly parts

of the grassland. Substantial portions of the Shortgrass Prairie remain in use for cattle grazing.

The fire regime of the grasslands prior to settlement and development for agriculture was one of

stand-replacing fires on a short return interval, every

year in some areas (DeBano and others 1998; Wright

and Bailey 1982). Ignitions due to lightning were

common (Higgins 1984), and Native Americans ignited

many fires (Wright and Bailey 1982). Prairie fires

were often vast, burning into the forest margins and

preventing tree invasion of grasslands (Reichman 1987).
12 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

Postfire Succession—In prairie grasslands, burning maintains dominance by fire-adapted grasses and

forbs. Fire also maintains the productivity of grasslands, supplying fresh, nutritious vegetation that is

used by herbivores. Fire effects are strongly influenced by season of burning and moisture conditions.

Burning outside the growing season causes little change

in biomass yield or species dominance; fire during the

growing season is likely to reduce yield and change

species dominance. Postfire recovery is delayed if a

site is burned during drought or, where annuals dominate, before seed set (DeBano and others 1998).

All grassland communities are subject to invasion

by shrubs and trees in the absence of fire. Invading

species include oaks, pines, junipers, mesquite, and

aspen (Wright and Bailey 1982).

Western Forests ________________

Rocky Mountain Forest

Vegetation and Fire Regimes—The Rocky Mountain Forest Region occupies inland mountain ranges

and plateaus from New Mexico to Alberta and British

Columbia. Vegetation patterns are complex and varied due to climatic differences that arise from variation in elevation and topography and the great latitudinal extent of the region. The forests are mainly

coniferous. Important dominant trees include ponderosa pine, lodgepole pine, spruces, and firs. West of

the Rocky Mountains, from Idaho north into British

Columbia, Douglas-fir, western larch, and grand fir

are dominant tree species.

Rocky Mountain forests in past centuries had a

variety of fire regimes: understory, mixed-severity,

and stand-replacement. At low elevations, understory

fires maintained large areas of ponderosa pine and

Douglas-fir in an open, parklike structure for thousands of years prior to the 1900s. Fires on these sites

increased grass and forb production. Stand-replacing

fires and complex mixed-severity fires were common

in subalpine spruce-fir and lodgepole pine forests;

understory fires also occurred, especially on dry sites.

Presettlement mean fire return intervals in Rocky

Mountain forests ranged from less than 10 years

(Arno 1976) to more than 300 years (Romme 1980).

Forests with a multistoried structure, including dense

thickets of young conifers, were more likely to experience stand-replacing fire than open, parklike stands.

When ignition occurred in lodgepole pine forests, old

stands were more likely to burn than young stands

(Romme and Despain 1989).

Postfire Succession—Stand-replacing fires were

unusual in ponderosa pine during presettlement times

but are now more common because of increased

surface fuels and “ladder” fuels (shrubs and young

trees that provide continuous fine material from the

forest floor into the crowns of dominant trees). In

presettlement times, repeated understory fire maintained an open forest with grasses and forbs on the

forest floor and scattered patches of conifer regeneration. Fires occasionally killed large, old trees, creating

openings where the exposed mineral soil provided a

seedbed suitable for pine reproduction (Weaver 1974).

Many forests were composed of multiple patches of

even-aged trees.

Higher-elevation spruce-fir forests experience occasional stand-replacing fire. Conifer seedlings and deciduous shrubs sprout after being top-killed by fire

and dominate regrowth within a few years after fire.

Regenerating stands often produce large volumes of

browse until the tree canopy closes, 25 or more years

after fire. In the Northern Rocky Mountains, where

lodgepole pine forests are mixed with spruce-fir,

serotinous lodgepole pine cones open after being

heated by fire. Fire thus simultaneously creates a

good seedbed for pine and produces a rain of seed. The

result is quick regeneration of lodgepole pine, often in

dense stands.

Sierra Forest

Vegetation and Fire Regimes—Mixed conifer

forests occur in the Sierra Nevada of California. Important species include Douglas-fir, incense cedar,

sugar pine, white fir, and California red fir.

Sierra forests are famous for their groves of giant

sequoia trees. Understory fires typically burned these

forests at average intervals of 3 to 25 years. This fire

regime produced an open structure with a grass and

forb understory and scattered tree regeneration, similar to the structure of Rocky Mountain ponderosa

pine forests.

Sierra Nevada forests also include ponderosa pine,

with a presettlement regime of frequent understory

fire; montane forests with a complex mixture of conifer

species; and subalpine forests of lodgepole pine,

whitebark pine, and California red fir. Montane and

subalpine forests had a complex presettlement fire

regime that included infrequent understory fire, mixedseverity fire, and stand-replacement fires of all sizes

(Kilgore 1981; Taylor and Halpern 1991).

Postfire Succession—The understory fires characteristic of Sierra mixed conifer and ponderosa pine

forests maintained open structures with little accumulation of debris on the ground (Kilgore 1981; Weaver

1974). Understory fire maintained dominance by pines

and giant sequoias, with understory species including

manzanita, deerbrush, wedgeleaf ceanothus, and bitter cherry (Wright and Bailey 1982). In the absence of
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 13

fire, less fire-resistant species, including white fir and

incense cedar, invade and develop into dense, tangled

patches of young trees.

Pacific Coast Maritime Forest

Vegetation and Fire Regimes—The Pacific Coast

Maritime Forest is the most productive forest type in

the world (Agee 1993). The area is ecologically important because of the many species, including animals,

that depend on old age classes of trees. The area is

economically important because of its rapid rates of

tree growth and biomass accumulation. The Pacific

Coast region has wet winters and dry periods in the

summer. In late summer, fire danger can become high,

leading to stand-replacing crown fires with awesome

intensity as described by Weaver (1974) for the 1933

Tillamook fire in Oregon.

Sitka spruce, western hemlock, and coast redwood

dominate Pacific Coast Maritime forests. In past centuries, fires occurred infrequently in Sitka spruce and

coastal forests of western hemlock, although most

western hemlock forests show evidence that they were

initiated following fire. Inland western hemlock forests probably burned in a regime of somewhat more

frequent, mixed-severity fire. In redwood forests on

relatively dry sites, fires of all kinds—understory,

mixed-severity, and stand-replacement—were more

common, occurring as frequently as every 50 years

(Agee 1993).

Near the coast, long fire rotations in presettlement

times resulted in a large proportion, probably about

two-thirds, of the forest in mature and old age classes

at any one time. There was thus ample habitat for flora

and fauna that prefer or can survive in old growth.

Postfire Succession—Douglas-fir is important over

much of the Pacific Coast Maritime Region because it

is resistant to fire as an old tree, is able to colonize

disturbed sites, and has a life span of several hundred

years. On upland sites in the region, stand-replacing

fire can be followed by dense shrub communities

dominated by salmonberry, salal, red huckleberry,

and vine maple (Agee 1993). Even where Douglas-fir

becomes established immediately after fire, red alder

may overtop it for many years (Wright and Bailey

1982). Postfire shrubfields sometimes persist indefinitely and sometimes are replaced by shade-tolerant

conifers that regenerate beneath the shrub canopy.

Understory fires tend to eliminate most trees except

large Douglas-fir and coastal redwood, if present (Agee

1993). Shade-tolerant trees regenerate under the remaining canopy. Mixed-severity fires produce gaps in

which Douglas-fir regenerates and grows rapidly.

Where redwood grows on alluvial sites, mixed-severity fire favors development of large, old redwood trees

along with dense redwood regeneration.

Western Woodlands,

Shrublands, and Grasslands ______

Pinyon-Juniper

Vegetation and Fire Regimes—Pinyon-juniper

woodlands are dry, open forests occurring in the

Southwestern United States. Prominent overstory

species include the pinyon pines, Utah juniper, oneseed juniper, and alligator juniper. Pinyon-juniper

woodland occupies elevations between higher oak woodlands and lower grass- and shrub-dominated areas

(Wright and Bailey 1982). Because of the open nature

of pinyon-juniper woodland, grasses and shrubs are

prominent in the understory.

In presettlement times, stand-replacing fires

probably occurred at intervals averaging less than

50 years in pinyon-juniper woodlands. Because of

fire, areas with mature pinyon-juniper cover were

somewhat restricted to locations with rocky soils

and rough topography, which inhibited fire spread

(Bradley and others 1992; Kilgore 1981; Wright and

Bailey 1982). Where livestock grazing reduced herbaceous fuels, fire occurrence decreased and pinyonjuniper woodlands expanded. In mature, closed

stands, fire spreads poorly because surface fuels are

sparse. High winds and a high proportion of pine to

juniper increase the potential for fire spread (Wright

and Bailey 1982). Fire-caused tree mortality is likely

to be great where fine fuels are dense or tumbleweeds

have accumulated.

Postfire Succession—The impact of fire depends

on tree density and the amount of grass and litter in

the stand. For a few years after fire, pinyon-juniper

woodlands present a stark landscape of dead trees and

nearly bare soil. Annual plants become established in

a few years. These are followed by perennial grasses

and forbs. Invading plants often include weedy species, especially on bare soil. Junipers and shrubs

typically reestablish in 4 to 6 years. After 40 to 60

years, the shrubs are replaced by a new stand of

juniper (Barney and Frischnecht 1974; Koniak

1985). Development of a mixture of mature pinyons

and junipers can require up to 300 years (West and

Van Pelt 1987).

Chaparral and Western Oak Woodlands

Vegetation and Fire Regimes—Chaparral and

western oak woodlands include broad-leafed shrub

and tree species that are well adapted to fire. These

plant communities occur in dry mountains and foothills throughout the Southwestern United States. The

largest area is in southwestern California and the

foothills of the Sierra Nevada, where chaparral is notorious for its frequent, fast-spreading, stand-replacing
14 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

fires. Following fire, chaparral species sprout and

also establish vigorously from seed. Many species

have seed that germinates best after being heated by

fire (DeBano and others 1998). In California chaparral, stand-replacing fires have occurred every 20 to

40 years for hundreds of years (Kilgore 1981). Fires

were less frequent in Arizona chaparral, at higher

elevations in California, and on northern aspects.

Oak woodlands are characterized by species that

resprout vigorously after fire; Gambel oak dominates

many such woodlands in Utah and Colorado. Oak

woodlands had understory fire regimes with occasional stand-replacing fire in presettlement times

(Wright and Bailey 1982). Fire frequency was reduced

in areas where grazing reduced surface fuels (Bradley

and others 1992).

Postfire Succession—Annual and perennial herbs

flourish after fire in chaparral, along with seedling

and resprouting shrubs. Herbs are gradually eliminated as the dense overstory of large shrubs matures

(DeBano and others 1998).

Browse productivity in chaparral increases dramatically during the first 4 to 6 years after burning

(Wright and Bailey 1982) but declines thereafter. For

a decade or two after fire, chaparral is quite fire

resistant (Wright 1986). Burning at 20- to 30-year

intervals maintains a diverse mixture of species. If the

fire return interval is longer, sprouting species will

dominate, reducing plant species diversity.

In oak woodlands, fire either underburns or topkills the dominant species and stimulates suckering

at the bases of oaks. It thus changes the structure of

oak woodlands, stimulates other shrubs, and produces a 2- to 3-year increase in productivity of grasses

and forbs. Perpetuation of oaks and optimization of

mast are wildlife management objectives in some

locations because of widespread wildlife use of mast.

In California, acorns are eaten by nearly 100 species of

animals, including California quail, wild turkey,

deer, and bear (McDonald and Huber 1995).

Sagebrush and Sagebrush Grasslands

Vegetation and Fire Regimes—Sagebrush dominates large areas in the Western United States in

dense shrub stands and mixtures with grasslands. A

common associate is antelope bitterbrush. Grasses

and forbs are abundant. Sagebrush grasslands often intermix with forest cover, especially at higher

elevations.

In presettlement times, fires burned sagebrush

grasslands at intervals as short as 17 years and as

long as 100 years (Wright and Bailey 1982). Fire

severity in sagebrush varied, depending on the occurrence of sufficient grass and litter to carry fire. If

fuel was sufficient, fires were stand-replacing and

severe, burning through the shrub crowns. Where

fuels were sparse, fires were patchy. Varied patterns

of vegetation and seasonal differences in burning

conditions produced substantial differences in fire

severity and effects.

Cheatgrass, an exotic annual, is favored by frequent

burns, especially spring burns (Wright and Bailey

1982). Cheatgrass provides an accumulation of fine

fuel that burns readily, so it alters the fire regime in

sagebrush grasslands to much more frequent, standreplacing fire (Kilgore 1981; Knick 1999). This disturbance reduces shrub cover severely and eliminates the

patchy pattern formerly characteristic of sagebrushdominated landscapes.

Postfire Succession—Fires in sagebrush grasslands reduce woody shrub species. Big sagebrush, a

valuable wildlife browse species, is highly susceptible

to injury from fire. Its recovery depends on season

and severity of burn, summer precipitation, and frequency of burning. Big sagebrush may take more than

50 years to recover preburn dominance (Wright 1986).

Antelope bitterbrush may be killed or only top-killed

by fire, depending on the ecotype present and fire

severity (Bedunah and others 1995).

Many grass and forb species thrive after fire and

may delay regrowth of shrubs. Fires occurring every

few years reduce perennial grasses and favor annuals,

including cheatgrass. Shrubs reinvade during wet

years. Sagebrush grasslands occasionally undergo

severe droughts, which provide a major setback to

shrub vegetation even in the absence of fire (Wright

1986).

Deserts

Vegetation and Fire Regimes—North American

deserts occur in two separate areas of dry climate.

The larger of the two areas extends from Baja

California north through the Great Basin to central

Idaho and Oregon (Humphrey 1974). This large region supports three floristically distinct deserts: the

Sonoran Desert in the south, the Mojave Desert in

southeastern California and southern Nevada, and

the large Great Basin Desert to the north. The second

North American desert area is the Chihuahuan

Desert, located in the northern interior of Mexico

and southern New Mexico.

In deserts with woody plants and tall cacti, fire

severity in presettlement times depended on fuel

loading and continuity. Severe fire was possible only

after a moist, productive growing season; mixedseverity fire was more likely at other times. Fire was

most frequent and widespread in the Great Basin

Desert because of its greater shrub biomass (sagebrush) and because grass biomass was usually

sufficient to carry fire between clumps of shrubs
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 15

(Kilgore 1981). Next to the Great Basin Desert, the

Chihuahuan Desert was the most prone to fire, while

the Sonoran and Mojave Deserts only had enough

ground cover to carry fire after occasional, unusually

wet growing seasons (Humphrey 1974). In a review of

fire effects on succulent plants, Thomas (1991) estimates that intervals between fires prior to European

American settlement were as short as 3 years in some

desert grasslands and more than 250 years in dry

areas such as the Sonoran Desert.

Seasonal weather and grazing influence fire potential in deserts (Wright and Bailey 1982). A wet year

produces large quantities of grasses and forbs, which

provide fuel to carry fire. Grazing reduces these fine

fuels, thus reducing potential fire spread.

Postfire Succession—Regrowth following fire depends on the availability of moisture. If burning is

followed by a wet season, production of perennial

grasses and some forbs may increase (Wright and

Bailey 1982). In the most arid desert areas, fires may

reduce density of shrubs and cacti for 50 to 100 years

(Wright 1986). However, studies have shown substantial differences between species and also complex

interactions among available moisture, grazing, and

plant species (Cable 1967; MacPhee 1991; Wright and

Bailey 1982). Several studies report increases in exotic annual grasses, including red brome and red

stork’s bill, after fire in desert ecosystems; both frequency and intensity of fires may have increased since

the introduction of these grasses (Rogers and Steele

1980; Young and Evans 1973).

Subtropical Ecosystems _________

Florida Wetlands

Vegetation and Fire Regimes—The subtropical

region of Florida is underlain by an expanse of limestone bedrock that is almost level and barely above sea

level. Due to the area’s flat surface and high annual

rainfall—59 inches, 149 cm (Wright and Bailey 1982)—

wetland covers much of the area. Lower places in the

bedrock surface accumulate peat and support vegetation dominated by sawgrass. Where elevations

are slightly higher, fresh water swamp or wet prairie

vegetation occurs. Dry land occurs as knolls called

“hammocks,” which support mixed hardwood forest.

Despite its extensive wetlands, fire has always influenced the ecology of southern Florida.

Postfire Succession—Burning has apparently

maintained coastal marshes against mangrove invasion. Frequent fires in sawgrass kept fuel loadings low

and prevented severe fires that would consume peat

deposits. As peat accumulates, tree distribution expands out from the hammocks, increasing habitat for

terrestrial fauna and decreasing habitat for wetland

animals (Wright and Bailey 1982). Understory fires,

occurring about five times per century on the average,

maintained cypress stands by killing young hardwoods and suppressing hardwood regeneration (Ewel

1990). Severe fires after logging or draining swamps

alter successional pathways, enhancing willows and

eventual succession to hardwood forest.
16 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

Notes

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 17

L. Jack Lyon

Edmund S. Telfer

David Scott Schreiner

Chapter 3:

Direct Effects of Fire and

Animal Responses

This chapter summarizes current knowledge

about the immediate and short term (days to weeks)

effects of fire on terrestrial vertebrates: fire related

mortality, emigration, and immigration. Within these

topics, we describe fire effects mainly for two animal

classes—birds and mammals. Information regarding

reptiles, amphibians, and invertebrates is included if

available in the literature.

Most animal species respond predictably to the passage of fire (Komarek 1969; Lyon and others 1978).

These responses vary widely among species. Many

vertebrate species flee or seek refuge, but some vertebrates and many insects are attracted to burning

areas. Other behavioral responses to fire include rescuing young from burrows, approaching flames and

smoke to forage, and entering recent burns to feed on

charcoal and ash (Komarek 1969).

Injury and Mortality ______________

Despite the perception by the general public that

wildland fire is devastating to animals, fires generally

kill and injure a relatively small proportion of animal

populations. Ambient temperatures over 145 °F are

lethal to small mammals (Howard and others 1959),

and it is reasonable to assume the threshold does not

differ greatly for large mammals or birds. Most fires

thus have the potential to injure or kill fauna, and

large, intense fires are certainly dangerous to animals

caught in their path (Bendell 1974; Singer and

Schullery 1989). Animals with limited mobility living

above ground appear to be most vulnerable to firecaused injury and mortality, but occasionally even

large mammals are killed by fire. The large fires of

1988 in the Greater Yellowstone Area killed about

1 percent of the area’s elk population (Singer and

Schullery 1989). Fire effects on habitat influenced

the species’ population much more dramatically than

did direct mortality. Because of drought during the

summer of 1988 and forage loss on burned winter

range, elk mortality was high in the winter of 1988 to

1989, as high as 40 percent at one location (Singer and

others 1989; Vales and Peek 1996).

Fire may threaten a population that is already

small if the species is limited in range and mobility or

has specialized reproductive habits (Smith and Fischer

1997). The now extinct heath hen was restricted to

Martha’s Vineyard for many years before its extirpation, where scrub fires probably accelerated its demise

(Lloyd 1938).
18 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

Season of burn is often an important variable in

fauna mortality. Burning during nesting season appears to be most detrimental to bird and small mammal populations (Erwin and Stasiak 1979). Following

the burning of a reestablished prairie in Nebraska,

mortality of harvest mice in their aboveground nests

was evident, and many nests of ground-nesting birds

were found burned. Nestlings and juveniles of small

mammals are not always killed by fire, however.

Komarek (1969) observed adult cotton rats carrying

young with eyes still closed out of an area while fire

approached. While fire-caused mortality may sometimes be high for rodent species, their high reproductive potential enables them to increase rapidly in

favorable environments and disperse readily into

burned areas. Kaufman and others (1988b) describe

this pattern for deer mouse and western harvest

mouse populations in Kansas tallgrass prairie.

Birds

Fire-caused bird mortality depends on the season,

uniformity, and severity of burning (Kruse and Piehl

1986; Lehman and Allendorf 1989; Robbins and Myers

1992). Mortality of adult songbirds is usually considered minor, but mortality of nestlings and fledglings

does occur. In addition, a review by Finch and others

(1997) points out that reproductive success may be

reduced in the first postfire year because of food

reductions from spring fires. Nest destruction and

mortality of young have been reported for several

ground-nesting species, including ruffed, spruce, and

sharp-tailed grouse (Grange 1948), northern harrier

(Kruse and Piehl 1986), and greater prairie-chicken

(Svedarsky and others 1986). While eggs and young of

ground-nesting birds are vulnerable to spring fires,

long-term fire effects on bird populations depend

partly on their tendency to renest. According to a

review by Robbins and Myers (1992), wild turkeys

rarely renest if their nests are destroyed after 2 to 3

weeks of incubation, while northern bobwhite may

renest two or three times during a summer. For this

reason, many biologists consider turkeys more vulnerable to fire. A mixed-grass prairie habitat in North

Dakota was burned during the nesting season, but

69 percent of active clutches survived the fire and 37

percent eventually hatched. Nesting success was attributed in part to areas skipped by the fire as it

burned in a mosaic pattern (Kruse and Piehl 1986).

Underground nests, such as that of the burrowing owl,

are probably safe from most fires.

In forested areas, fire effects on birds depend largely

on fire severity. The young of birds nesting on the

ground and in low vegetation are vulnerable even to

understory fire during nesting season. Species nesting

in the canopy could be injured by intense surface fire

and crown fire, but this kind of fire behavior is more

common in late summer than during the nesting

season.

Mammals

The ability of mammals to survive fire depends on

their mobility and on the uniformity, severity, size,

and duration of the fire (Wright and Bailey 1982).

Most small mammals seek refuge underground or in

sheltered places within the burn, whereas large mammals must find a safe location in unburned patches or

outside the burn. Lyon and others (1978) observe that

small animals are somewhat more likely to panic in

response to fire than large, highly mobile animals,

which tend to move calmly about the periphery of a fire

(fig. 5).

Most small mammals avoid fire by using underground tunnels, pathways under moist forest litter,

stump and root holes, and spaces under rock, talus,

and large dead wood (Ford and others 1999). Not all

survive. Ver Steeg and others (1983), for instance,

found numerous dead meadow voles after an early

spring fire in Illinois grassland. Adequate ventilation

inside burrows is essential for animal survival (Bendell

1974). Burrows with multiple entrances may be better

ventilated than those with just one entrance (Geluso

and others 1986). Small mammals living in burrows

survived stand-replacing fire during summer in an

ungrazed sagebrush-bunchgrass community in southeastern Washington (Hedlund and Rickard 1981).

Most voles survived a prescribed burn in Nebraska

grassland (Geluso and others 1986). Several retreated

underground at the approach of the fire and returned

to the surface after the fire had passed, apparently

unharmed. Others remained aboveground, moving

quickly through dense vegetation to outrun the fire.

One individual sought refuge upon a raised mound of

soil created by plains pocket gophers and was adequately sheltered there from heat and flame.

Small rodents that construct surface-level nests,

such as brush rabbits, harvest mice, and woodrats

(dusky-footed, desert, and white-throated), are more

vulnerable to fire-caused mortality than deeper-nesting species, especially because their nests are constructed of dry, flammable materials (Kaufman and

others 1988b; Quinn 1979; Simons 1991). Woodrats

are particularly susceptible to fire mortality because

of their reluctance to leave their houses even when a

fire is actively burning (Simons 1991).

Direct fire-caused mortality has been reported for

large as well as small mammals, including coyote,

white-tailed deer, mule deer, elk, bison, black bear,

and moose (French and French 1996; Gasaway and

DuBois 1985; Hines 1973; Kramp and others 1983;

Oliver and others 1998). Large mammal mortality is

most likely when fire fronts are wide and fast moving,

fires are actively crowning, and thick ground smoke
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 19

occurs. Singer and Schullery (1989) report that most of

the large animals killed by the Yellowstone fires of

1988 died of smoke inhalation. Because mortality

rates of large mammals are low, direct fire-caused

mortality has little influence on populations of these

species as a whole (French and French 1996). Animal

mortality, of course, provides food for scavenger fauna

(fig. 6). The largest group of fire-killed elk in Yellowstone

National Park was monitored for several months after

it burned. Grizzly bears, black bears, coyotes, bald

Figure 5—Bison foraging and resting near burning area, Yellowstone National Park. Photo by Jeff

Henry, courtesy of National Park Service.

Figure 6—Fire-killed deer after stand-replacing fire, Yellowstone National Park. Note “whitewash”

on the deer’s flanks, evidence of use by scavenging birds. Photo courtesy of National Park Service.
20 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

eagles, golden eagles, and common ravens fed on the

carcasses (French and French 1996). According to

Blanchard and Knight (1996), the increased availability of carcasses benefited grizzly bears because drought

had reduced other food sources.

The stand-replacing and mixed-severity fires of 1988

in the Greater Yellowstone Area, which occurred mostly

in lodgepole pine-dominated forest, provided opportunities to study animal behavior during burns. Most

thoroughly studied were large mammals, including

bison, elk, bear, moose, and deer. French and French

(1996) observed no large mammals fleeing a fire, and

most appeared “indifferent” even to crowning fires.

Singer and Schullery (1989) concluded that large

mammals were sufficiently mobile to simply move

away from danger during the fires. Bison, elk, and

other ungulates grazed and rested within sight of

flames, often 100 m or less from burning trees.

Reptiles and Amphibians

According to a review by Russell and others (1999),

there are few reports of fire-caused injury to

herpetofauna, even though many of these animals,

particularly amphibians, have limited mobility. In a

review of literature in the Southeastern States,

Means and Campbell (1981) reported only one species

that may suffer substantial population losses from

fire, the eastern glass lizard. Another review (Scott

1996) mentions the box turtle as being vulnerable to

fire, but there are many reports of box turtles and

other turtle species surviving fires by burrowing into

the soil (Russell and others 1999). No dead amphibians or reptiles were found after understory burning in

a longleaf pine forest in Florida (Means and Campbell

1981). The vulnerability of snakes to fire may increase

while they are in ecdysis (the process of shedding

skin); of 68 eastern diamondback rattlesnakes marked

before a fire in Florida, the only two killed were in

mid-ecdysis.

Many reptiles and amphibians live in mesic habitat.

Woodland salamanders in the southern Appalachians,

for instance, use riparian sites and sites with plentiful, moist leaf litter. These sites are likely to burn less

often and less severely than upland sites. The resulting microsite variation within burns may account for

observations that fire has little effect on populations

of these species (Ford and others 1999). Wetlands

may provide refuge from fire, and activities such as

breeding by aquatic species may be carried out with

little interruption by fire (Russell and others 1999).

Many desert and semidesert habitats burned infrequently in past centuries because of sparse fuels.

In these areas, as in mesic sites, patchy fire spread

may protect herpetofauna from fire-caused injury and

mortality. A growing concern is the conversion of

vegetation in desert and semidesert, which burned

infrequently in past centuries, to vegetation that now

burns every few years due to invasion of weedy species

(see “Effects of Altered Fire Regimes” in chapter 4).

Animals in these ecosystems may not be adapted to

avoid fire.

Invertebrates

The vulnerability of insects and other invertebrates

to fire depends on their location at the time of fire.

While adult forms can burrow or fly to escape injury,

species with immobile life stages that occur in surface

litter or aboveground plant tissue are more vulnerable. However, aboveground microsites, such the unburned center of a grass clump, can provide protection

(Robbins and Myers 1992). Seasonality of fire no doubt

interacts with phenology for many invertebrates. Research is needed on fire effects at all stages of insect

life cycles, even though larval stages may be more

difficult to track than adult stages (Pickering 1997).

An August understory burn in South Carolina forest

reduced the soil mesofauna as measured the day after

fire, but annually burned plots had generally higher

populations of soil mesofauna than did plots that had

not been burned in 3 years or more (Metz and Farrier

1971).

Escape and Emigration __________

A second popular concept regarding animals’ response to fire is that they leave the area permanently

as soon as fire is detected. While non-burrowing mammals and most birds do leave their habitat while it is

burning, many return within hours or days. Others

emigrate because the food and cover they require are

not available in the burn. The length of time before

these species return depends on how much fire altered

the habitat structure and food supply.

Birds

Many birds leave burning areas to avoid injury.

Some return to take advantage of the altered habitat,

but others abandon burned areas because the habitat

does not provide the structure or foods that they

require to survive and reproduce. While some raptors

are attracted to fire (see “Immigration” below), others

move out of an area immediately after fire. After the

large Marble-Cone fire in California, the spotted

owls in Miller Canyon abandoned their habitat (Elliott

1985). Spotted owls in south-central Washington continued to use areas burned by understory fire but

avoided stand-replacement burns, probably because

their prey had been reduced (Bevis and others 1997).

Structural features make recent burns unsuitable
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 21

habitat for some species. Although stand-replacing

fire in a Douglas-fir forest in western Montana favored birds that feed on insects, at least one insect

feeder, the Swainson’s thrush, abandoned the burn

immediately (Lyon and Marzluff 1985), probably due

to its need for cover.

Several studies report declines in bird abundance or

species diversity in the first year or two after standreplacing fire, but few reports are available for the

months immediately following fire. After a late October fire in 1980 in coastal chaparral, California, fewer

birds of all species were seen in November. Three

months later, the bird population remained 26 percent

below average (McClure 1981). The number of bird

populations absent or declining in postfire years 1

and 2 has been reported to exceed the number of

populations remaining stable or increasing after fires

in Saskatchewan grassland (Pylypec 1991), Kansas

shrub-grassland (Zimmerman 1992), California coastal

sage scrub (Stanton 1986), and Wyoming spruce-firlodgepole pine forest (Taylor and Barmore 1980). Many

bird species return to burned habitat 2 to 3 years after

fire (Lyon and Marzluff 1985; Wirtz 1979).

Mammals

Because large mammals, such as moose and deer,

depend on vegetation for forage, bedding, cover, and

thermal protection, they abandon burned areas if fire

removes many of the habitat features they need. Thus

stand-replacing fires and understory burns that are

severe enough to top-kill shrubs and young trees seem

more likely to trigger high rates of emigration than

patchy or low-severity fires. Woodland caribou in

southeastern Manitoba avoided boreal forest burned

by stand-replacing fire in favor of bog communities,

lakes, and other unburned areas. Caribou may continue to avoid burns for 50 years or more, until lichens

become reestablished in the new forest (Schaefer and

Pruitt 1991; Thomas and others 1995). If recent burns

provide some, but not all, habitat requirements for a

species, the animals may stay near the edges of a burn.

Immediately following large, stand-replacing fires in

chapparal, Ashcraft (1979) reported mule deer grazing no farther than 300 feet (90 m) from cover.

Many small mammal species also leave burned

habitats. Based on intensive trapping results, Vacanti

and Geluso (1985) found that most voles survived a

prescribed burn in Nebraska grassland but left the

burned area until a new litter layer had accumulated,

about two growing seasons later. Possible reasons for

emigration included decreased protection from predators, decreased food availability, and more interactions among individuals. In the year after prescribed

understory burns in conifer woodland with a shrubby

understory in California, the abundance of small

mammals was almost three times greater on unburned than burned plots, even though species composition did not vary significantly between burned and

unburned areas (Blankenship 1982). Densities of western harvest mouse decreased the first year after

tallgrass prairie was burned because their aboveground

nests were destroyed and they left the area. During

the same period, deer mice increased, apparently

attracted by sparse ground cover that made seeds easy

to find. Western harvest mouse densities in the burn

increased the following spring and summer, with the

populations on unburned sites serving as sources of

dispersing individuals (Kaufman and others 1988b).

In a southwestern Idaho shadscale-winterfat community, fire reduced the abundance of small mammals in

the first postfire year. A decline in American badger

numbers on the burn accompanied the small mammal

decline (Groves and Steenhof 1988).

The effects of fire on mammal species are related to

the uniformity and pattern of fire on the landscape.

Fire has been cited by many authors as detrimental to

American marten food and habitat (see Koehler and

Hornocker 1977). However, a mixed-severity fire in an

area of lodgepole pine, spruce, and fir in northern

Idaho left a mosaic of forest types that supported a

diversity of cover and food types favorable for marten

(Koehler and Hornocker 1977). During summer and

fall, American marten feed on ground squirrels, fruits,

and insects in areas burned by stand-replacing fire.

They require dense forest during most winters, but

they use open forest during mild winters. Thus

while large, uniform burns do not favor American

marten, a mosaic of vegetation shaped in part by

recent fire may do so.

Immigration ____________________

Many animals are actually attracted to fire, smoke,

and recently burned areas. Some of the most interesting research regarding immigration in response to fire

is in the field of insect ecology. The beetles of the

subgenus Melanophila (“dark loving”), for instance,

use infrared radiation sensors to find burning trees,

where they mate and lay eggs (Hart 1998). Most birds

and mammals that immigrate in response to fire are

attracted by food resources.

Birds

A few bird species are attracted to active burns, and

many increase in the days and weeks that follow fire.

Parker (1974) reports that black vulture, northern

harrier, red-shouldered hawk, and American kestrel

were attracted to an agricultural (corn stubble) fire in

Kansas. In the Southwest, raptor and scavenger species that are attracted to fire or use recent burns for
22 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

hunting include northern harrier, American kestrel,

red-tailed hawk, red-shouldered hawk, Cooper’s hawk,

and turkey and black vultures (Dodd 1988). After the

large, severe Marble-Cone fire in California, western

screech-owls moved into the burned area (Elliott 1985).

Many species of birds were attracted to a 440-acre

(180-ha) burn on the Superior National Forest, Minnesota. About 10 weeks after the fire, the area was alive

with bird activity. Species included sparrows, American robin, barn swallow, common grackle, American

kestrel, northern flicker, common raven, hairy woodpecker, great blue heron, eastern bluebird, and blackbacked woodpecker (Stensaas 1989).

Predators and scavengers are often attracted to

burns because their food is more abundant or more

exposed than on unburned sites. During small prescribed burns in Texas bunchgrass and mesquitegrass stands, white-tailed hawks were attracted to

grasshoppers chased from cover by the fires. Turkey

vultures and crested caracaras fed on small mammals that had died in the fire (Tewes 1984). Standreplacing and mixed-severity fire in a Douglas-fir

forest in western Montana favored birds feeding on

insects (Lyon and Marzluff 1985). Immediately after

the fire, intense activity by wood-boring insects, parasites of wood borers, and predaceous flies occurred,

accompanied by “almost frenetic” feeding by warblers

and woodpeckers. In another study of grassland fire,

American kestrel and red-tailed hawk increased after

burning (Crowner and Barrett 1979). During a grassland fire in Florida, both cattle egrets and American

kestrels foraged close to the flames. Apparently the

egrets were attracted to vertebrates and invertebrates,

and the kestrels were preying exclusively on insects

as they flew out of the fire, into the wind (Smallwood

and others 1982).

Several studies show that woodpeckers are particularly attracted to burned areas. Black-backed woodpeckers are almost restricted to standing dead, burned

forests in the Northern Rocky Mountains (Caton 1996;

Hutto 1995; Lyon and Marzluff 1985) (fig. 7). Schardien

and Jackson (1978) found pileated woodpeckers foraging extensively on logs in an area in Mississippi that

had burned 2 weeks earlier; an abundant food supply

of wood-boring beetles appeared to be the primary

attraction. Woodpeckers were attracted to a standreplacement burn in coastal sage scrub, probably to

feed on insects in the fire-killed cover (Moriarty and

others 1985).

When small mammals are attracted to abundant

new growth in the months following fire, predators

and scavengers are attracted too. Abundant prey attracted golden eagles and peregrine falcons to recently

burned areas in New Mexico and southern California

(Lehman and Allendorf 1989). Following standreplacing fire in chaparral, common raptors and

ravens were studied for an increase in numbers.

Only ravens increased, probably because of increased

scavenging opportunities (Wirtz 1979). In Great Basin

and Chihuahuan Desert shrubsteppe, patchy burns

probably favor species that require perches and cover

above the ground (Bock and Bock 1990).

Mammals

Most mammals travel at least occasionally to seek

food and shelter, and some make lengthy migrations

every year. Mammal species can readily move into

burned areas. Some use burned areas exclusively, and

some use them seasonally or as part of their home

range.

Reports of mammals moving into burned areas immediately after fire are mainly anecdotal. Lloyd (1938)

describes movement of large animals into burned

areas to seek protection from insects. In California

chaparral, mountain lions are attracted to the edges

of recent burns where deer tend to congregate (Quinn

Figure 7—Male black-backed woodpecker at nest hole in

fire-killed lodgepole pine. Photo by Richard L. Hutto.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 23

1990). Crowner and Barrett (1979) report red fox

hunting in a recent burn in an Ohio field.

Many studies describe movement by large mammals to recently burned areas because of food quantity

or quality. Courtney (1989) reports migration of pronghorn to a northern mixed prairie in Alberta 2 months

after a July fire. The pronghorn fed on pricklypear

cactus, which was succulent and singed, with many of

the spines burned off. The following spring, pronghorn

moved into the burn because vegetation there began

growing approximately 3 weeks earlier than on unburned range. When the Delta caribou herd had its

calves in Alaska in 1982, the caribou preferred a

recently burned snow-free area to an unburned snowfree area and a snow-covered area (Davis and

Valkenburg 1983). Seven months after a standreplacing fire in boreal forest, northern Minnesota,

yearling moose had moved into the burn, apparently

attracted by increased forage and a low-density resident moose population (Peek 1972, 1974). Moose density increased from 0.5 per square mile a few months

after the fire to more than 2 per square mile two

growing seasons after the fire. Moose temporarily left

the area during the winter, when the forage that had

sprouted in response to fire was covered with snow

(Peek 1972).

Large mammals may move into burned habitat

simply because of familiarity with the area before fire.

A behavioral study of Alaskan moose after standreplacing and mixed-severity fire indicated that increased use of burned areas depended heavily on

prefire travel patterns and awareness by the moose

population of the area (Gasaway and others 1989).

Visibility of predators may be another reason for

large ungulates to move into burned areas. Desert

bighorn sheep abandoned areas from which fire was

excluded (Etchberger 1990). Mazaika and others (1992)

recommend prescribed burning in the Santa Catalina

Mountains, Arizona, to clear large shrubs and maintain seasonal diet quality for bighorn sheep.

Most small mammals are able to migrate readily in

response to increased food supplies, so many species

repopulate burns quickly after fire. Removal of litter

and standing dead vegetation, rather than increased

growth of vegetation, seemed to attract deer mice to

burned prairie within 5 weeks of a spring fire (Kaufman

and others 1988a). Increased food availability apparently outweighed the increased danger of predation

(Kaufman and others 1988b). After fire in Arizona

chaparral, recolonization was “rapid” for the species

that prefer grassy habitat, including voles, pocket

mice, and harvest mice (Bock and Bock 1990).

Two landscape-related aspects of fire, size and homogeneity, influence colonization and populations of

small mammals on recent burns. Research by Schwilk

and Keeley (1998) showed a positive relationship between deer mouse abundance and distance from unburned edge, perhaps in response to food provided by

postfire annual plants growing in the middle of

burned areas. The fires, which burned in California

chaparral and coastal sage scrub, left some “lightly

burned” patches in canyon bottoms. These refugia

may have enabled small mammals to colonize severely

burned sites during the first 6 months after fire (Schwilk

and Keeley 1998).

Reptiles and Amphibians

Little is known about amphibian and reptile emigration and immigration after fire. A study of low-severity

prescribed fires in hardwood-pine stands of the South

Carolina Piedmont found no evidence that herpetofauna emigrated in response to fire (Russell 1999).

Western fence lizards in chaparral take refuge under

surface objects at the time of fire; after the fire, they

invade the burned site from unburned patches

(Lillywhite and North 1974). Komarek (1969) reports

seeing southern diamondback rattlesnakes sunning

themselves in areas blackened by recent fire. Frequent lightning-season fires promote growth of the

bunchgrasses that flatwoods salamanders seek out

for laying their eggs. Fire exclusion reduces the

grasses in favor of closed slash or pond pine forest

(Carlile 1997).
24 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

Notes

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 25

L. Jack Lyon

Mark H. Huff

Edmund S. Telfer

David Scott Schreiner

Jane Kapler Smith

Chapter 4:

Fire Effects on Animal

Populations

The literature describing animals’ behavioral responses to fire, discussed in chapter 3, is limited.

Furthermore, short-term responses do not provide

insights about the vigor or sustainability of the species

in an area. Studies of animal populations and communities are more helpful in providing such insights.

Most research regarding fire effects on fauna focuses

on the population level, reporting changes in abundance and reproductive success of particular species

following fire. Population changes are the net result

of the behavioral and short-term responses discussed

in chapter 3, plus longer term responses (years to

decades).

Numerous population studies report abundance

and density of animals in relation to fire, but information on productivity and other demographic factors

may be essential for understanding population responses. Research on the threatened Florida scrub-jay

provides an example. The scrub-jay requires scrub oak

associations (myrtle, Chapman, and sand live oak,

ericaceous shrubs, and saw palmetto), often in areas

with open pine cover (less than 15 percent), where pine

densities are kept low by frequent understory fires.

The best vegetation for the jays consists of a mosaic of

different age classes of scrub, most of which have

burned within the last 20 years. Optimum scrub

height is about 4.5 feet (1.5 m), interspersed with

shorter scrub (Breininger and others in press;

Woolfenden 1973). Without fire, the oaks become too

tall and the habitat too dense for the Florida scrub-jay

because predators are not easily seen (Breininger and

others 1995). Florida scrub-jay densities in areas with

tall shrubs are sometimes greater than in areas with

optimum-height shrubs. However, jay mortality in

tall scrub exceeds reproductive success; the jay is

unable to sustain a population in tall scrub, as it can

in shorter scrub (Breininger and others in press).

Changes in Animal Populations ___

Birds

Bird populations respond to changes in food, cover,

and nesting habitat caused by fire. The season of

burning is important to birds in two ways: Fires

during the nesting season may reduce populations

more than fires in other seasons; and migratory populations may be affected only indirectly, or not at all, by

burns that occur before their arrival in spring or after

their departure in fall.

Most raptor populations are unaffected or respond

favorably to burned habitat. Fires often favor raptors by
26 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

reducing hiding cover and exposing prey populations.

When prey species increase in response to postfire

increases in forage, raptors are also favored. Dodd

(1988) describes beneficial effects from fire on populations of burrowing owl in desert grassland, sharpshinned and Cooper’s hawk in chaparral, and northern goshawk and sharp-shinned hawk in ponderosa

pine forest.

Fire effects on insect- and plant-eating bird populations depend on alterations in food and cover. The

canyon towhee, which eats insects and seeds, increased after stand-replacing fire in chaparral, foraging for food in the recent burn (McClure 1981). Wirtz

(1977) reports that swallows, swifts, and flycatchers

were more abundant over burned than unburned

chaparral during the first postfire year. California gnatcatchers in coastal sage scrub, however, require the

structure and cover provided by mature scrub. They

avoid burns for the first 4 to 5 years after fire (Beyers

and Wirtz 1997). In the northern Rocky Mountains,

Hutto (1995) found 15 bird species more abundant in

communities recently burned by stand-replacing fire

than in other habitat; most were bark-probing insect

eaters. On a site burned 19 years previously by standreplacing fire in Olympic National Park, hummingbirds were probably more abundant than anywhere

else in the area because the burn provided abundant

nectar-producing forbs and shrubs and also open space

for courtship (Huff and others 1985). After mixedseverity and stand-replacement burns in central Idaho,

lazuli buntings and chipping sparrows, both seed

eaters, were the most abundant songbirds (Saab and

Dudley 1998). Fire in marshes usually increases areas

of open water and enhances forage for shorebirds

and waterfowl (Vogl 1967; Ward 1968).

Bird nest site selection, territory establishment, and

nesting success can be affected by season of fire.

Spring burns may destroy active nests (Ward 1968).

Duck nesting success in mixed-grass prairie in North

Dakota was significantly lower in areas burned in

spring than fall (Higgins 1986). Blue-winged teal,

northern shoveler, and American wigeon showed the

lowest nesting success on spring burns. The differences were short-lived, however. Duck nesting response to fall- and spring-burned areas was similar in

the third postfire year.

Nesting success also depends on the quality of the

habitat before fire. Most birds nesting in areas

burned by stand-replacing fire in the northern Rocky

Mountains used broken-topped snags that were

present before the fire (Hutto 1995). Many species of

woodpeckers show substantial population increases

and disperse into areas burned by stand-replacing

fire (Hejl and McFadzen 1998; Hutto 1995; Saab and

Dudley 1998). After mixed-severity and standreplacement burns in central Idaho, nest abundance

for nine cavity-nesting species increased through

postfire year 4. On burned, unlogged sites, all species

had nesting success above 50 percent, and three Forest Service-sensitive species had 100 percent success

(table 1) (Saab and Dudley 1998).

Ground-dwelling bird populations are likely to be

affected by fires of any severity, whereas canopy-dwelling populations may not be affected by understory fire.

Table 1—Success of cavity-nesting species after stand-replacing and mixed-severity fires in ponderosa pine/Douglas-fir forest in

central Idaho (Saab and Dudley 1998).

No. nests/km

surveyed in 1996, Nesting success,

Species all treatments* unlogged stands Characteristics of preferred nesting habitat

Percent

Lewis’ woodpecker** 0.70 100 Highest nesting success on standard logged sites,

selected the largest, most heavily decayed snags

Hairy woodpecker 0.58 92 Highest nesting success on unlogged sites

Northern flicker 0.40 75 Highest nesting success on wildlife logged sites,

selected the largest, most heavily decayed snags

Western bluebird 0.63 60 Highest nesting success on wildlife logged sites

Mountain bluebird 0.64 56 Highest nesting success on unlogged sites

American kestrel 0.29 not reported Nested mainly on standard logged sites, selected

heavily decayed snags

European starling 0.13 100

White-headed woodpecker** 0.03 100 Selected heavily decayed snags

Black-backed woodpecker** 0.10 100 Favor unlogged sites, locations with high tree density,

selected hard snags

* 1996 was postfire year 2 for sites in mixed-severity burn, postfire year 4 for sites in stand replacing burn. Three treatments were studied: standard

salvage logged; wildlife logged (approximately 50 percent salvaged logged); and unlogged.

** Species listed as sensitive by Forest Service in Regions 1, 2, 4, or 6.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 27

After a fall fire on prairie in Saskatchewan, populations of ground-dwelling birds declined significantly.

Savannah sparrows and clay-colored sparrows, the

two most common species, were both adversely affected by the burn. These species rely on shrubs,

specifically western snowberry and silverberry, for

nesting habitat (Pylypec 1991). The year after fire, the

abundance of breeding pairs in the burn was less than

half the abundance in unburned areas. The third

postfire year, savannah sparrows had recovered to a

breeding pair abundance 68 percent of that on unburned sites, but clay-colored sparrow abundance had

not changed substantially.

Woodpeckers generally nest in snags or in the forest

canopy. Reports indicate that populations of woodpecker using forests with understory fire regimes tend

to be unaffected by underburns. Thinning from below,

designed to emulate understory fire in reducing fuels

in an old-growth forest in Oregon, did not alter use of

the site by pileated woodpeckers or Vaux’s swifts,

another bird that uses the tree canopy in old-growth

forests (Bull and others 1995). Pileated woodpeckers’

ability to use underburned sites probably depends on

fire severity. Fires that reduce logs, stumps, and snags

could have adverse effects by decreasing insect availability. The endangered red-cockaded woodpecker

inhabits open longleaf, loblolly and shortleaf pine

forests with few hardwoods in the midstory. Winter

and growing season understory fires every 2 to 5 years

are essential for retarding the development of a hardwood midstory in red-cockaded woodpecker habitat

(Carlile 1997; U.S. Department of the Interior, Fish

and Wildlife Service 1985) (fig. 8). If a hardwood

midstory does develop, the woodpecker abandons its

territory (Loeb and others 1992). The most abundant

red-cockaded woodpecker populations now occur in

areas with a long history of aggressive prescribed

burning (Costa and Escano 1989).

Bird populations may exhibit some plasticity in

relation to postfire habitat use and nest site selection.

Brewer’s sparrows and sage sparrows have been described as specifically requiring large patches of dense

sagebrush (Knick and Rotenberry 1995; Wiens and

Rotenberry 1981), but evidence from burned areas

suggests some adaptability. The Brewer’s sparrow

population declined after fire in big sagebrush in

Idaho; however, this decline was neither severe nor

long-lived (Petersen and Best 1987). Return rates of

banded male Brewer’s and male and female sage

sparrows the first 4 years after fire did not differ

between burned and unburned areas, except the second year after fire when fewer male sage sparrows

returned to the burn. The burn may have benefited

the sage sparrow population indirectly, since new

males used the burn to establish their territories. Nest

placement by Brewer’s sparrow was examined in big

sagebrush rangeland before and after a prescribed fire

in southeastern Idaho (Winter and Best 1985). Before

the burn, all nests were located in sagebrush canopies.

The prescribed fire burned about 65 percent of the

vegetation, leaving a mosaic of burned and unburned

sagebrush. After fire, there was a significant shift in

nest placement: 21 percent were placed close to the

ground. Fire may have reduced the number of tall

shrubs, influencing some sparrows to nest beneath

shrubs to obtain cover and concealment. Eastern kingbird populations in Michigan forests show an adaptable response to stand-replacing fire. In undisturbed

riparian areas, eastern kingbirds nest in woody vegetation, which provides foliage for concealment, but

they also nest successfully in the charred trunks and

branches of burned jack pines (Hamas 1983). Several

nests occurred in cupped depressions formed by embers that burned into heartwood.

Figure 8—Prescribed fire to improve red-cockaded woodpecker habitat. Fire is backing past a cavity tree on the Osceola

National Forest, Florida. Photo by Dale Wade.
28 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

Fires influence bird populations indirectly by

altering the populations of associated invertebrate

species. Chigger infestation in the bird community

increased in a chaparral stand during the months

following stand-replacing fire. Feather mites were

reduced, perhaps because silicon dioxide in the ash

killed the mites (McClure 1981).

Mammals

Most of the literature describing fire effects on

small mammal populations is from studies of standreplacement and mixed-severity fire. Like birds, mammals respond directly to fire-caused changes in cover

and food. Spring fires may impact mammal populations more than fires in other seasons because of

limited mobility of young. The species with the most

vulnerable young are small mammals, most of which

also have high reproductive rates; if postfire habitat

provides food and shelter for them, their populations

recover rapidly.

Ream (1981) summarized information in 237 references about small mammals and fire. She concluded

that populations of ground squirrels, pocket gophers,

and deer mice generally increase after stand-replacing

fire. Kaufman and others (1982) also report that the

deer mouse population increased after fire. They found

more deer mice on 1- and 2-year-old burns in tallgrass

prairie than in unburned areas. In the same study,

western harvest mice were more abundant on unburned sites. One year after stand-replacing fire in

shrub-steppe habitat in Idaho, the total number of

small mammals was lower in burned plots than in

unburned plots (Groves and Steenhof 1988), and most

of the animals in the burn were deer mice.

Rabbits, showshoe hare, red squirrel, northern flying squirrel, and voles generally avoid recent standreplacement burns, according to Ream (1981). Shrews

avoid burned areas from which most of the litter and

duff have been removed. Of 25 animal populations

common in chaparral brushlands, two were more

abundant in mature, closed chaparral than in recently

burned sites: Townsend’s chipmunk and dusky-footed

woodrat (Biswell 1989). Northern red-backed voles

avoided a stand-replacement burn in black spruce for

1 year and finally established a resident population in

postfire year 4, coinciding with the first year of berry

production in the burn (West 1982). In the first year

after stand-replacing fire in California grassland and

chaparral, populations of agile kangaroo rat, California pocket mouse, deer mouse, and California mouse

were either unchanged or greater on burned than

unburned areas. Populations of brush mouse, western

harvest mouse, and woodrat species decreased or

disappeared in burned chaparral and grasslands

(Wirtz 1977). Mixed-severity fire had little impact on

populations of small mammals in pitch pine forests of

the southern Appalachians (Ford and others 1999).

Animals that are dormant or estivating in underground burrows during and immediately after fire

are particularly well protected from direct fire effects. Populations of Townsend’s ground squirrels,

dormant below ground at the time of stand-replacing

fire in a sagebrush-grass community in southeastern

Washington, seemed unaffected by the fire (Hedlund

and Rickard 1981). Research after a stand-replacing

fire in chaparral found that the only burrowing rodents, Heerman and agile kangaroo rats, were also the

only rodents to survive in substantial numbers, probably because their burrows protected them from heat

(Quinn 1979).

Population responses of small mammals to fire are

related to fire uniformity. Most reports of woodrat

responses to fire indicate that they usually suffer

relatively high mortality because their nests are above

ground (Simons 1991). However, populations of woodrats were “unexpectedly high” in burned areas observed by Schwilk and Keeley (1998). These burns left

patches of “lightly burned” vegetation in California

chaparral and coastal sage scrub, which may have

provided refugia for woodrat populations.

Ungulate species often benefit from increased food

and nutrition on recent burns. Because ungulates are

sensitive to alterations in vegetation structure, however, their net response to fire depends on its severity

and uniformity. In Lava Beds National Monument,

northern California, mule deer populations were little

affected by fire; home ranges were neither abandoned

nor extended as a result of burning (Purcell and others

1984). Mule deer populations in chaparral burned by

stand-replacing fire often increase, benefiting from

increased availability of browse. Mule deer density in

climax chaparral was estimated at 25 per square mile,

while density in a severely burned area was 56 per

square mile (Ashcraft 1979). Fawn production the

second spring after burning was 1.15 fawns per doe

compared to 0.7 fawns per doe in climax chaparral.

Biswell (1961) reported an even more dramatic increase: deer density in chamise chaparral rose from

30 deer per square mile in unburned brush to 120 deer

per square mile the first year after stand-replacing

fire. Density decreased each year after that until it

reached preburn levels in 5 to 12 years. In contrast,

Stager and Klebenow (1987) report that mule deer

preferred pinyon-juniper stands 24 and 115 years

after stand-replacing fire to recently burned stands.

The difference may be attributable to the drier conditions in pinyon-juniper, which slow vegetation recovery from fire.

Most other large ungulates either respond neutrally

or positively to postfire changes in habitat. Elk rely on

browse in seral shrub fields during winter and use
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 29

dense, pole-sized forest heavily in fall (Irwin and Peek

1983). Moose also rely on seral shrubs in many areas,

especially where shrubfields are interspersed with

closed-canopy forest. In two areas converted from

sagebrush dominance to grassland with shrub patches,

pronghorn were present after fire but not before; they

had been absent from one site for 60 years prior to the

burn (Deming 1963; Yoakum 1980). Bison may avoid

burned areas for a short time, until regrowth of forage

begins (Moore 1972). Several studies indicate that

bison prefer foraging in recently burned areas the

summer after fire (Boyce and Merrill 1991; Shaw and

Carter 1990; Vinton and others 1993) (fig. 9). Whitetailed deer prefer to browse on recent burns if cover is

close by. Management recommendations for whitetailed deer for specific geographic regions often list a

maximum opening size or minimum distance to cover

(for example, see Ivey and Causey 1984; Keay and

Peek 1980).

Large carnivores and omnivores are opportunistic

species with large home ranges. Their populations

change little in response to fire, but they tend to thrive

in areas where their preferred prey or forage is most

plentiful—often, in recent burns. Fire has been recommended for improving black bear (Landers 1987) and

grizzly bear habitat (Hamer 1995; Morgan and others

1994) (fig. 10). In Minnesota, enough early postfire

plant communities must exist within a gray wolf

pack’s territory to support a surplus of deer, moose,

and American beaver for prey (Heinselman 1973).

American beaver populations invade streamside

habitat where fire has stimulated regrowth of aspen or

willow species (Kelleyhouse 1979; Ream 1981). Burned

areas in New York had more beaver colonies and a

higher annual occupancy than unburned areas

(Prachar and others 1988).

Fire may indirectly reduce disease rates in large

mammal populations. Following a stand-replacing fire

in spruce-lodgepole pine and bunchgrass mosaic in

Glacier National Park, Montana, bighorn sheep

tended to disperse, which may have reduced lungworm infections in the population (Peek and others

1985).

Reptiles and Amphibians

Fire-caused changes in plant species composition

and habitat structure influence reptile and amphibian

populations (Means and Campbell 1981; Russell and

others 1999). In chaparral, reptiles were more abundant in recently burned areas than in areas with

mature, dense cover. Individual populations responded

to the developing structure of the vegetation (Simovich

1979). Species that preferred open sites increased

slightly during the first 3 years after fire. During the

same time, species that used or could tolerate dense

Figure 9—Bison grazing in area converted by stand-replacing

fire from shrub-dominated to forb- and grass-dominated cover.

Photo by Jim Peaco, courtesy of National Park Service.

vegetation decreased but were not eliminated. As the

chaparral becomes a dense, mature layer, reptile abundance is likely to decrease.

Amphibians in forested areas are closely tied to

debris quantities—the litter and woody material that

accumulate slowly in the decades and centuries after

stand-replacing fire. In forests of British Columbia,

the proportion of nonmammalian vertebrates (mainly

amphibians) using woody debris was positively correlated with the length of the fire rotation (Bunnell

1995).

Many herpetofauna populations show little response

to understory and mixed-severity fire. After mixedseverity fire in pitch pine stands in the southern

Appalachian Mountains, populations of woodland salamanders were generally unchanged (Ford and others

1999). Low-intensity underburns in hardwood-pine

stands of the South Carolina Piedmont did not significantly alter species richness of herpetofauna;
30 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

amphibians were significantly more abundant on

burned plots due to greater numbers of Fowler’s toad

and red-spotted newts (Russell 1999). Although the

slash pine habitat of the flatwoods salamander in

Florida was underburned during winter, its breeding

season, the population showed no sign of decline (Means

and Campbell 1981).

In longleaf pine forests and slash pine plantations in

the Florida sandhills, the threatened gopher tortoise

(fig. 11) requires a sparse tree canopy and open, grassy

ground cover for optimum food and nesting (Carlile

1997; Means and Campbell 1981), conditions that are

provided by understory burning. Fires during the

growing season may increase nest sites and enhance

food supplies for new hatchlings (Carlile 1997). More

than 300 other species use the gopher tortoise’s burrow, including numerous arthropods, reptiles, and

amphibians, so fire effects on the tortoise impact many

other populations in the faunal community (Carlile

1997; Means and Campbell 1981; Russell and others

1999; Witz and Wilson 1991).

Figure 10—Grizzly bears foraging in lodgepole pine regeneration following stand-replacing fire, Yellowstone National Park. Photo

by Jim Peaco, courtesy of National Park Service.

A review by Russell and others (1999) explains that

fire in isolated wetlands usually increases areas of

open water and enhances vegetation structure favored

by many aquatic and semiaquatic herpetofauna.

Invertebrates

At least 40 species of arthropods are attracted to

fires (Evans 1971), alerted by stimuli including heat,

smoke, and increased levels of carbon dioxide. Many

use burned trees for breeding. When the larvae hatch,

they feed on the burned trees.

Soil protects most soil macrofauna and pupae of

many insects from fire. The level of protection depends

on depth of the organism and depth of heat penetration, which in turn depend on duff consumption (Schmid

and others 1981). Insect abundance above ground

decreases immediately after fire in prairies but then

increases as fresh, young plant growth becomes available (Robbins and Myers 1992).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 31

Effects of Altered Fire

Regimes _______________________

Understory Fire Regimes

Exclusion of fire can cause changes in faunal abundance and community composition in forests adapted

to understory fire, but studies designed to examine the

long-term effects of fire exclusion are rare. In the

Southeast, the red-cockaded woodpecker requires

longleaf pine habitat with an open midstory, maintained in past centuries by frequent understory fire.

When a dense hardwood midstory develops due to fire

exclusion, the woodpecker abandons its territory (Loeb

and others 1992). Bird populations were monitored for

15 years in a loblolly and shortleaf pine stand in

northwest Florida, comparing a site underburned annually to one from which fire had been excluded

(Engstrom and others 1984). After 15 years of fire

exclusion, the unburned plot had 20 times more trees

and less than one-third the ground cover of the annually burned plot. In the fire-excluded stand, the bird

community changed continuously in response to structural changes. Species that require open habitat disappeared within 5 years of fire exclusion. During

years 3 to 7, another group of species reached maximum numbers (common yellowthroat, indigo bunting,

Figure 11—Two gopher tortoises graze on new grass shoots after a prescribed fire. Photo by Larry Landers.

eastern towhee, white-eyed vireo, and northern cardinal). After saplings began to mature in the understory,

species associated with mesic woods were observed.

Populations of canopy-dwelling birds such as the eastern wood-pewee, great crested flycatcher, blue jay,

and summer tanager were affected little by 15 years

of succession.

Saab and Dudley (1998) hypothesize the effects of

three future fire regimes on ponderosa pine-Douglasfir forests with presettlement fire return intervals of

5 to 22 years (table 2). High intensity, stand-replacing

fires would favor seven of the 11 cavity nesting bird

species studied and would negatively affect four species. Continued fire suppression, accompanied by increasing forest density, would favor only two species.

The third possibility discussed is a combination of

silvicultural treatment and prescribed fire, which

theoretically would favor eight species and negatively

affect only two species. The table offers a framework

for testing whether management to replace a presettlement regime of frequent understory fire with a

combination of thinning and management-ignited

understory fire can produce benefits similar to those

from presettlement fire regimes to the species listed.

To assess potential changes throughout the faunal

community, such a table would need to include at least

all indicator species and species of special concern.
32 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

Stand-Replacement Fire Regimes

Fire exclusion from areas with stand-replacing

fire regimes has contributed to loss of habitat and

population declines in several raptor and predator

species. Examples include the golden eagle in the

Appalachian Mountains (Spofford 1971) and shorteared owl along the eastern grassland-forest interface

(Lehman and Allendorf 1989). A review by Nichols and

Menke (1984) explains that several raptors (red-tailed

hawks, Cooper’s hawks, sharp-shinned hawks, and

great horned owls) are more abundant in recently

burned chaparral than in unburned areas due to

greater visibility of prey.

Frequent, stand-replacing fire in presettlement times

maintained a “virtually treeless landscape” on the

Great Plains (Bidwell 1994). Fire exclusion, tree planting, and enhancement of waterways have encouraged

woodlands to develop, fragmenting the prairies. The

greater prairie-chicken, Henslow’s sparrow, and upland sandpiper all decline where habitat is fragmented

(Bidwell 1994). To increase abundance of the greater

prairie-chicken in North Dakota, Kobriger and others

(1988) recommend use of prescribed burning. To maintain nest and brood habitat for the prairie-chicken,

Kirsch (1974) recommends burning large plots (at

least 0.5 mile, 800 m, across) at 3- to 5-year intervals.

Grasslands left undisturbed for more than 10 years

are not desirable.

Prairie dog colonies once covered hundreds of

thousands of acres of the Great Plains that burned

frequently (Bidwell 1994). Prairie dogs prefer burned

to unburned areas for feeding and establishing colonies (Bone and Klukas 1990). The prairie dog is essential prey for the black-footed ferret, and prairie dog

colonies provide for needs of more than 100 other

Table 2—Predicted responses by cavity-nesting birds to three possible fire regimes compared with

the presettlement low intensity, high frequency fire regime in Idaho ponderosa pine/

Douglas-fir forests (Saab and Dudley 1998), presented as a framework of hypotheses to

be tested. + = more favorable than presettlement regime, 0 = no different, - = less favorable.

Potential new fire regime

High intensity stand- Complete fire Prescribed fire with

Bird species replacement fire suppression stand management

American kestrel + - +

Lewis’ woodpecker + - +

Red-naped sapsucker - 0 +

Downy woodpecker - 0 +

Hairy woodpecker + 0 +

Black-backed woodpecker + - 0

White-headed woodpecker - - +

Northern flicker + + -

Pileated woodpecker - + -

Western bluebird + - +

Mountain bluebird + - +

vertebrate species in some way (Scott 1996; Sharps

and Uresk 1990). Prairie dog grazing and waste alter

the soil and vegetation near colonies, favoring early

successional forb species, stimulating growth of grass

and forbs, and increasing the nitrogen content of

forage (Bidwell 1994; Sharps and Uresk 1990). Improved grass forage attracts bison, and increased forb

cover attracts pronghorn. Bison, in turn, trample the

areas where they graze (Yoakum and others 1996).

Bison impact on prairie dog colonies is reduced when

recent burns are available for grazing (Coppock and

others 1983).

Invasion by nonnative annual plants has increased

fire frequency in many semidesert ecosystems that

were characterized by stand-replacement fire regimes

in presettlement times. Exotic annuals, particularly

cheatgrass in sagebrush ecosystems, increase fuel

load and continuity. The result is increased fire frequency, followed by greater area of bare soil that is

colonized by greater numbers of exotic annuals (U.S.

Department of the Interior 1996; Whisenant 1990).

The impact of exotic annuals is exacerbated in sagebrush ecosystems because fire exclusion and overgrazing since the mid-1800s increased sagebrush dominance at the expense of native herbaceous species.

Loss of sagebrush cover and disruption of the historic

balance of shrubs, native grasses, and forbs threatens

the viability of sage grouse, sage sparrow, Brewer’s

sparrow, and sage thrasher populations (Knick and

Rotenberry 1995; Sveum and others 1998). In the

Snake River Birds of Prey National Conservation

Area, big sagebrush has declined from more than

80 percent cover in the 1800s to less than 15

percent in 1996 (U.S. Department of the Interior

1996). Areas that have burned in the last 15 years
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 33

have less than 3 percent sagebrush cover. Models

predict complete loss of shrublands in 25 to 50 years

without fire suppression in cheatgrass areas. Loss of

sagebrush is contributing to a steady decline in blacktailed jackrabbit populations and increased fluctuations in Townsend’s ground squirrel populations. Prairie falcons and golden eagles rely on these two prey

species, so increased fire frequency is reducing the

density and reproductive success of both species

(Wicklow-Howard 1989; U.S. Department of the Interior 1996). Other animals in Idaho that prey on the

Townsend’s ground squirrel—red-tailed hawks, American badgers, western rattlesnakes, and gopher

snakes—may also be affected (Yensen and others

1992).

Lodgepole pine and aspen communities in the Western States provide two examples of effects of fire

exclusion on forests with stand-replacement fire regimes. In lodgepole pine-spruce-fir forests, the most

productive period for bird communities appears to be

the first 30 postfire years. Thirteen species regularly

breed only in the first 30 years after fire. Conversely,

just two species breed exclusively in forests more than

30 years old (Taylor 1969, 1979; Taylor and Barmore

1980). Species that breed exclusively in the first 30

years after fire may be difficult to maintain in the

ecosystem without fire. Fire exclusion and postfire

salvage of dead trees after fire may reduce populations

of these species over large geographic areas.

Aspen stands provide more forage and a greater

diversity of understory plants than the spruce and fir

communities that generally replace them in the absence of fire. Fires of moderate to high severity can

regenerate aspen, but the moderate to high intensity

fire necessary to stimulate vigorous suckering of aspen is often difficult to achieve (Brown and DeByle

1982; Severson and Rinne 1990).

DeMaynadier and Hunter’s (1995) review points

out that most research on effects of fire on amphibians

and reptiles has been done in Florida. They and other

authors (Russell and others 1999) caution against

extending the results of this research to ecosystems

where frequent fire was not part of the presettlement

disturbance regime.

Mixed-Severity Fire Regimes

Not enough information is available to generalize

about effects of changing fire regimes in areas with

presettlement patterns of mixed-severity fire. Exclusion of fire from mixed-conifer and Douglas-fir forests

in the Southwest has led to increased fuel loads and

increasing risk of large, uniformly severe fire (Fiedler

and Cully 1995; Lehman and Allendorf 1989; U.S.

Department of the Interior, Fish and Wildlife Service

1995). Severe fire is likely to destroy nesting trees

and the dense forest structure required by Mexican

and California spotted owls. Prescribed understory

fire has been recommended to reduce fuels in areas

near spotted owl nest trees and to break up fuel

continuity in large areas of continuous dense forest,

reducing the likelihood of large, stand-replacing fires

in the future (Fiedler and Cully 1995; U.S. Department of the Interior, Fish and Wildlife Service 1995;

Weatherspoon and others 1992).

Grazing and fire exclusion have converted some

desert grasslands to open woodlands. This constitutes

loss of habitat for species such as pronghorn and Ord’s

kangaroo rat but increases habitat for mule deer

(Longland 1995; MacPhee 1991).

Animal Influences on Postfire

Habitat ________________________

Most of the literature regarding the relationship

between fire and fauna focuses on fire-caused changes

in vegetation and how habitat changes influence animal populations. A related topic is the effect of animal

populations on the process of postfire succession. In

this brief section, we provide a few examples of such

relationships for animals and plants native to North

America.

The jay-sized Clark’s nutcracker (fig. 12) is responsible for most whitebark pine regeneration (Tomback

1986). The nutcracker prefers to cache seed in open

sites with highly visible landmarks, conditions available within recent burns (Murray and others 1997)

(fig. 13). Tomback and others (1996) studied whitebark

pine regeneration after the 1988 fires in the Greater

Yellowstone area. Areas burned by mixed-severity

Figure 12—Clark’s nutcrackers cache seed of whitebark pines.

Unrecovered seed from these caches accounts for most

whitebark pine regeneration. Photo by Bob Keane.
34 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

and stand-replacement fire had greater whitebark

pine regeneration than did unburned sites.

Bison not only prefer burned to unburned grassland

for grazing during the growing season, they also contribute to the pattern of burning in prairie. In tallgrass

Figure 13—Whitebark pine regeneration in an area burned by

stand-replacing fire 30 years previous to the photo. Photo by

Stephen F. Arno.

prairie in northeastern Kansas, bison selected

patches with low forb cover dominated by big bluestem,

and grazed larger patches in burned than unburned

habitat (Vinton and others 1993). Ungrazed forbs in

areas adjacent to heavily grazed patches were thriving, producing greater biomass than in larger, ungrazed

portions of the study area. The increased variability in

vegetation productivity may act as feedback to fire

behavior, increasing variation in patchiness and variable severity of subsequent fires. During the centuries

before European American settlement, bison populations may have been controlled by Native American

hunting, which would have reduced the effects of

grazing on fuel continuity (Kay 1998).

Kangaroo rats and pocket mice may enhance postfire dominance of Indian ricegrass in sagebrush grassland ecosystems. These rodent species gather and

hoard large numbers of seed, with a clear preference

for Indian ricegrass. On burned sites with abundant

populations of these rodents, Indian ricegrass seed

had been deposited in scatter-hoards before fire even

though the species was not dominant. Indian ricegrass

dominated soon after fire. Six years after fire, density

of Indian ricegrass was more than tenfold greater on

burned than unburned sites (Longland 1994, 1995).

Although fire causes high mortality for antelope

bitterbrush, it also creates litter-free sites, in which

bitterbrush germination rates are high. Most antelope bitterbrush seedlings originate in rodent seed

caches, and rodents apparently retrieve fewer seed

from sites with limited cover (such as burned areas)

than from sites with better protection (Bedunah and

others 1995; Evans and others 1983).

Grazing and browsing on postfire sites, whether by

wild ungulates or domestic grazers, can alter postfire

succession. For example, if aspen is treated by fire to

regenerate the stand but then repeatedly browsed by

wildlife, it often deteriorates more rapidly than without treatment (Bartos 1998; Basile 1979). Such intense effects of feeding by large ungulates only occur

where the animal populations are food limited. Where

Native American predation kept populations of these

animals in check, such effects are unlikely (Kay 1998).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 35

Mark H. Huff

Jane Kapler Smith

Chapter 5:

Fire Effects on Animal

Communities

Many animal-fire studies depict a “reorganization”

of animal communities resulting from fire, with increases in some species accompanied by decreases in

others. Descriptions of faunal communities after fire,

however, are much less prevalent than descriptions of

population changes. The literature about fire and bird

communities is more complete than the literature

about other kinds of animals. In this chapter, we use

the literature about fire and birds to search for response patterns in the relationship between fire regime and changes in bird community composition.

The literature does not at this time provide enough

studies of mammal communities to complete a similar analysis.

Each animal species in a community is likely to

respond differently to fire and subsequent habitat

changes. To synthesize information about these responses, we modified Rowe’s (1983) classification of

plant responses to fit animal responses to fire. Rowe’s

approach was to assign to each plant species an adaptation category based on reproduction and regeneration attributes in the context of fire. Using similar

categories in our evaluation of the animal-fire literature, we classified species’ responses (not species themselves) for a given study location using observed changes

in animal abundance. Mean changes in species abundance before and the first few years after fire, or in

burned versus unburned areas, can be classified into

one of six categories (table 3). Possible community

response patterns using these six categories include:

A. Increasers predominate: A high proportion of

invader and/or exploiter responses. This pattern

represents an upward shift in abundance, especially for opportunistic species.

B. Decreasers predominate: A high proportion

of avoider and/or endurer responses. This pattern represents a downward shift in abundance

and unsuitable or poor habitat conditions for

species established prior to the burn.

C. Most populations change: An equitably high

proportion of invader and/or exploiter responses

and of avoider and/or endurer responses. This

pattern represents a small change in total abundance but a large shift in abundance of many

individual species.

D. Few populations change: A high proportion of

resister responses and a low proportion of other

responses. This pattern represents little change

in species composition and relatively minor fire

effects on the animal community.

E. Intermediate change: A high proportion of

resister, endurer, and exploiter responses; low

proportion of invader and avoider responses.
36 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

This chapter presents bird community responses to

fire according to the fire regimes as described in

chapter 1: understory, stand-replacement, and mixedseverity. Understory fire regimes occur only in forest

cover types. Stand-replacing fire regimes are divided

according to vegetation type: grassland, shrub-grassland, shrubland, and forest. Finally, we discuss

mixed-severity fires (also limited to forest types) that

leave at least 40 percent cover of large trees.

Analysis of the literature using the framework described above shows that fire effects on bird communities are related to the amount of structural change in

vegetation. In burned grasslands, bird communities

tend to return to prefire structure and composition by

postfire year 3. Postfire shrub communities are generally in flux until the shrub canopy is reestablished,

often 20 years or more after fire. In forests, understory

fire usually disrupts the bird community for 1 year or

less. Stand-replacing fire generally alters bird communities for 30 years or more. However, variation is

great. Many bird communities conform only loosely to

this pattern.

Many studies of fire effects on bird communities

report species richness or other indices of diversity.

Conserving all species is obviously essential for sustaining ecosystem patterns and processes, but maximizing diversity in a given location does not necessarily sustain the ecosystem (Telfer 1993). Bird responses

to fire in Southeastern scrub communities provide an

example. Many bird species (for instance, the Carolina

wren and northern cardinal) are negatively affected

by regimes of frequent fire in these scrub communities. Increasing fire frequency may reduce these species, thus reducing species richness. But the populations reduced by frequent fire represent forest edge

species common in Eastern North America. In contrast, increasing fire frequency favors the threatened

Florida scrub-jay and other scrub specialists, which

have a narrow geographic range and are the species

that make Florida scrub habitat unique (Breininger

and others in press). Their habitat is declining because

fire frequencies have declined, and these changes have

long-lasting effects on habitat structure even when

fires later return to the system (Duncan and others

1999).

Table 3—Classification of changes in bird abundance into response categories.

Response

 category Before fire After fire

Invader Not detected Detected (minimum number)

Exploiter Detected >50% increase

Resister Detected <50% increase or decrease

Endurer Detected >50% decrease

Avoider Detected Not detected or very low numbers

Vacillator Detected/not detected Inconsistent, wide fluctuations

Understory Fire Regimes _________

Understory fires burn beneath the tree canopy,

mostly through surface and understory fuels. Prescribed understory burns are commonly used to reduce fuel hazards and maintain open forest structure

in areas that had high-frequency, low-intensity fire

regimes in presettlement times, such as southeastern

pines and ponderosa pine (see Biswell 1989). Understory fires often disrupt the bird community during

the first postfire year, but by postfire year 2,

underburned forests are generally returning to

preburn bird community structure and composition.

The time since burn and the interval between understory fires influence fire effects on bird populations. In

oak scrub and slash pine communities along the central east coast of Florida, for example, Carolina wren

and white-eyed vireo had highest densities in areas

that had not burned for more than 10 years. Common

yellowthroat and rufous-sided towhee preferred areas

burned 4 years previously, and few shrub-dwelling

birds used understory burns less than 2 years old

(Breininger and Smith 1992). Positive correlations

between densities of shrub-dwelling birds and mean

shrub height suggest that some shrub dwellers would

decline under a regime of fire every 7 years or less.

However, this decline would not be expected if some

patches of habitat remained unburned (Breininger

and Schmalzer 1990). Much scrub occurs as patchy

mosaic within other vegetation types that have a

greater propensity to burn, so burns are naturally

patchy.

Frequent Understory Fires

Understory fires occurring at short (5- to 10-year)

intervals usually cause minor changes to vegetation composition and structure and likewise to bird

communities. Several studies have shown that many

bird species resist changes in abundance in frequently

underburned forests. Emlen (1970) reports few changes

in the bird community during the first 5 months after

understory burning in a 20-year-old slash pine forest

in Everglades National Park. The fire removed most of

the ground cover, dead grass, and litter; defoliated
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 37

most shrubs; and scorched small trees. Trees in the

middle and upper canopy were undamaged. Grass and

herbs recovered quickly. Over 70 percent of bird species responses were classified as resister, showing

little or no change in abundance. No species showed

invader responses after the fire. In southeast Arizona

ponderosa pine stands, moderately intense prescribed

understory fires (with flame lengths up to 4 feet, 1.2 m)

consumed nearly half of all snags more than 6 inches

(15 cm) dbh, resulting in a net 45 percent decrease in

potential nest trees the first year after treatment

(Horton and Mannan 1988). Cavity-nesting bird species abundances changed little, however. In contrast

to the above studies, a review by Finch and others

(1997) reports considerable community change after

“cool” understory burns in ponderosa pine. Seed eaters, timber drilling birds, and some aerial insect eaters

increased, while timber and foliage gleaners generally

decreased.

In the first 2 years after “cool” prescribed understory fires in the Black Hills, the bird community

showed mainly resister and exploiter responses (Bock

and Bock 1983). Bird abundance in postfire year 1

was nearly twice that in the unburned area, yet in

postfire year 2 abundances were similar in burned and

unburned areas. Such rapid shifts could not be explained by changes in vegetation structure or composition. Most likely, temporary, rapid increases in food

resources attracted bird species to burned areas and

resulted in a quick surge in their abundances.

The severity of understory fire affects the composition and abundance of the bird community after fire.

In loblolly pine-bottomland hardwood forests of

Alabama’s Piedmont, high-intensity understory fire

removed vegetation from the middle of the canopy

down, while low-intensity understory fires had a

“patchy” effect, with live and dead understory vegetation interspersed. Significantly more birds used the

low-intensity burns than the high-intensity burns in

the 4 months after treatment (Barron 1992). Bark,

canopy, and shrub gleaners were more abundant on

the low-intensity burn, while ground foragers were

more abundant on the high-intensity burn.

Infrequent Understory Fires

More substantial changes in forest structure and

the bird community may occur after fire in areas

with infrequent understory fire (intervals greater

than 10 years). Populations of the most common breeding birds decreased after severe understory fires in

Yosemite National Park, while less common species

increased substantially (Granholm 1982). Two understory fires were examined: a prescribed fire in

white fir-mixed conifer forest and a naturally ignited

understory fire in a California red fir forest. In

presettlement times, these forest types underburned

every 17 to 65 years (Taylor and Halpern 1991).

Trees up to 40 feet (12 m) tall were killed by the fires.

Bird communities in the two burns showed similar

responses. The highest proportion of species responses

was in the resister category. No species avoided the

burns, and more than 70 percent of the responses were

classified as resister, exploiter, or invader. Hermit

thrush and Hammond’s flycatcher populations were

reduced most by the fires, and woodpecker populations increased most.

Vegetation usually responds more slowly after fire

in dry forests, including pinyon-juniper, than in more

productive, frequently burned forests. The bird community may likewise be slow to return to its prefire

composition and structure. In pinyon-juniper forests

of Nevada, understory fires occurred in the past much

less frequently (once every several decades) than in

the southern pines and ponderosa pine (Wright and

Bailey 1982). In central Nevada, more than 60 percent

of bird species the first 2 years after prescribed understory fire showed vacillator (showing wide population

fluctuations) or exploiter responses (Mason 1981).

Species using resources near the ground increased

most after burning. Savannah and black-chinned sparrows were found only on burned areas.

Stand-Replacement Fire

Regimes _______________________

Research in the literature indicates that bird communities are disrupted for at least 2 years by standreplacing fire. A few studies show signs that the

community is returning to its preburn structure in

postfire years 3 and 4, but others do not. The changes

can be positive for insect-eating and seed-eating species and negative for species that require a dense,

closed canopy such as bark and foliage gleaners.

Grasslands

Grasslands with few or no shrubs have a relatively

simple aboveground vegetation structure, which is

consumed almost completely by fire. Vegetation change

following fire is rapid. Conditions similar to preburn

vegetation composition and structure reestablish by

postfire year 2 or 3 (for example, see Launchbaugh

1972). Although grasses dominate the vegetation,

forbs often increase in density and cover immediately

after fire, so plant diversity may be highest within

the first 2 years after fire. Bird species that nest and

use grasslands seem to be well adapted to rapid,

predictable changes in habitat characteristics associated with fires, even though such fires often remove

avian nest substrates and hiding cover.
38 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

Bird communities in a South Dakota prairie 2 to

3 months after fire showed dramatic population

changes, with a high proportion of invader, endurer,

and avoider responses (Huber and Steuter 1984). This

was the only grassland study that showed such a high

proportion of invader responses, which may be due to

the short duration of the study and the fact it was

conducted soon after fire. Upland sandpiper and western meadowlark showed substantial increases compared to unburned areas, while grasshopper sparrow

and red-winged blackbird had much lower abundances

on the burn.

Other research on postfire bird communities was

done over longer periods than the above study. During

the first 2 years after grassland fires in southeastern

Arizona, most bird populations changed, but few species abandoned or were completely new to the area

(Bock and Bock 1978). Nearly 75 percent of the species

responses were classified as vacillator, endurer, and

exploiter.

In Saskatchewan, the bird community also changed

in the first 2 years after grassland fire (Zimmerman

1992). More than half of the bird populations showed

resister responses. No responses were classified as

avoider, and only a few responses were invader and

exploiter. Abundance of key species such as claycolored and savannah sparrows were still substantially below the unburned levels in year 3, so overall

abundance was consistently lower in the burned area.

Recovery was slower than in other grasslands studied.

The cool climate and short growing season of

Saskatchewan may slow the recovery process for

some prairie species.

The same bird species may respond differently to

fire in different habitats. For example, field sparrows

in central Illinois prefer to breed in grasslands overgrown with shrubs and young deciduous trees (shrubgrassland), but they also breed in grasslands without

brush and in open woodlands (Best 1979). After burning, field sparrows used shrub-grassland more and

burned grassland less than they had during the same

period the previous year. Thus the response of field

sparrow populations in grasslands was endurer, and

the response in shrub-grasslands was exploiter. Fire

evidently caused field sparrows to use the preferred

habitat more intensively than the less-preferred

habitat.

Climatic interactions with fire and habitat suitability are not well understood, but adaptation to periodic

drought may be essential for a bird species to persist

in grass-dominated communities (Zimmerman 1992).

In average and wet years, food resources increased in

Kansas prairie after fire, yet bird abundance did not.

This indicated that the bird community was saturated

(Zimmerman 1992). When drought and fire overlapped

and resources were reduced, even drought-adapted

species decreased in abundance, although no species

disappeared from the community.

Shrub-Grasslands

We differentiate between grasslands and shrubgrasslands because grass-dominated areas with

shrubs have more complex habitat structure than

grasslands. The only shrub-grasslands discussed here

are those in which shrubs were present before fire or

in unburned areas used as controls. Shrub-grasslands

are likely to have more niches available to birds and to

recover their preburn structure more slowly after fire

than grasslands. The two 1-year studies examined

here indicate that annual burning causes substantial

changes in bird communities in shrub-grasslands.

Annual burning of a Kansas prairie for more than 10

years led to a significant decrease in bird species

richness. Annual burning maintained the prairie

with low coverage of woody vegetation, rendering it

unsuitable for woody-dependent core species and most

other species. Among the bird species present every

year, response to fire was almost 90 percent resister,

endurer, and avoider (Zimmerman 1992). Annual burning virtually eliminated habitat characteristics needed

by Henslow’s sparrow and common yellowthroat.

Most species abundances changed in response to fire

on a southwestern Florida dry shrub-grassland. In the

first postfire year, most species responses on burned

plots (with shrub cover ranging from 34 to 82 percent)

were invader and avoider, compared to plots without

fire for more than 15 years that had a closed shrub

canopy (Fitzgerald and Tanner 1992). Species showing an invader response were mostly ground feeders

(for example, Bachman’s sparrow and common grounddove), whereas shrub-dwelling species showed the

avoider response (for example, northern cardinal and

gray catbird). Burned plots provided better avian

habitat than mechanically treated plots (in which

shrubs were chopped). Birds colonized the burned

plots much sooner than the mechanically treated

plots. Shrubs killed by fire provided a more complex

habitat structure than shrubs in the mechanical treatment. Annual burning would ultimately exclude

shrubs, so the bird community response would probably resemble that after mechanical treatment.

The importance of shrubs as perches in shrubgrasslands is illustrated by a study in Kansas tallgrass

prairie (Knodel-Montz 1981). Forty artificial perches

were placed in burned and unburned prairie. Comparisons were made among plots annually burned

and unburned, with and without artificial perches.

Artificial perches on the burn were used nearly twice

as often as those on the unburned plot, although the

difference was not statistically significant. In the

unburned plot, birds seemed to prefer natural perches

to artificial ones.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 39

Shrublands

Shrublands usually occur in dry environments and

are characterized by sparse to dense shrubs with few

or no trees. Examples are the extensive sagebrush

lands of the Interior West and California chaparral.

Fuels in shrublands tend to burn rapidly. Fires typically move swiftly and are difficult to control. Most

aboveground vegetation is consumed by fire, so the

structure of the vegetation is altered dramatically.

Recovery time ranges from years to decades, depending largely on the resprouting ability of the species

burned. Bird populations often decline after shrubland

fires, but declines may be offset by populations that

rebound if fire spread is patchy, leaving some areas

unburned, and if species usually associated with grassland communities invade the burn.

Numbers of bird species and individuals were much

lower where fire burned a California coastal sage

scrub community dominated by California sagebrush

than in an unburned area nearby (Stanton 1986). This

was most noticeable the first 18 months after the fire

(Moriarty and others 1985). The fire killed all but a

few large shrubs and trees. Among the 37 bird species

observed by Stanton (excluding raptors), more than

70 percent responded as resister and endurer, and few

as avoider or invader. Significant differences in foraging activity between seasons and between burned

and unburned areas were observed. All birds except

the flycatchers spent more time actively foraging in

the unburned than in the burned area. Permanent

residents foraged more in the burned area during

spring and early summer than during the rest of the

year. Birds tended to perch rather than forage in the

burned area.

Lower bird populations also predominated after a

fire in big sagebrush in south-central Montana that

killed nearly 100 percent of the sagebrush (Bock and

Bock 1987). In postfire year 3, grass and herb cover

were much higher on the burn than in a similar

unburned area, but no recruitment of new sagebrush

was detected. Of the few species detected, responses

were mostly avoider, plus either endurer or resister.

Lark sparrow, lark bunting, and Brewer’s sparrow all

avoided the burn. During the breeding season, these

three species occupied patches of significantly more

shrub canopy in the unburned area than available

randomly. Grasshopper sparrows were classified as

endurer. The only resister response was by the western meadowlark, an adaptive grassland bird.

Because sagebrush does not sprout from underground buds after fire, sagebrush communities require several decades to establish postfire vegetation

composition and structure similar to that on unburned sites. Incomplete burning, characteristic of

sagebrush stands, appears to be important to the

development of these communities. Unburned islands

of sagebrush are important sources of sagebrush seed

after fire and retain habitat features vital to species

associated with shrubs, such as sage grouse and

Brewer’s sparrow.

In southeastern Idaho, more than 50 percent of the

species responses were classified as resister for a

postburn bird community in big sagebrush (Petersen

and Best 1987), where the prescribed fire was incomplete, killing about 50 percent of the shrubs. The first

year after fire, total bird abundance declined significantly (22 percent). In years 2 and 3, there were no

significant abundance differences between burned

and unburned areas. In year 4, significantly more

birds were detected on the burn. Species showing

resister responses may have used different parts of the

patchy postfire habitat. No species avoided the burn

during the 4 years of the study.

Nonuniform burning was used to explain bird community changes after a fire in sagebrush in northcentral Utah; 3 and 4 years after the fire, few bird

species showed appreciable declines. Bird abundance

in the burned area (with about 90 percent of aboveground vegetation burned and 80 percent of shrubs

killed) was compared to abundance in an unburned

site plowed 17 years before the study (Castrale 1982).

Total bird density and number of breeding species

were similar on the two sites. Breeding bird responses

3 to 4 years after fire were primarily exploiter and

resister, with no avoider or invader responses. Burning was associated with increases of western meadowlark, a grassland species. Brewer’s sparrow and sage

thrasher, which nest above the ground in shrubs, were

associated with unburned islands of sagebrush and

did not use the grassy portions of the burned site. If the

fire had killed all the shrubs, Brewer’s sparrow, which

can be eliminated from sagebrush habitats with chemical control of shrubs (Schroeder and Sturges 1975),

probably would have been absent.

Completeness of burn influenced fire effects in central Florida’s oak scrub (Breininger and Schmalzer

1990). During the winter and spring following a November burn, stations with more than 95 percent of

the vegetation burned had low numbers of permanent

residents, while stations with 10 to 25 percent of the

vegetation burned had bird counts similar to unburned stations.

The nature of habitat adjacent to burned shrublands sometimes influences bird community responses. Lawrence (1966) sampled bird communities

in interior California chaparral dominated by

buckbrush before a prescribed fire and 3 years afterward. Observation transects crossed chaparral, grassland, and pine-oak woodland. Most chaparral species

responses were classified as resister and endurer,

with no avoider or exploiter responses. California

quail and scrub jay declined sharply after fire.
40 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

Research in California shrublands indicates that

fire does not reduce species diversity but does alter

species composition. During the year following a

stand-replacing fire in coastal sage scrub, southern

California, the species richness of birds in the burned

area gradually increased. By the end of the first year,

species richness on the burn was 70 to 90 percent

similar to that on an adjacent unburned area (Moriarty

and others 1985). The species most abundant in the

burn were those typically associated with open areas,

whereas the species most abundant in unburned

areas typically avoid open areas.

Forests and Woodlands

Stand-replacing fires in forests and woodlands are

either severe surface fires or crown fires; more than

80 percent of the trees are top-killed or killed. The

contrasts between prefire and postfire environments

are much sharper than after understory fire, and the

time needed for the vegetation to develop structure

and composition resembling the preburn forest is

measured in decades to centuries. During this time,

many forces can alter the trajectory of succession, so

the mature forest may differ substantially from the

preburn forest. A stand-replacing fire is likely to

result in many or most of the bird species present

before fire being replaced by new species (Finch and

others 1997). Some species use habitat that occurs

only for a short time after stand-replacing fire. In

Yellowstone and Grand Teton National Parks, more

species were unique to the postfire period (1 to 17

years) than to later stages of succession (111 to 304

years after stand-replacing fire) (Taylor and Barmore

1980).

In this section we first describe bird response to

fire in the short term (less than 5 years after fire) and

then in the long-term (5 years or longer). Short-term

studies typically included control plots, either sampled

before the fire or after the fire in a similar, unburned

area. Long-term studies covered early to late stages of

vegetation succession. Some examined succession from

6 to 60 years after fire, when canopy closure occurred.

Others examined a chronosequence of similar sites at

different locations from early to late seral conditions.

Short term—The few studies available indicate

that changes in habitat characteristics caused by

stand-replacing fire cause postfire avian communities

to differ substantially in the short term from prefire

communities. High turnover occurs in the first 5 years

after stand-replacing fire. The predominant response

categories are invader and avoider. These responses

usually describe 50 to 90 percent of postfire bird

populations. Few species responses are classified as

resister after crown fire, often less than 20 percent of

the species present in the first 2 years postfire; some

studies show no resister responses to fire. This

community response to fire differs substantially from

the response generally observed in understory fire

regime types, where a high proportion of the postfire

bird community consists of resister species. Most

studies of understory fire regimes showed at least a

third of the species responses as resister, with some

over 70 percent.

In western hemlock forests of western Washington,

which has a stand-replacing fire return interval spanning several centuries, more than half the bird populations showed invader and avoider responses during

the first 2 years after a severe crown fire. The bird

community composition shifted from domination by

canopy-dwelling species to species nesting and foraging near the ground (Huff and others 1985).

Bird community response to stand-replacing fire in

ponderosa pine forests of Arizona was similar to that

in western hemlock forests (Lowe and others 1978),

even though the climate and presettlement fire regimes of the two communities differ. Nearly 60 percent

of the species responses were classified as invader and

avoider 1 year after fire.

Substantial species turnover also characterized a

dense 200-year-old spruce-fir-lodgepole pine forest in

Grand Teton National Park, Wyoming, which burned

in stand-replacing fire. More than 80 percent of bird

population responses were avoider and invader during the first 3 years postfire (Taylor and Barmore

1980). Few species showed resister responses. As in

western Washington, a shift in the bird community

from canopy dwellers to ground/brush dwellers occurred. Patterns observed nearby in Yellowstone National Park were similar (Pfister 1980). In 250-yearold lodgepole pine-spruce-fir forest, about three-fourths

of the bird community responses were classified as

invader in years 2 to 3 after stand-replacing fire. The

increased bird diversity in comparison with unburned

forest was associated with rapid changes in forest

structure and composition after the fire, which attracted several species uncharacteristic of the unburned forest.

A shift from canopy dwelling to ground- and shrubdwelling species also occurred after stand-replacing

fire in northern Minnesota. Apfelbaum and Haney

(1981) sampled birds before and after crown fire in a

73-year-old jack pine/black spruce forest. The fire

burned severely in an upland pine-dominated area

while only lightly burning the hardwood draws. The

number of breeding territories decreased by more

than half the first year after fire. Tree canopy dwellers were most abundant before the fire, while groundand shrub-dwelling species predominated afterward.

The bird community showed high species turnover;

70 percent of species responses were avoider and

invader. The black-backed woodpecker was an
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 41

important species showing the invader response, comprising about 13 percent of total bird abundance after

the fire. Ovenbird, the most important ground- and

brush-dwelling species prior to fire, avoided the burned

area, where the moss ground cover was replaced by

lush herbs and jack pine seedlings.

Long term—Oliver and others (1998) show how a

“landscape” disturbance is likely to affect bird abundance in three groups of species: those that reside in

structurally complex old-growth stands with abundant understory, those that prefer edges between

dense and open vegetation, and those that prefer open

habitat (fig. 14). The diagram reflects some patterns

reported in long-term studies of birds in forested

ecosystems, although it does not account for the complex role of fire in producing and destroying snags

(see “Snags and Dead Wood” in chapter 1). The predictions are in agreement with Finch and others’ (1997)

review of the general pattern of species change in

southwestern ponderosa pine forests, whether burned

by understory or stand-replacing fire: Granivores, tree

drilling birds, and some aerial insectivores usually

increase after fires, while tree- and foliage-gleaning

birds generally decrease. Birds more closely tied to

foliage availability, such as hermit thrush and blueheaded vireo, begin recovering as foliage volume increases in subsequent years. Finch and others (1997)

add that woodpecker abundance often peaks in the

first decade after fire, then gradually declines.

Figure 14 depicts a period early in succession after

stand-replacement fire when birds are abundant, and

also a time of transition when dominance by open- and

Figure 14—Hypothetical patterns of change in populations of

species dependent on three features of forest structure: dense

understory/old growth, edge, and open sites. Shaded areas

are discussed in the text. Adapted from Oliver and others

(1998).

edge-using species gives way to dominance by understory- and canopy-using species. Three studies of bird

community dynamics after stand-replacing fire provide some insight regarding the species and habitat

requirements that account for these changes. Research on bird community response to succession in

the long-term requires either commitment to a longterm research program or use of a chronosequence, a

series of sites similar in all characteristics except time

since fire. The former method was used for a study in

the California Sierra Nevada (Bock and Lynch 1970;

Bock and others 1978; Raphael and others 1987). The

two other studies discussed here are based on

chronosequences. These three studies indicate that (1)

early seral conditions foster high bird diversity, (2)

more bird species breed exclusively in early seral

stages than in mature forests, and (3) snags are a key

habitat feature for avian diversity and abundance.

In the Sierra Nevada study, burned and unburned

plots were established in 1966, 6 years after a large

(approximately 37,000 acres, 15,000 ha) standreplacing fire in a mixed-conifer forest dominated by

Jeffrey pine and white fir. The fire killed nearly all the

overstory and understory trees, although small pockets of trees were alive in postfire year 6. Birds were

sampled every year except one for the next 20 years.

Changes in the avian community in the burn were

primarily related to changes in vegetation structure

with succession (Raphael and others 1987). In postfire years 6 to 8, bird abundance on burned plots

was similar to that on unburned plots, but species

composition differed. Species nesting and foraging on

living trees were most abundant on unburned plots,

while species characteristic of low brush and open

ground predominated in the burned area (Bock and

Lynch 1970). Primary cavity excavators (woodpeckers) were more abundant on the burn; even higher

numbers may have been present during the 6 years

before the study was initiated. Of 32 regularly breeding species, 28 percent were unique to the burn, while

19 percent were unique to the unburned area.

Bird diversity decreased in the burn from postfire

years 8 to 15, to less than in the unburned area (Bock

and others 1978). By postfire year 15, fewer dead

snags were standing and the ground cover was more

dense, resembling a shrubfield. Six of 11 hole-nesting

species declined during this period. Species that require some open ground, such as dark-eyed juncos—

the most abundant breeding species in postfire year

8—were replaced by species indicative of shrubfields,

including fox sparrows.

Shrub cover doubled between postfire years 8 and

23, snag density declined 90 percent, and cover of herb

and grass-like species decreased significantly. At the

end of this period, large snags (more than 15 inches,

38 cm dbh) were 2.5 times more abundant in the
42 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

unburned area than in the burn. Birds that feed and

nest in shrubs increased in abundance by more than

500 percent. Woodpeckers declined steadily. At the

end of the 25 years, woodpecker abundance on the

burn was similar to that in the unburned area. At

postfire year 25, vegetation characteristic of a closedcanopy forest still had not developed in the burn.

The transition from open- to closed-site species

(postulated in fig. 14) was beginning, but the bird

community was likely to continue changing and not

likely to closely resemble either the unburned area or

the burn anytime soon.

Western hemlock forests of different ages (times

since stand-replacing fire) were sampled in western

Washington (Huff 1984; Huff and others 1985). The

ages of stands in this chronosequence were 1 to 3

years, 19, 110, 181, and 515 years. Year 19 of the sere

had the highest bird diversity and least resembled

other successional stages examined. Lowest bird diversity and abundance occurred at the 110-year-old

site where, comparatively, the tree vertical structure

was simple, snag density low, and understory composition and structure poorly developed. This stand age

may represent the transition from open to closed

structure depicted in figure 14. Huff and others (1985)

note that rate of forest reestablishment may be slower

in western hemlock forests than in the Jeffrey pinewhite fir forests of the Sierra Nevada (described above).

If so, a longer period of high diversity associated with

early seral conditions can be expected for the western

Washington sere. Once a full canopy develops in the

western hemlock sere, few changes occur in bird species composition. Because the fire return interval is

long, species composition may change relatively little

for centuries.

A large-scale examination of avian successional relationships after crown fire was conducted in

Yellowstone and Grand Teton National Parks, Wyoming, by comparing recent burns to older burns and to

areas unburned for at least 300 years (Taylor 1969,

1979; Taylor and Barmore 1980). The most obvious

changes in species composition the first few years after

fire were surges in abundance of black-backed and

northern three-toed woodpeckers. (Prior to the 1974

Waterfall Canyon fire, the black-backed woodpecker

was not even known to occur in the Grand Tetons.)

Breeding bird density in postfire years 5 to 29 was

more than 50 percent greater than in lodgepole pine

stands more than 40 years old with closed canopy. In

postfire years 5 to 25, following an influx of cavity

excavators, the number of secondary cavity nesters

increased rapidly. Two of these species, the tree swallow and the mountain bluebird, dominated the avifauna. They consistently comprised 30 percent or

more of postfire birds during the first 30 years after

fire. In the second decade after fire, they comprised

55 to 64 percent of the total bird population. By about

postfire year 30, mountain bluebirds and tree swallows started to decline at a rate that depended on the

loss of standing snags with nest cavities. During this

period, vegetation structure and succession made a

transition from shrubland to young forest.

The most important event in succession for the

postfire bird community was the transition from open

to closed canopy, which occurred between postfire

years 30 and 50. With the onset of this event, species

abundance decreased by more than 60 percent. Species characteristic of later seral stages gradually appeared as the trees got taller. From about postfire

year 50 to year 100, change in forest composition and

structure stagnated. Over the next 200 years, lodgepole pine in the canopy gave way to shade-tolerant

spruce and fir. The bird community changed little

during this 250 years, with bird abundance lower than

that in earlier successional stages. Bird density and

diversity in 300-year-old and older spruce-fir forest is

higher than in the previous 250 years.

Mixed-Severity Fire Regimes ______

Little is known about the effects of fire on bird

populations in mixed-severity fire regimes. One might

expect the bird community response to mixed-severity

fire to be intermediate between responses to understory and stand-replacement fire. Both mixed-severity and stand-replacement fire occurred in Grand

Teton National Park, Wyoming, in a 250 year old

spruce-fir forest (Taylor and Barmore 1980). Half the

species responses were invader and exploiter for the

first 3 years after fire. Some canopy-dwelling species

typical of unburned areas occurred in the mixedseverity burn but were absent from the stand-replacing burn. These included western tanager, goldencrowned kinglet, red-breasted nuthatch, and

yellow-rumped warbler. The mixed-severity burn had

less species turnover than the stand-replacement burn

in the first 2 years postfire. Almost half the species

responses to the stand-replacement fires were avoider,

yet no avoider responses were recorded in the mixedseverity burns.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 43

L. Jack Lyon

Mark H. Huff

Jane Kapler Smith

Chapter 6:

Fire Effects on Fauna at

Landscape Scales

Studies of disturbance and succession have been

a major focus of ecology over the past century (McIntosh

1985). These are studies of temporal pattern. The

study of the spatial patterns associated with temporal patterns has blossomed only since about 1990

(Turner 1990). At the landscape scale (25,000 acres,

10,000 ha or more), a complex web of interactions and

relationships unfolds. Interactions at this scale are

widely accepted as important aspects of ecosystems

(see, for example, Agee 1998; Lerzman and others

1998). However, knowledge gained at finer scales of

resolution (for example, stand or homogeneous patch)

is often difficult to apply at a landscape scale (Schmoldt

and others 1999). Including landscape considerations in management demands new approaches to

planning, analysis, and design (Diaz and Apostol

1992).

Landscapes are spatially heterogeneous, characterized by structure, function, and temporal variation

(Forman and Godron 1986). Landscape structure encompasses the spatial characteristics of biotic and

abiotic components in an area and is described by the

arrangement, size, shape, number, and kind of patches

(homogeneous units). Landscape function is defined

by interactions among biotic and abiotic components.

Temporal variation of a landscape is expressed by

changes in structure and function over time. Configuration of patches affects the occurrence and spread of

subsequent fires, so landscape-level feedback is an

important part of fire effects at landscape scales (Agee

1998).

Fire’s most obvious function in landscapes is to

create and maintain a mosaic of different kinds of

vegetation (Mushinsky and Gibson 1991). This includes size, composition, and structure of patches, as

well as connectivity (linkages and flows) among

patches. Within a large (200 mi2

, 500 km2) burn in

Alaska, Gasaway and DuBois (1985) reported substantial variation in fire severity and many unburned

patches, resulting in variation in plant mortality and

perpetuation of the mosaic nature of the landscape.

Over time, a mixture of a few large burns with many

small burns and variation within them produces

relatively small homogeneous areas. One study in

northern Manitoba reported an average stand size of

10 acres (4 ha) (Miller 1976 in Telfer 1993). Standreplacing fires in boreal forest may skip as much as

15 to 20 percent of the area within their perimeters.

The 1988 fires in the Greater Yellowstone Area, well

publicized because of their size and severe fire behavior, actually consisted of a complex patchwork containing areas burned by crown fire, areas burned by
44 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

severe surface fire, underburned sites, and unburned

areas (Rothermel and others 1994) (fig. 15, table 4).

The majority of severely burned area was within 650

feet (200 m) of unburned or “lightly burned” areas.

Landscape-scale fire effects on fauna include (1)

changes in availability of habitat patches and heterogeneity within them, (2) changes in the composition

and structure of larger areas, such as watersheds,

which provide the spatial context for habitat patches,

and (3) changes in connections among habitat patches.

During the course of postfire succession, all three of

these landscape features are in flux.

Fire changes the proportions and arrangement of

habitat patches on the landscape. When fire increases

heterogeneity on the landscape, animal species have

increased opportunities to select from a variety of

habitat conditions and successional stages. Fires often

burn with varying severity, increasing heterogeneity.

Adjacent unburned areas (which may surround or be

embedded in the burn) serve as both sinks and sources

for animal populations, and also influence animal

Figure 15—Aerial photo shows variation in fire severity over the landscape after the 1988 fires in the Greater Yellowstone

Ecosystem. Black patches were burned by crown fire. Most of these are surrounded by red and gray areas where trees were

killed by severe surface fire. Green cover represents a combination of unburned forest and areas burned by understory fire.

Photo by Jim Peaco, courtesy of National Park Service.

Table 4—Proportion of area burned at four severities within

the perimeter burned each day in the Greater

Yellowstone Area, 1988 (Turner and others 1994).

Percent of area burned daily

 Severity level June 1-July 31 Aug. 20-Sept. 15

Unburned 29.3 28.2

Underburned,

 “light” burn 18.9 14.5

Severe (stand-replacing)

 surface fire 26.6 24.4

Crown fire 25.1 32.8

emigration and immigration patterns (see Pulliam

1988). Bird diversity after stand-replacing fire may

be higher on patchy or small burns than on large,

uniform burns because the small areas are accessible

to canopy and edge species as well as species that

use open areas. A small (300 acre, 122 ha) standreplacing fire in Douglas-fir forest in western Montana
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 45

left many unburned patches. The burn attracted

wood-boring insects, woodpeckers, and warblers. The

burn itself was not used by Swainson’s thrushes, but

they remained abundant in nearby unburned areas

(Lyon and Marzluff 1985).

Two management examples show how understanding of the relationship of individual species to landscape heterogeneity can be applied. The Karner blue

butterfly (fig. 16) requires wild lupine, a forb growing

in fire-dependent oak savanna and prairie, to complete its life cycle. The larva itself (fig. 17), however, is

very sensitive to fire. To protect the butterfly at Indiana Dunes National Lakeshore, managers divide the

landscape so that every burn area contains patches

from which fire is excluded; these patches serve as

refugia from which the butterfly can repopulate the

burn (Kwilosz and Knutson 1999).

The sage grouse is sensitive to fire effects on the

arrangement of habitat components on the landscape.

Stand-replacing fire in sagebrush changes the proportions and arrangement of sage grouse habitat components. It is this arrangement that determines

whether fire benefits or damages the species. Sage

grouse use various successional stages of the sagebrush sere as lekking, nesting, brooding, and wintering grounds. Forb and insect availability are the driving factors in sage grouse productivity (Drut and

others 1994). Fires increase openings, which often

increases forb production. Fires may also enhance

the nutritional value of browse and provide new

lekking sites (Benson and others 1991; Martin 1990;

Pyle and Crawford 1996). If burns cover large tracts of

sagebrush or remove sagebrush from key wintering

areas, however, they may damage sage grouse populations (Fischer and others 1996; Gregg and others 1994;

Klebenow 1969, 1973; Welch and others 1990). Neither extensive dense sagebrush nor extensive open

areas constitute optimal habitat for the species. While

burning sometimes succeeds in restoring the balance

of plant community components in sage grouse habitat, it is accompanied by the risk of increasing

cheatgrass productivity, which may cause the area to

reburn before sagebrush recovers (Crawford 1999).

Researchers in many ecosystems recommend addressing the size and spatial arrangement of patches

in planning for specific objectives. In Southeastern

forests, Dunaway (1976) recommends interspersion of

underburned areas in longleaf pine, which have low

ground cover and provide successful foraging for

northern bobwhite chicks, with unburned areas for

Figure 16—Karner blue butterfly, an endangered species whose

larval form feeds exclusively on a the fire-dependent wild lupine.

Photo by Robert Carr, courtesy of the Michigan Chapter, The

Nature Conservancy.

Figure 17—Karner blue butterfly larva feeding on its sole food

source, the fire-dependent wild lupine. Ants protect the larvae

from predation and feed on “honeydew,” a high-sugar liquid

exuded by the larvae. Photo by Catherine Papp Herms, courtesy of the Michigan Chapter, The Nature Conservancy.
46 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

escape cover and sheltering broods. In the Western

States, Belsky (1996) suggests that a mosaic of pinyon-juniper woodland, grassland, and intermediate

seral communities would optimize biodiversity in arid

Western ecosystems. In the Southwest, Reynolds and

others (1992) quantify the proportions of a landscape

in ponderosa pine forests, characterized in presettlement times by understory fire regimes, that seem

desirable for sustaining northern goshawk populations (fig. 18). Recommendations for sustaining habitat and prey for the northern goshawk in Utah and

the Rocky Mountains include increasing the predominance of early-seral and midseral species, increasing

the numbers of large trees in the landscape, and maintaining connectivity among habitat patches (Graham

and others 1997, 1999).

Some animals require habitat that contains different features at different scales. Wright (1996) found

many patches of old-growth ponderosa pine and Douglasfir in western Montana that seemed suitable for occupation by flammulated owls, but the owls occupied

fewer than half of them. The explanation lies in the

landscape context for the patches of old growth. Occupied patches (fig. 19) were embedded in a landscape

with many grassy openings and some dense thickets of

Douglas-fir; unoccupied patches (fig. 20) were typically embedded in a landscape of closed, mature forest.

Understory fire may enhance old growth for nesting,

openings for foraging, and the landscape context for

nest sites. However, a homogeneous underburned

landscape without Douglas-fir thickets would reduce

the quality of habitat for the owl.

Nesting patterns of the Florida scrub-jay provide a

second example of habitat requirements that vary

according to scale. Optimum habitat for Florida scrubjays consists of open oak patches 10 years or more after

fire containing openings that often result from more

recent fires. Scrub-jays use these openings for caching

acorns (Breininger and others 1995). The oak patches

preferred for nesting occur within a matrix of pinescrub habitat, which is not used directly by the jay but

indirectly serves its needs by providing prey species,

enabling jays to see predatory birds from a long distance, and spreading fires into oak-dominated areas,

which often burn poorly. Management that favors

open oak without considering the more flammable

adjacent habitat can result in a loss of openings and an

increase in shrub height and tree densities, and eventually a Florida scrub-jay decline (Breininger and

others 1995).

Corridors and connectivity influence habitat use by

migratory fauna such as bison (Campbell and Hinkes

1983) and caribou (Thomas and others 1995), and for

many predators, including fisher (Powell and Zielinski

1994), lynx (Koehler and Aubry 1994) and spotted owl

(Laymon 1985; Thomas and others 1990). Connectivity is a crucial consideration for aquatic fauna as well

(Rieman and others 1997). Although research design

considerations may make it difficult to demonstrate

conclusively that wildlife corridors benefit fauna (Beier

and Noss 1998), some species definitely require landscapes with little fragmentation and high connectivity

(Bunnell 1995). Bighorn sheep in Alberta foraged in

spruce-pine forest burned 10 years previously by standreplacing fire significantly more than in unburned

areas, probably because the burned sites had a more

open structure adjacent to escape areas (Bentz and

Woodard 1988). Connectivity accounted in part for the

expansion of a bison herd in Alaska after fire. Standreplacing fire in black spruce forest produced extensive sedge-grasslands, a type that bison depend upon

for winter range (Campbell and Hinkes 1983). The

authors comment that winter range expansion was

enhanced because the burned area was contiguous

with summer range and areas used for winter range

prior to the fire, so access to burned range was relatively easy. Where black spruce and shrublands fragment sedge-grasslands, bison have difficulty accessing their winter range because of deep snowpack.

In landscapes that contained a fine-grained mosaic

of structures and age classes during presettlement

times, native fauna could readily find most kinds of

habitat. In contrast, in landscapes where large, standreplacing fires were common, fauna sometimes traveled great distances in search of habitat. Ecosystems with large, stand-replacing fire regimes and

Figure 18—Proportions of a home range or landscape desirable for sustaining goshawks in forests with understory fire

regimes (Reynolds and others 1992), example of a set of

recommended stand descriptors to be implemented at landscape scales.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 47

Figure 19—Not all habitat that seems suitable for flammulated owls at the stand level is occupied by the owl; suitability

at the landscape is also important. This photo depicts a typical landscape in western Montana where flammulated owls

were detected. Photo by Vita Wright.

Figure 20—Typical landscape where flammulated owls were not detected, western Montana. Photo by Vita Wright.
48 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

ecosystems that are now highly fragmented probably

require more attention to connectivity than areas

retaining a fine-grained mosaic. As fire and fire exclusion further alter landscapes, corridors, and entry/exit

areas near corridors, fauna that require large, continuous areas of any structure—whether early seral or

old growth—may not readily find new habitat.

Despite the tendency of natural fire regimes to

provide habitat with a variety of structures at a variety of successional stages, one cannot assume that

landscapes prior to European American settlement

were at equilibrium, even at a landscape scale (Agee

1998). Ecosystems characterized in past centuries by

infrequent large, severe fire are especially unlikely to

exhibit a steady-state age structure because large

fires have a long-term effect on the distribution of age

classes on the landscape. Examples include aspenblack spruce forests in Alberta (Cumming and others

1996) and lodgepole pine in the Greater Yellowstone

Area (Turner and others 1994). Presettlement fire

regimes are an important part of the context for

management. The spatial and temporal variability in

these regimes, though difficult to identify and apply,

may be a crucial aspect of effective management

(Lertzman and others 1998).

Figure 21—Kirtland’s warblers at nest. Photo by Betty Cotrille, courtesy of the Michigan Chapter, The

Nature Conservancy.

Effects of Altered Fire Regimes ____

Excluding fire from a landscape, unless it is being

intensively managed for fiber production, has two

major effects on animal habitat. First, it increases the

abundance and continuity of late-successional stages.

Second, it changes fuel quantities and fuel arrangement, at least for a time.

Extensive changes in habitat associated with decades of fire exclusion are most evident in areas

influenced by frequent fires during presettlement times

(Gruell and others 1982). Understory fire regimes in

southeastern forests provide one example. Fire exclusion increases dominance by less flammable vegetation, converting pine to hardwood forest (Engstrom

and others 1984). Loss of early seral structures on

sandy sites contributes to the decline of several reptile

species, including sand skinks, six-lined racerunners,

mole skinks, and central Florida crowned snakes

(Russell and others 1999).

Kirtland’s warbler/jack pine ecology in Michigan

provides another example of fire exclusion’s effects at

landscape scales. Jack pine forests were characterized

in presettlement times by relatively frequent, standreplacement fire. The Kirtland’s warbler (fig. 21)
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 49

nests on the ground in dense jack pine regeneration 5

to 24 years after stand-replacing fire or harvesting

(Mayfield 1963; Probst and Weinrich 1993). The warbler was nearly extirpated during the 1960s and 1970s

because of nest parasitism by the brown-headed cowbird, fire exclusion, and tree regeneration practices in

jack pine forests of Michigan (Mayfield 1963, 1993).

Extensive use of fire and harvesting to provide breeding habitat have kept the Kirtland’s warbler from

extinction, although uncertainty still exists about

habitat attributes that actually limit population growth

(nest sites, lower branch cover for fledglings, and

foliage volume for foraging). Habitat modeling and

management planning need to integrate habitat requirements, dynamics of disturbance and succession

over large areas, and population dynamics of the

warbler itself (Probst and Weinrich 1993).

In many Western ecosystems, landscape changes

due to fire exclusion have changed fuel quantities and

arrangement, increasing the likelihood of increased

fire size and severity (Lehman and Allendorf 1989). In

interior ponderosa pine/Douglas-fir forest, for example,

exclusion of understory fire has led to the development

of landscapes with extensive ladder fuels, nearly continuous thickets of dense tree regeneration, and large

areas of late successional forest infested with root

disease (Mutch and others 1993). These changes not

only constitute habitat loss for species that require

open old-growth stands and early seral stages; they

also may increase the likelihood of large, severe fires

in the future. Where fire exclusion has caused a shift

in species composition and fuel arrays over large

areas, subsequent fires without prior fuel modification are not likely to restore presettlement vegetation

and habitat (Agee 1998).

The effects of fire exclusion on fauna that require

late-seral and old-growth habitat originally established by fire are largely unknown. Although pileated

woodpeckers do not nest in recent stand-replacement

burns, they do prefer to nest in western larch, a fire

dependent tree, in the Northern Rocky Mountains

(McClelland 1977). If altered fire regimes reduce the

abundance of large, old western larch, they are likely

to impact the woodpecker as well. In presettlement

times, the spotted owl occupied landscapes that consisted of large areas of forest at different stages of

succession, characterized by Gaines and others (1997)

as a “very dynamic” landscape. The owl prefers oldgrowth forest within this landscape, so fire exclusion

has enhanced owl habitat, at least in some parts of

the owl’s range (Thomas and others 1990). Large,

severe fires now would reduce the species’ habitat and

reduce connectivity between remaining old-growth

stands (Thomas and others 1990). Protection of the

owl may include fuel reduction in areas adjacent to

occupied stands to reduce the likelihood of standreplacement fire.

Several ecosystems in Western North America experience more frequent fire now than they did in the past

because of invasive species. Where cheatgrass dominates areas formerly covered by large patches of sagebrush and grassland, for example, fires now occur

almost annually and shrub cover is declining. Knick

and Rotenberry (1995) report that site selection by

sage sparrow, Brewer’s sparrow, and sage thrasher is

positively correlated with sagebrush cover. In addition, sage sparrow and sage thrasher prefer large to

small patches of shrubs. The sage grouse requires

mature sagebrush as part of its habitat, so extensive

stand-replacing burns are likely to reduce its populations (Benson and others 1991). Increased fire frequency and cheatgrass cover have increased landscape-level heterogeneity by reducing sagebrush cover

and patch size, lowering the value of even remnant

sagebrush patches as habitat for native birds (Knick

1999).
50 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

Notes

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 51

L. Jack Lyon

Robert G. Hooper

Edmund S. Telfer

David Scott Schreiner

Chapter 7:

Fire Effects on Wildlife Foods

Fire’s influence on wildlife food is probably the most

thoroughly researched aspect of the relationship between fire and fauna. It has generated a vast literature showing a great variety of results. The literature

is not balanced among faunal classes or geographic

regions, since most studies focus on a species of concern to managers in a particular geographic area. To

summarize this literature, we return to the vegetation

communities described in chapter 2, organizing the

discussion according to the five geographic regions

presented there. This structure also corresponds to

the broad outline of “Effects of Fire on Flora” in the

Rainbow Series.

For each vegetation type, we summarize what is

known about changes in quantity and quality of forage

following fire. In addition, where information is available, we summarize changes in availability of seeds,

mast, and insects.

The factors that affect postfire changes in vegetation

quantity and nutritional quality include soil, vegetation type, age and structure of vegetation prior to

burning, rainfall before and after burning, severity of

the fire, season of burning, time since fire, and presettlement disturbance regime. In general, the literature regarding fire effects on wildlife food indicate

that:

• Burning sets back plant development and succession, often increasing or improving forage for

wildlife from a few years to more than 100 years,

depending on vegetation type.

• Fires usually increase habitat patchiness, providing wildlife with a diversity of vegetation

conditions from which to select food and cover.

• The biomass of forage plants usually increases

after burning in all but dry ecosystems.

• The production of seeds by grasses and legumes

is usually enhanced by annual or biennial fires.

Mast production is usually enhanced by a 5-year

or longer burning cycle.

• Burning sometimes, but not always, increases

the nutritional content and digestibility of plants.

This effect is short-lived, typically lasting only

one or two growing seasons.

• Some wildlife species select a more nutritious

diet from burned areas even though the average

nutrient content of burned plants does not differ

from that of unburned plants.

It is impossible to generalize about fire effects on

wildlife foods that apply throughout all of North

America. Furthermore, while improving and increasing food for particular wildlife species may be an
52 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

important objective, it is important that these goals

not be accomplished at the expense of ecosystem sustainability (Provencher and others 1998).

Northern Ecosystems ____________

Boreal Forest

Fire and the Quantity of Forage—During the

first two growing seasons after a mixed-severity fire

with large areas of stand-replacement in Minnesota

boreal forest, most herbaceous and low shrub species

increased rapidly in biomass. Production leveled off

during the next 3 years (Irwin 1985; Ohmann and

Grigal 1979).

Stand-replacing fire in boreal forest can greatly

increase the production of woody browse for moose

(MacCracken and Viereck 1990; Oldemeyer and others 1977; Wolff 1978). Prefire stand age and species

composition play a significant role in plant response to

fire (Auclair 1983; Furyaev and others 1983;

MacCracken and Viereck 1990; Viereck 1983). Aspen

stands that were 70 years old before stand-replacing

fire produced 10 times as much browse in the first

postburn year than did birch and spruce stands that

were 180 years old before fire. Spruce stands that were

70 years old before fire produced three times as much

browse after burning than did similar stands 180

years old before burning. The benefits of burning to

moose may peak 20 to 25 years after stand-replacing

fire (MacCracken and Viereck 1990; Oldemeyer and

others 1977) and last less than 50 years (Schwartz

and Franzmann 1989).

In boreal forests, stand-replacing fire reduces the

lichens that caribou use as forage in winter; lichens

may be reduced for up to 50 years after fire. Caribou

prefer open forests burned 150 to 250 years ago. Their

preference is related not only to abundance of food, but

also to snow cover, visibility of predators and other

herd members, and nearness to traditional travel

routes (Thomas and others 1995). Lichens decline in

old stands (200 years or more), indicating that fires of

moderate to high severity may be essential for maintaining forage for caribou in the long-term (Auclair

1983; Klein 1982; Schaefer and Pruitt 1991).

Fire and Nutritional Quality—Stand-replacing

fire in boreal forest increases the protein, phosphorus,

calcium, magnesium, and potassium content of woody

browse for moose, but this effect is probably gone by

the third growing season (MacCracken and Viereck

1990; Oldemeyer and others 1977).

Laurentian Forest

Fire and the Quantity of Forage—Quaking aspen and paper birch, two of the most important browse

plants for white-tailed deer and moose in the Northern and Eastern States, both sprout well after most

fires. Paper birch reaches peak browse production 10

to 16 years after stand-replacing fire (Safford and

others 1990). Twenty-five years after prescribed fire in

quaking aspen in northern Minnesota, aspen productivity was 111 percent of productivity on unburned

stands (Perala 1995).

Fire and Nutritional Quality—Low-intensity understory fires in aspen stands in southern Ontario

increased the levels of nitrogen, calcium, phosphorus,

magnesium, and potassium in aspen leaves the first

growing season after burning (James and Smith 1977).

The level of potassium in twigs was lower in burned

than unburned stands.

Increases in some nutrients have been reported

after severe fire. Ohmann and Grigal (1979) reported

the effects of a mixed-severity fire (with large areas of

stand-replacement) in forests of jack pine, quaking

aspen, and paper birch in northern Minnesota. Concentrations of potassium, calcium, and magnesium

increased during the first 5 years after fire, generally

exceeding levels on unburned sites. Phosphorus on

burned sites also exceeded that on unburned sites.

Nitrogen concentrations were higher on burned than

unburned sites but declined during the first five growing seasons after fire.

Eastern Ecosystems and the

Great Plains ____________________

Eastern Deciduous Forests

Fire and the Quantity of Forage—Biomass of

herbs and shrubs usually increases after fire in Eastern deciduous forests. The fire frequency needed for

maximum productivity differs among vegetation types.

Prescribed understory burns at 10- and 15-year intervals did not affect shrub or herbaceous cover in New

Jersey pine stands in comparison with unburned sites

(Buell and Cantlon 1953). As intervals between burns

decreased to 5, 4, 3, 2, and 1 years, however, the shrub

cover decreased and cover of herbaceous plants, mosses

and lichens increased. Two and three growing seasons

after late winter-early spring prescribed burns in oakhickory stands in West Virginia (Pack and others

1988), herbaceous vegetation increased, although response to burning varied considerably. Results suggested that thinning stands prior to prescribed burning was necessary to increase understory productivity.

Fire and Nutritional Quality—Many studies

from the Northeast and Midwest report increased

nutrition of wildlife foods after fire, but the duration

of increases varies. Prescribed understory burns in

April increased the July levels of crude protein in
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 53

scrub oak foliage in central Pennsylvania for 4 years

(Hallisey and Wood 1976). Crude protein also increased in blueberry foliage, but only during the first

growing season. Phosphorus, magnesium, and calcium levels were higher in scrub oak foliage, and

magnesium was higher in teaberry the first growing

season after fire. During the growing season after an

April understory burn in mixed-oak forests in Wisconsin, the concentration of nitrogen, phosphorus, and

potassium in leaves increased (Reich and others 1990).

The increase was thought to be due to increased

availability of nutrients in the soil. For most nutrients

in most plant species, the effect decreased throughout

the growing season.

Southeastern Forests

Fire and the Quantity of Forage—Several studies indicate that, in general, understory fire in Southern forests does not increase the biomass of forage

but often increases the proportion of herbage to

browse (Evans and others 1991; Stransky and Harlow

1981; White and others 1991) (fig. 22). Other studies

have found that biomass increased after burning

only under some conditions. Gilliam (1991) reported

an increase in herbaceous biomass after burning a

Figure 22—Biennial prescribed burn plots, St. Marks National Wildlife Refuge, Florida. Area in foreground is

burned every other August and dominated by runner oak, a mast-producing species. Area in background is

burned every other April and shows wiregrass flowering. This is a species favored by seed-eating animals.

Photo courtesy of Dale Wade.

pine-bluestem range in Louisiana that had not been

burned in 40 years. Prescribed burning combined with

herbicides significantly increased the amount of forage in oak-hickory stands in northeastern Oklahoma

(Thompson and others 1991). Grasses and legumes

increased after fire reduced the canopy in oak-pine

stands in Oklahoma and Arkansas (Masters and others 1993). Understory burning at 1- and 2-year intervals favored herbaceous cover, while understory burning at 3- and 4-year intervals favored a mixture of

herbs and shrubs.

Fire and Nutritional Quality—Based on a review of 16 studies of fire in Southern forests, Stransky

and Harlow (1981) propose several generalizations

about the effects of fire on plant nutrition. Burning

typically increases the crude protein and phosphorus

content of grasses, forbs, and browse the first postfire

year. Increases in nutritive quality are greatest at the

beginning of the growing season and decline rapidly,

so protein and phosphorus levels are usually similar in

burned and unburned areas by winter. Calcium content of plants after burning is highly variable among

studies. The palatability of forage generally improves

after fire, at least until growth stops and lignin

content increases. Seldom are effects of burning on
54 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

the nutritional content of plants detected after the

first growing season (Christensen 1977; Stransky and

Harlow 1981; Thill and others 1987). However, highintensity fires in the Florida Keys, oak communities,

and pine-oak communities in the Southeast have extended fire’s positive effects on plant nutrition for at

least 1 year beyond the first growing season (Carlson

and others 1993; DeWitt and Derby 1955; Thackston

and others 1982). Exceptions to the pattern of nutrient

increases after fire include reports from Florida

sandridge habitat (Abrahamson and Abrahamson

1989) and eastern Texas pine-hardwood (O’Halloran

and others 1987). These studies report no substantial

increase in plant or fungus nutrient levels after fire.

Fire and the Quantity of Seeds and Mast—Seed

and mast production generally increases after fire in

Southern forests. According to Harlow and Van Lear

(1989), seed production by legumes, grasses and

spurges is significantly greater on annually burned

areas than on areas burned less frequently. Production of berries, drupes, and pomes peaks 2 to 4 years

after burning for most of 20 species of shrubs and small

trees. A stand-replacement fire in pine and hardwood

in the mountains of Virginia greatly increased the

production of blueberries the second growing season

after burning. Production declined by year 5 but remained higher than that on unburned plots (Coggins

and Engle 1971). Season and frequency of burning

influence berry production. Waldrop and others (1987)

found that annual and biennial summer fires in

loblolly pine forests on the Coastal Plain of South

Carolina reduced the numbers of blueberry plants

after 30 years of burning, whereas winter burning did

not. According to a review by Robbins and Myers

(1992), frequent growing season burns reduce mastproducing species except runner oak and some blueberry species.

Hard mast in the Southeast is used by a variety of

birds and mammals, including northern bobwhite,

wild turkey, sapsuckers, squirrels, black bear, and

white-tailed deer. The current lack of frequent fires

(both understory and stand-replacement) in the southern Appalachians is thought to be responsible for the

replacement of oaks by other species (Van Lear 1991;

Van Lear and Watt 1993). Small oaks resprout after

fires, and large oaks have fire-resistant bark that

enables these large trees to survive fire better than

their competitors if frequently underburned. However,

when fires are excluded for long periods, competing

species such as tuliptree also develop fire-resistant

bark. These competitors can survive fire, but they

have much less potential than oaks for producing

hard mast. Thirty years of prescribed burning in the

Coastal Plain of South Carolina had no effect on the

number of mast-producing hardwoods more than 5 inches

(12.5 cm) dbh (oaks, hickories, blackgum and others)

(Waldrop and Lloyd 1991). Annual summer fires nearly

eliminated small (less than 1 inch, 2.5 cm dbh) hardwoods, but all other burning treatments produced

increases in the mast-producing species (Waldrop and

others 1987).

Recommendations for use of prescribed fire often

focus on specific wildlife-related objectives. Johnson

and Landers (1978) recommend understory fire at 3-

year intervals to optimize fruit production in open

slash pines in Georgia, with some use of longer intervals to promote mast-producing species. The animals

favored by this practice include white-tailed deer,

common gray fox, northern bobwhite, wild turkey,

raccoon, and songbirds. Hamilton (1981) recommends

understory burning pine-hardwood habitats in winter

at 5- to 10-year intervals to provide ample berries and

mast for black bears. Harlow and Van Lear (1989)

suggest that annual burning may not be desirable for

the majority of wildlife species because mast production for most shrubs and small trees peaks 2 to 6 years

after burning. However, annual burning would benefit northern bobwhite, mourning doves, some songbirds, and rodents.

Fire and Availability of Invertebrates—Fire

effects on invertebrates relate not only to preburn

conditions and fire severity but also to the life cycles

and population patterns of the specific invertebrates

studied. In the sandhills of the Florida panhandle,

longleaf pine stands with dense turkey oak and sand

live oak in the understory were underburned during

the growing season. Arthropod density and biomass

increased significantly, especially populations of grasshoppers, which constituted more than 90 percent of

the arthropod biomass (Provencher and others 1998).

Such increases are likely to benefit the northern bobwhite, a bird that feeds in the ground and herb layer,

and hawking birds such as the loggerhead shrike and

kestrels. Dunaway (1976) found that annual understory burning in longleaf pine did not increase the

number of insects available to foraging birds, but may

have provided open conditions conducive to successful

hunting by northern bobwhite chicks. In central Florida

sandhills, increased frequency of understory burning

was positively correlated with the colony density of

southern harvester ants (McCoy and Kaiser 1990).

Understory fire in loblolly pine-shortleaf pine forest

in east-central Mississippi increased invertebrates

available to northern bobwhite and turkeys for up to

3 years (Hurst 1971, 1978).

While some insects are attracted to fire and increase

rapidly in burns, fire reduces others. For example, fire

is used in the southern Appalachian Mountains to

control insects that prey on oak seedlings and mast

(Van Lear and Watt 1993).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 55

Prairie Grassland

Fire and the Quantity of Forage—In presettlement times, frequent fires in grasslands kept tree

cover in check. Studies in many regions describe the

invasion of prairie by trees in the absence of fire

(Gruell 1979; Reichman 1987; Sieg and Severson 1996).

Where prairie fires eliminate trees, fires increase the

amount of forage available to fauna simply by increasing the area covered by prairie. In addition, grassland

fire can cause early green-up of warm-season grasses,

improved seed germination, and greater production of

grasses and forbs (Hulbert 1986, 1988; Svejcar 1990).

Dramatic increases in yield during the first postfire

year have been reported for dominant prairie grasses

including prairie dropseed, big bluestem, western

wheatgrass, bluebunch wheatgrass, and Indiangrass

(Bushey 1987; Dix and Butler 1954; Hulbert 1988;

Svejcar 1990). Many studies that report increased

yield also describe some circumstances under which

yield is reduced. In general, fires followed by drought

and fires in areas with less than 11 inches (300 mm)

of summer rainfall may cause decreased forage production (Kucera 1981).

Fire and Nutritional Quality—Fire often increases the percentage of protein and minerals in

prairie grasses and shrubs, although effects vary with

season of burning (Daubenmire 1968). Forage quality

in mountain shrub and grassland communities is

enhanced by increased availability of mineral nitrogen (Hobbs and Schimel 1984). The effects of fire on

grassland nutrients interact with the effects of grazing. Grazed patches in a tallgrass prairie contained

less biomass than ungrazed patches and therefore

lost less nitrogen to volatilization by fire (Hobbs and

others 1991). The differences were substantial enough

that grazing may control whether burning causes net

increases or decreases in nitrogen on a site. Grazing

also increases heterogeneity in grasslands, contributing to patchy fuels and thus variation in fire behavior

and severity.

Fire and Availability of Invertebrates—Reed

(1997) reviewed studies of fire effects on prairie arthropod communities. She found that fire modified

these communities, and the communities continued to

change with time after fire. Prairies with fires initiated in different years and different seasons are likely

to promote species richness. Fire in oak savannas,

studied over a 30-year period, did not significantly

alter arthropod diversity (Siemann and others 1997).

Fires in Texas grasslands did not significantly alter

arthropod abundance and availability to foraging birds

(Koerth and others 1986). The density of arachnids

and insect orders on Texas grasslands, however, differed significantly between burned and unburned areas at various times of year (table 5).

Beetle abundance declines immediately after fire in

prairies but may return to prefire levels within a

month (Rice 1932). On a tallgrass prairie in Kansas,

arthropod biomass was greater on annually burned

than unburned plots; cicada nymphs were more abundant on burned than unburned plots (Seastedt and

Table 5—Effects of fire on invertebrates in a Texas grassland after a January fire (Koerth and others

1986). Groups listed were significantly (p < 0.05) more (or less) abundant on burned than

unburned areas, as indicated by density (number/m2

).

First year after fire Second year after fire

More abundant Less abundant More abundant Less abundant

 Month on burn on burn on burn on burn

April Orthoptera Hemiptera

May Hemiptera

Homoptera

Coleoptera

June Diptera Arachnidae Homoptera

July Hymenoptera Hemiptera

Arachnidae Homoptera

August Coleoptera

Arachnidae

September Hymenoptera Coleoptera

Arachnidae

October Hymenoptera Hemiptera Orthoptera

Arachnidae Arachnidae
56 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

others 1986). Cicadas respond positively to increased

root productivity on burned sites, but they are relatively immobile so their feeding is unlikely to contribute to decline of their host plants.

Western Forests ________________

Rocky Mountain Forest

Fire and the Quantity of Forage—Most studies

of fire and wildlife foods in Western forests focus on

ungulates. This research generally indicates that burning produces positive results for elk and mule deer.

During the first 5 to 10 years following stand-replacing fire, grass and forb biomass generally increases.

Grass and forb biomass decreased the first growing

season after fire in aspen stands in Wyoming but

increased the second and third growing seasons to

above preburn levels (Bartos and Mueggler 1981). On

“heavily burned” sites, grass recovered more slowly

than forbs. Forage increased three-fold after both

understory and stand-replacement fire in a ponderosa

pine forest in Arizona (Oswald and Covington 1983).

The increase persisted 9 years in underburned stands,

but grazing, perhaps combined with severe fire effects, reduced forage after 2 years in areas burned by

stand-replacing fire. Climax bunchgrass stands have

been recommended for bighorn sheep winter range,

but bighorn sheep in western Montana preferred seral

forest with elk sedge and pinegrass openings (Riggs

and Peek 1980).

Although total biomass of grasses and forbs often

increases following fire, the quantity of useable forage may actually be less on burned areas if species

composition shifts to domination by relatively unpalatable species. Prescribed understory burning failed

to improve forage in some Southwestern ponderosa

pine stands because, although herbage increased dramatically, flannel mullein, an unpalatable species,

dominated the understory after fire (Ffolliott and

Guertin 1990).

Burning brush fields in northern Idaho greatly increased the browse available to wintering elk the

following year (Leege and Hickey 1971). In British

Columbia, elk wintered primarily in postfire grass and

shrub communities, except during severe weather

when conifer stands were used (Peck and Peek 1991).

In Idaho, mule deer foraged primarily in burned habitats in winter, while white-tailed deer foraged primarily in unburned habitats (Keay and Peek 1980). An

intense prescribed fire in Douglas-fir in Idaho improved forage for mule deer and elk. The benefits were

expected to last more than 20 years (Lyon 1971).

Positive effects of fire on grazing and browse productivity generally last less than 30 years (Oswald and

Covington 1983; Pearson and others 1972).

Mixed-severity and stand-replacement fires often

increase berry-producing shrubs and their productivity 20 to 60 years after fire. These changes benefit

birds, small mammals, and bear. Increased production of forb foliage and tuberous roots after the 1988

Yellowstone fires benefited grizzly bears (Blanchard

and Knight 1996). A mathematical model predicts

increased wintering populations of elk and bison in

Yellowstone for 20 to 30 years postfire (Boyce and

Merrill 1991). Large, intense burns may be necessary

for long-term maintenance of natural forest succession patterns of some forest types and for habitat

diversity in others (Finch and others 1997). While fires

top-kill huckleberry plants and kill many whitebark

pines, two species that provide important forage for

grizzly bears, they also rejuvenate decadent huckleberry stands and prevent subalpine fir from replacing

whitebark pine in many high elevation forests (Agee

1993).

Fire and Nutritional Quality—Fires usually increase some nutrients in Rocky Mountain forests and

the pine forests of Arizona and New Mexico for 1 to 3

years (Severson and Medina 1983). Stand-replacement fall burning in Wyoming aspen stands increased

crude protein and phosphorus of forage during the

first summer after treatment (DeByle and others 1989).

In vitro dry matter digestibility was also higher in

burned areas, and calcium content was lower. By late

summer, only crude protein levels were different and,

in the second postfire year, forage quality was similar

on burned and unburned areas. Burning improves the

nutritional qualities of forage plants in ponderosa

pine forest for one to three growing seasons (Meneely

and Schemnitz 1981; Pearson and others 1972;

Rowland and others 1983). In western larch/Douglasfir stands in Montana that had been burned with

understory fire 3 years previously, nutrient content of

plants was compared with samples from stands not

burned for 70 years (Stark and Steele 1977). Sodium

levels were higher for several species in stands where

at least half of the duff was consumed by fire. Iron

concentration was significantly greater in some species on burned than unburned sites, and calcium and

phosphorus were significantly lower. The plant species tested showed no significant differences in nitrogen, magnesium, or copper between burned and unburned sites. Scouler’s willow in underburned

ponderosa pine/Douglas-fir forests in Montana contained higher concentrations of phosphorus and crude

protein, and lower lignin concentration, than willows

in unburned stands (Bedunah and others 1995).

Some research reports no significant changes in

nutrient levels after fire. Seip and Bunnell (1985)

found no differences in the nutritive quality of forage

on frequently burned alpine range and unburned
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 57

range used by Dall’s sheep in British Columbia. The

authors thought that sheep on burned range were in

better physical condition than those on unburned

range because of the quantity of forage rather than its

nutritive quality. Stand-replacing prescribed fire in

Idaho aspen forests in September produced little change

in the nutritive content of forage the first, second, and

fourth growing seasons after burning (Canon and

others 1987). However, elk preferred to forage in the

burned areas, possibly because preferred species were

consistently available and foraging was more efficient.

Sierra Forest

Fire and the Quantity of Forage and Seed—

Wildlife forage species in Sierra Nevada forests include many plants that dominate in chaparral to the

west and more mesic forests to the north. Deerbrush

and greenleaf manzanita are chaparral species but are

also important components of the understory of Sierra

forests. Forage of deerbrush and other Ceanothus

species, which is high quality food for ungulates

(Sampson and Jesperson 1963; Stubbendieck and others 1992), is abundant after fire because it reproduces

from seed that is scarified by burning (Burcham 1974).

Early spring burning in the Sierra Nevada increases

palatability of foliage for wildlife (Kauffman and Martin 1985). Thimbleberry is an understory species characteristic of mesic Sierra forests; it generally increases

after fire (Hamilton and Yearsley 1988).

Pacific Coast Maritime Forest

Fire and the Quantity of Forage—Salmonberry,

an important understory species in Pacific Coast forests, is used by numerous wildlife species. Deer, elk,

mountain goats, and moose browse on its buds and

twigs; songbirds, gallinaceous birds, bears, and coyote

feed on its fruit. Salmonberry sprouts prolifically and

grows rapidly in the first years after fire, although

severe fire may reduce sprouting (Tappeiner and others 1988; Zasada and others 1989).

Western Woodlands, Shrublands,

and Grasslands _________________

Pinyon-Juniper

Fire and the Quantity of Forage—Severson and

Medina (1983) and Severson and Rinne (1990) review

the effects of fire on forage production and wildlife

habitat in the Southwest. While they demonstrate the

important role of fire in improving Southwestern vegetation types for wildlife, they emphasize the need for

a balance between burned and unburned areas. Fire

intensity varies greatly in pinyon-juniper woodlands,

and the early successional effects of fires are difficult

to predict (Severson and Rinne 1990). Often, fire may not

have much effect unless combined with other treatments (Wittie and McDaniel 1990). When conditions

are favorable for stand-replacing fire, burning kills

most of the pinyon-juniper overstory and increases

diversity in the plant community, with some effects

lasting up to 115 years after fire (McCulloch 1969;

Severson and Medina 1983; Severson and Rinne 1990;

Stager and Klebenow 1987). Shortly after fire, burns

are usually dominated by forbs, with grasses becoming abundant a few years later. In an Ashe’s juniper

community burned during a moist winter and spring,

grasses recovered quickly and soil erosion was minimal (Wink and Wright 1973). Similar treatments

during a dry winter and spring, however, reduced

herbaceous yields and increased erosion.

Chaparral and Western Oak Woodlands

Fire and the Quantity of Forage—Intense fires

in chaparral result in a flush of herbaceous plants

and shrubs for 1 to 5 years (Biswell 1974; Christensen

and Muller 1975; Klinger and others 1989; Taber

and Dasmann 1958). In Gambel oak rangeland in

Colorado, fire did not significantly change the biomass

of forbs and shrubs 2, 5, and 10 years after fall mixedseverity fire, but grass biomass was greater on burned

than unburned sites during postfire year 10 (Kufeld

1983).

Fire and Nutritional Quality—Most studies of

postfire nutrients in Western ecosystems report

some changes, but the plant species and the nutrients

affected vary. Stand-replacing fires in chaparral increased the protein content of leaves for one to two

growing seasons and the phosphorus content for up to

6 years (Rundel and Parsons 1980; Taber and Dasmann

1958). Two growing seasons after fall mixed-severity burns in Gambel oak rangeland in Colorado, zinc

and copper levels were higher in plants on burned

than unburned sites. However, no differences were

found in the protein, lignin, calcium, or phosphorus

content of forbs, grasses or shrubs growing on burned

and unburned areas (Kufeld 1983).

Where postfire nutrient changes vary among the

plant species available to fauna, animals may select

the more nutritious foods. September prescribed burns

in mountain shrub and grassland habitats in Colorado

increased the level of protein and in vitro digestible

organic matter in winter diets of bighorn sheep and

mule deer (Hobbs and Spowart 1984). Burning had no

detectable effect on spring diets. The effects of burning

on crude protein in the diet persisted for 2 years in

both communities. The effect on digestible matter was

present only in the mountain shrub habitat the second

year. The increase in the nutritional quality of diets
58 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

was greater than the apparent increase in the quality

of browse and forage, indicating that sheep and deer

foraged selectively for the plants that were more

nutritious.

Fire and the Quantity of Seeds and Mast—The

acorns produced by Western oak woodlands are used

by birds, small mammals, and ungulates. Oaks that

have been severely damaged by fire may produce

“massive” seed crops (Rouse 1986).

Sagebrush and Sagebrush Grasslands

Fire and the Quantity of Forage—Some studies

report no increases in grass and sagebrush productivity due to fire but do report other changes favorable to

ungulates. Burning big sagebrush-bluebunch wheatgrass winter range in Wyoming decreased sagebrush

for the 4 years of study but did not increase wheatgrass. Annual forbs were more abundant on the burned

area only the second year after burning. Nonetheless,

bighorn sheep and possibly mule deer made greater

use of the burned areas than the unburned areas (Peek

and others 1979). Prescribed burning reduced plant

litter that inhibited grazing by elk on a Montana

fescue-wheatgrass winter range. Fire did not significantly change the forb, shrub, and grass standing

crops, however, except that rough fescue, the preferred winter forage, was reduced the first year after

burning (Jourdonnais and Bedunah 1990).

Fire and Availability of Invertebrates—Standreplacing and mixed-severity fire in big sagebrush in

Oregon did not affect populations of darkling beetles

or June beetles (Pyle and Crawford 1996). Fire did not

appreciably alter their food and cover on the ground

surface.

Deserts

Fire and the Quantity of Forage—Fire reduces

most shrubs in the Great Basin Desert for at least a

few years (Humphrey 1974). In the first year after fire,

perennial grasses and forbs have reduced vigor and

annuals are abundant. By the third year, total herbage often reaches a maximum, exceeding production

on unburned sites, and grasses and herbs flower

profusely. The dominant grasses are thickspike wheatgrass, plains reedgrass, and bluebunch wheatgrass;

other grasses, including bluegrass species and Idaho

fescue, do not recover to preburn production until the

second decade after fire.

Fire in the Mojave Desert is likely only after a season

of heavy production by annual plants. The moisture

levels of woody and perennial plants determines the

level of mortality. If conditions are excessively dry,

damage is severe.

In the Sonoran and Chihuahuan Deserts, fire is

uncommon because of the widely spaced, openbranched vegetation. In wet years, fires occur in grasslands and their interface with desert, killing woody

plants, such as velvet mesquite, and expanding the

grassland. Fires that burn off the spines from cacti

(cholla, pricklypear, and barrel cactus) make the plants

available as forage for livestock and rabbits. Fires at

the grassland-woodland ecotone may remove woody

vegetation without increasing ground cover (Bock and

Bock 1990). In desert grasslands, fire is likely to

reduce yield for 1 to 2 years, with productivity recovering to preburn levels by the third year (Jameson

1962; Wright 1980). Where black grama is dominant,

fire effects vary. Productivity may be reduced for 10

years or longer (Wright 1980). Tobosagrass production

increased two to threefold after early spring burns

followed by rain, but burning in a dry spring reduced

yield (Wright 1973).

Subtropical Ecosystems _________

Florida Wetlands

Fire and the Quantity of Forage—In Florida

wetlands, fires increase open aquatic areas and reduce

the encroachment of pine hammocks, thus altering the

balance between terrestrial and aquatic habitat. Burning opens up cattail stands by removing years of

accumulated litter. Fire eliminates litter in sawgrass

stands and reduces plant height for a year or two. To

maintain fruit production for white-tailed deer, Fults

(1991) recommends burning saw-palmetto understories every 3 to 5 years.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 59

L. Jack Lyon

Jane Kapler Smith

Chapter 8:

Management and Research

Implications

Management Implications ________

Only a few places in North America, or the world,

exist where fire has not shaped the vegetation or

influenced the faunal community. In many areas of

North America, managers have successfully prevented

or limited the occurrence of this natural process for

nearly 100 years, and that century of fire exclusion has

probably caused many changes in habitat and wildlife

populations of which we are not even aware. It is likely

that some faunal populations and communities present

in today’s landscapes could not have developed under

pre-1900 fire regimes. Many researchers and managers agree, however, that the success of fire exclusion

cannot continue (Fiedler and others 1998; Fule and

Covington 1995) and, indeed, is already beginning to

fail (Barbouletos and others 1998; Wicklow-Howard

1989; Williams and others 1998). Fire is most likely

to increase in wildlands in the future. This likelihood carries with it two broad implications for the

relationships between fire and fauna.

One: Alternatives in Managing Fire

Managers are increasingly likely to have to choose

among:

• Massive fire suppression (with increasing hazards and increasing costs).

• Uncontrolled, possibly uncontrollable fires.

• A combination of prescribed fires and wildland

fires used to achieve resource objectives.

The implications of these choices for animal communities in North American wildlands are significant.

Most North American fauna communities have developed under pressure from repeated fires of specific

severities and frequencies. Alteration of that pressure

for the past 100 to 500 years has changed the abundance and geographic distribution of many kinds of

habitat and the animals that depend on it.

Even more important than changes in past centuries, however, is the likelihood that fires in the immediate future will deviate substantially from what might

be considered normal or natural in many areas of

North America. While restoration of presettlement

fire regimes may be desirable for habitat protection,

this may be impossible in many areas because of fuel

accumulation, structural change due to fire exclusion,

and climate change (see discussion of this topic in

“Effects of Fire on Flora” in the Rainbow Series).

Even if habitat restoration is successful, animal

populations may be slow to colonize treated areas, so
60 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

perpetuation of existing habitat is a more reliable

management strategy than restoration of degraded

habitat. Managers attempting to restore habitat by

emulating presettlement fire regimes will not only

encounter increased fuel loads and increased continuity of fuels, but also resistance from the public because

of the immediate increased risks to human life, health,

property, and welfare. The altered vegetation may

need to be burned under conditions that would not

normally incur extensive fire spread. For many fauna

species, this practice can produce site and landscape

conditions completely outside the range of those under

which the species evolved. Because spatial and temporal variation are important aspects of presettlement

fire regimes, management plans should address these

features explicitly whenever possible (Lertzman and

others 1998).

Considering the many variables and unknowns that

impinge upon management choices in regard to fire,

careful consideration of the science and monitoring of

treatment results is important. As Rieman and others (1997) comment regarding fire effects on aquatic

fauna, “There is undoubtedly a point where the risk of

fire outweighs the risk of our management, but that

point needs to be discovered through careful evaluation and scientific study not through the opposing

powers of emotional or political rhetoric.”

Two: Integrating Management Objectives

Objectives of prescribed fires and use of wildland

fires for resource benefits must be clearly stated and

integrated with overall land management objectives,

addressing the potential for interaction among disturbances such as grazing, flood, windthrow, predation,

and insect and fungal infestation. In the past 10,000

years, fire has never operated in isolation from other

disturbances, nor has fire usually occurred independent of human influence (Kay 1998; Pyne 1982). During thousands of years prior to settlement of North

America by European Americans, Native Americans

influenced both fire regimes and animal populations.

In fact, populations of large ungulates may have been

limited by Native American predation rather than

food (Kay 1998). As Kay (1995) states, “Setting aside

an area as wilderness or a National Park today, and

then managing it by letting nature take its course will

not preserve some remnant of the past but instead

create conditions that have not existed for the last

10,000 yr.” As managers face ubiquitous needs for

addressing fire in land management, and as they

encounter increasing difficulty in managing habitat in

conditions near those under which faunal species

evolved, we believe it is of paramount importance to

have clear objectives for use of prescribed fire, wildland fire for resource benefits, and fire suppression,

based on understanding of past disturbance patterns

and human influence. It is important to avoid, if

possible, major deviations into ecological conditions

outside the range of variability that occurred in the

millennium prior to 1900.

When fire suppression and use are not integrated

with overall management programs, the potential for

unanticipated problems and failure increases. Management for aspen restoration and bighorn sheep

range improvement provide two examples. If aspen is

treated by fire to regenerate the stand but then repeatedly browsed by wildlife, it often deteriorates more

rapidly than without treatment (Bartos 1998; Basile

1979). The choice of treatment and the size and distribution of treated sites must in this case be integrated

with knowledge of wildlife use patterns and wildlife

management. Prescribed fire can negatively affect

bighorn sheep habitat when range condition is already

poor, when the burn leaves inadequate forage for the

winter, and when other species, especially elk, are

attracted to the burned habitat (Peek and others

1985). Again, fire management needs to be integrated

with wildlife information and management.

Understanding of fire history, potential fire behavior, and differing needs of multiple species must be

integrated in planning for prescribed fire. For example, since many small mammals use tunnels under

forest litter and in or near large pieces of dead wood as

refugia (Ford and others 1999), managers can influence the impact of fire on small mammals by including

moisture levels of these fuels in plans for fire use.

Salvaged logged sites in stand-replacement burns in

the Northern Rocky Mountains provide nesting opportunities for some cavity nesters (northern flicker,

hairy woodpecker, and mountain bluebird). Other bird

species (black-backed woodpecker, northern threetoed woodpecker, and brown creeper) occur almost

exclusively in burned, unlogged patches (Hejl and

McFadzen 1998). If salvage logging is considered after

a wildland fire, the needs of the specific bird community in the area must be considered.

Because funding and other resources for management will always be limited, it is important to use

objectives to shape clear priorities for fire suppression

and fire use. Is it more important to use limited resources on small areas that will benefit small, but

perhaps irreplaceable, populations of animals? Or is it

more important to restore large areas and address the

challenges of landscape-level planning? Only carefully

thought-out objectives can guide such choices well.

Needs for Further Understanding __

Research questions regarding fire effects on fauna

fall into two categories: (1) those regarding faunahabitat relationships and (2) those regarding presettlement fire regimes.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 61

Fauna-Habitat Relationships

Information involving relationships between fire

and animals is needed for all classes of fauna. Most of

the information currently available focuses on vertebrates, particularly mammals and birds. Studies of

landscape and community ecology are virtually limited to birds. Furthermore, most studies are limited to

population descriptors, while measurement of productivity may be essential for understanding fire effects

and predicting effects of management options. Given

the relative lack of information about fire effects on

herpetofauna and insects, studies in those areas may

be especially important (Pickering 1997; Russell and

others 1999). Future research should address microsite

conditions, patchiness within burns, and seasonality

of fire effects for specific ecosystems. Likewise, information about fire effects on aquatic fauna is sparse,

much of it originating from only a few ecosystems (for

example, see Bozek and Young 1994; Mihuc and others

1996; Minshall and others 1989; Rieman and others

1997). More information is needed regarding longterm effects, landscape effects, and effects of postfire

succession on aquatic fauna. (See also discussion of

this topic in “Effects of Fire on Soil and Water” in the

Rainbow Series.)

The need to fill information gaps will increase as

stands and landscapes continue to diverge from presettlement patterns and as managers increasingly use

fire for vegetation management. To improve long-term

management for sustaining ecosystems, information

is needed about the effects of fire on many kinds of

fauna, at different seasons and under different conditions, and over many decades. Information on the

interactions of burning season with life cycles of animal species, especially insects and herpetofauna, is

also important.

Site-Level Research Questions—At the site level,

managers need detailed information on the use of fire

to manage the structure of vegetation, especially in

shrublands and forest understories. Objectives for

this kind of management include maintaining nesting

habitat for birds, ensuring habitat features needed for

reproduction by herpetofauna and insects, providing

cover for small mammals, and enhancing local community diversity.

Also at the site level, managers need better designed, more comprehensive studies of fire impacts on

quantity and quality of forage for wildlife. A truly vast

literature addresses this subject, but much of it is hard

to apply because the investigators did not control for

factors other than burning and did not describe fire

severity or burning conditions in detail. Land managers in many localities currently use limited amounts of

prescribed fire to enhance wildlife habitat, but more

widespread use of fire in habitat management will

require more comprehensive knowledge than is currently available.

Landscape-Level Research Questions—At the

landscape level, we lack almost any knowledge of the

combination of mosaics and patterns best suited to

specific populations, and we have little understanding

of how to maintain the total landscape for regional

biodiversity. While habitat corridors are important for

sustaining some wildlife species (Beier and Noss 1998;

Oliver and others 1998), what are the implications of

fire and succession in corridors and the locations that

provide access to them? Some research of this kind is

under way, but limitations of time and money will

virtually assure that computer models rather than

landscape-level experiments will provide the greatest

progress (Schmoldt and others 1999).

Wildlife researchers often face a dilemma regarding

research priorities: Should we invest time and resources in learning more about faunal habitat, or

should we learn more about the species themselves?

The answer depends on the ecosystem under study.

Schultz and Crone (1998) developed a model for habitat change in the prairie habitat of the Fender’s blue

butterfly, a candidate for listing on the U.S. Endangered Species list. They report that lack of knowledge

about postfire habitat change limited the certainty of

the model’s predictions more than lack of knowledge

about the butterfly itself. In contrast, both Wright

(1996) and Telfer (1993) state that information about

the fauna species investigated (birds in both studies),

especially nesting success, currently limits our ability

to understand the effects of potential management

choices, including those regarding fire.

Presettlement Fire Regimes

Important knowledge gaps remain about the distribution and structure of vegetation in presettlement

times. Without this information, managers cannot

decide what proportion of forest land should be in

various age classes, structural classes, and cover types

to maintain biodiversity. Furthermore, managers need

methods for integrating current agricultural and

infrastructural elements in the landscape with remaining wildlands at large scales, approximating the

original fire-shaped mosaic and structure for an area

as well as possible. With this information, wildlands

can be used to the best advantage to maintain regional

biodiversity, increase numbers of particular wildlife

species, and achieve other environmental goals.

Human Dimension_______________

Finally, researchers and managers need to collaborate in assessing the comparative merits and drawbacks of various kinds of fire for natural resource
62 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

objectives across the landscape. What ecological and

social risks occur with prescribed fires, wildland fires

managed for resource objectives, and fire suppression? How can these risks be reduced? It is impossible

to know all the consequences of intervening in an

ecosystem, whether the intervention is active (prescribed fire, for example), or passive (such as fire

exclusion or landscape fragmentation). Monitoring

and comparison of monitoring results with predictions

are essential. Communication among researchers,

managers, and the public is also essential. Science

cannot be used until it is shared with and understood

by managers, whose job is to apply the results, and a

substantial proportion of the public, who add the

perspective of their values and experience. Policy,

according to Pyne (1982), “has to be based on broad

cultural perceptions and political paradigms, not solely

on ecological or economic investigations; scientific

research is only one component among many that

contribute to it.”
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 63

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 75

Appendices

Appendix A: Common and Scientific Names of Animal Species __________

Taxonomy for birds is from Ehrlich and others (1988); for reptiles and amphibians, Conant and Collins (1991)

and Stebbins (1985); for mammals, Jones and others (1992); for insects, Borror and White (1970).

Common name Scientific name

Agile kangaroo rat Dipodomys agilis

American Kestrel Falco sparverius

American marten Martes americana

American Robin Turdus migratorius

American Wigeon Anas americana

American badger Taxidea taxus

American beaver Castor canadensis

Arachnids (includes spiders

 and ticks) Arachnidae

Bachman’s Sparrow Aimophila aestivalis

Bald Eagle Haliaeetus leucocephalus

Barn Swallow Hirundo rustica

Beetles Coleoptera

Bewick’s Wren Thryomanes bewickii

Bighorn sheep Ovis canadensis

Bison Bison bison

Black bear Ursus americanus

Black Vulture Coragyps atratus

Black-backed Woodpecker Picoides arcticus

Black-chinned Sparrow Spizella atrogularis

Black-footed ferret Mustela nigripes

Black-tailed jackrabbit Lepus californicus

Blue Jay Cyanocitta cristata

Blue-headed Vireo Vireo solitarius

Blue-winged Teal Anas discors

Box turtle Terrapene carolina

Brewer’s Sparrow Spizella breweri

Brown Creeper Certhia americana

Brown-headed Nuthatch Sitta pusilla

Brush mouse Peromyscus boylii

Brush rabbit Sylvilagus bachmani

Bugs Hemiptera

Burrowing Owl Athene cunicularia

California Gnatcatcher Polioptila californica

California Quail Callipepla californica

California mouse Peromyscus californicus

California pocket mouse Chaetodipus californicus

Canyon Towhee Pipilo fuscus

Caribou Rangifer tarandus

Carolina Wren Thryothorus ludovicianus

Cattle Egret Bubulcus ibis

Central Florida crowned snake Tantilla relicta

Chigger Acarina

Chipping Sparrow Spizella passerina

Cicadas, hoppers, whiteflies, aphids,

 scale insects Homoptera
76 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

Clark’s Nutcracker Nucifraga columbiana

Clay-colored Sparrow Spizella pallida

Common Grackle Quiscalus quiscula

Common gray fox Urocyon cinereoargenteus

Common Ground-dove Columbina passerina

Common Raven Corvus corax

Common Yellowthroat Geothlypis trichas

Cooper’s Hawk Accipiter cooperii

Cotton rat Sigmodon spp.

Coyote Canis latrans

Crested Caracara Caracara plancus

Crossbills Loxia spp.

Dall’s sheep Ovis dalli

Dark-eyed Junco Junco hyemalis

Darkling beetles Tenebrionoidea

Deer mouse Peromyscus maniculatus

Desert woodrat Neotoma lepida

Desert tortoise Gopherus agassizii

Diamondback rattlesnake Crotalus admanteus

Downy Woodpecker Picoides pubescens

Dusky-footed woodrat Neotoma fuscipes

Eastern glass lizard Ophisaurus ventralis

Eastern Kingbird Tyrannus tyrannus

Eastern Wood-pewee Contopus virens

Eastern Bluebird Sialia sialis

Eastern Towhee Pipilo erythrophthalmus

Elk Cervus elaphus

European Starling Sturnus vulgaris

Feather Mite Acarina

Fender’s blue butterfly Icaricia icarioides fenderi

Ferruginous Hawk Buteo regalis

Field Sparrow Spizella pusilla

Flammulated Owl Otus flammeolus

Flatwoods salamander Ambystoma singulatum

Florida Scrub-Jay Aphelocoma coerulescens coerulescens

Fowler’s toad Bufo woodhousii

Fox Sparrow Passerella iliaca

Golden eagle Aquila chrysaetos

Golden-crowned Kinglet Regulus satrapa

Gopher tortoise Gopherus polyphemus

Gopher snake Pituophis melanoleucus

Grasshopper Sparrow Ammodramus savannarum

Grasshoppers, katydids, crickets,

 mantids, walkingsticks, and

 cockroaches Orthoptera

Gray wolf Canis lupus

Gray Catbird Dumetella carolinensis

Great Horned Owl Bubo virginianus

Great Crested Flycatcher Myiarchus crinitus

Great Blue Heron Ardea herodias

Greater Prairie-Chicken Tympanuchus cupido pinnatus

Grizzly bear Ursus arctos

Ground squirrel Spermophilus spp.

Appendix A

Common name Scientific name
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 77

Hairy Woodpecker Picoides villosus

Hammond’s Flycatcher Empidonax hammondii

Heath Hen Tympanuchus cupido cupido

Heerman kangaroo rat Dipodomys heermanni

Hermit Thrush Catharus guttatus

Hooded Warbler Wilsonia citrina

Indigo Bunting Passerina cyanea

June beetles Melolonthinae

Kangaroo rat species Dipodomys spp.

Karner blue butterfly Lycaeides melissa samuelis

Kirtland’s Warbler Dendroica kirtlandii

Lark Sparrow Chondestes grammacus

Lark Bunting Calamospiza melanocorys

Lazuli Bunting Passerina amoena

Lewis’ Woodpecker Melanerpes lewis

Loggerhead Shrike Lanius ludovicianus

Lynx Lynx lynx

Meadow vole Microtus pennsylvanicus

Mole skink Eumeces egregius

Moose Alces alces

Mountain lion Felis concolor

Mountain goat Oreamnos americanus

Mountain Bluebird Sialia currucoides

Mourning Dove Zenaida macroura

Mule deer Odocoileus hemionus

Northern Flicker Colaptes auratus

Northern flying squirrel Glaucomys sabrinus

Northern Goshawk Accipiter gentilis

Northern Harrier Circus cyaneus

Northern red-backed vole Clethrionomys rutilus

Northern Shoveler Anas clypeata

Northern Bobwhite Colinus virginianus

Northern Cardinal Cardinalis cardinalis

Nuthatches Sitta spp.

Ord’s kangaroo rat Dipodomys ordii

Ovenbird Seiurus aurocapillus

Peregrine Falcon Falco peregrinus

Pileated Woodpecker Dryocopus pileatus

Pine Warbler Dendroica pinus

Plains pocket gopher Geomys bursarius

Pocket mouse species Perognathus spp.

Pocket gopher species Thomomys, Geomys spp.

Prairie dog Cynomys ludovicianus

Prairie Falcon Falco mexicanus

Pronghorn Antilocapra americana

Rabbit Sylvilagus spp.

Raccoon Procyon lotor

Red squirrel Tamiasciurus hudsonicus

Red fox Vulpes vulpes

Red-bellied Woodpecker Melanerpes carolinus

Red-breasted Nuthatch Sitta canadensis

Red-cockaded Woodpecker Picoides borealis

Red-eyed Vireo Vireo olivaceus

Appendix A

Common name Scientific name
78 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

Red-naped Sapsucker Sphyrapicus nuchalis

Red-shouldered Hawk Buteo lineatus

Red-spotted newt Notophthalmus viridescns

Red-tailed Hawk Buteo jamaicensis

Red-winged Blackbird Agelaius phoeniceus

Ruffed Grouse Bonasa umbellus

Sage Grouse Centrocercus urophasianus

Sage Sparrow Amphispiza belli

Sage Thrasher Oreoscoptes montanus

Sand skink Neoseps reynoldsi

Sapsucker species Sphyrapicus spp.

Savannah Sparrow Passerculus sandwichensis

Scrub-Jay Aphelocoma spp.

Sharp-shinned Hawk Accipiter striatus

Sharp-tailed Grouse Tympanuchus phasianellus

Short-eared Owl Asio flammeus

Shrew species Sorex and Blarina spp.

Six-lined racerunner Cnemidophorus sexlineatus

Snowshoe hare Lepus americanus

Southern harvester ant Pogonomyrmex badius

Spotted Owl Strix occidentalis

Spruce budworm Choristoneura fumiferana

Spruce Grouse Falcipennis canadensis

Summer Tanager Piranga rubra

Swainson’s Thrush Catharus ustulatus

Three-toed Woodpecker Picoides tridactylus

Townsend’s chipmunk Tamias townsendii

Townsend’s ground squirrel Spermophilus townsendii

Tree Swallow Tachycineta bicolor

Turkey Vulture Cathartes aura

Upland Sandpiper Bartramia longicauda

Vaux’s Swift Chaetura vauxi

Vagrant shrew Sorex vagrans

Vole species Microtus, Clethrionomys, and Phenacomys spp.

Western Bluebird Sialia mexicana

Western fence lizard Sceloporus occidentalis

Western harvest mouse Reithrodontomys megalotis

Western Meadowlark Sturnella neglecta

Western rattlesnake Crotalus viridis

Western Screech-Owl Otus kennicottii

Western Tanager Piranga ludoviciana

White-eyed Vireo Vireo griseus

White-headed Woodpecker Picoides albolarvatus

White-tailed deer Odocoileus virginianus

White-tailed Hawk Buteo albicaudatus

White-throated woodrat Neotoma albigula

Wild Turkey Meleagris gallopavo

Wood Thrush Hylocichla mustelina

Woodland salamanders Desmognathus aeneus, Desmognathus ochrophaeus,

 Eurycea wilderae, Plethodon jordani

Woodrat species Neotoma spp.

Wrentit Chamaea fasciata

Yellow-rumped Warbler Dendroica coronata

Appendix A

Common name Scientific name
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 79

Appendix B: Common and Scientific Names of Plant Species____________

Names are from U.S. Department of Agriculture, Natural Resources Conservation Service (1999).

Common name Scientific name

alder Alnus spp.

alligator juniper Juniperus deppeana

American beech Fagus grandifolia

antelope bitterbrush Purshia tridentata

Ashe’s juniper Juniperus ashei

balsam fir Abies balsamea

barrel cactus Ferocactus spp.

big bluestem Andropogon gerardii

big sagebrush Artemisia tridentata

bigleaf maple Acer macrophyllum

bigtooth aspen Populus grandidentata

bitter cherry Prunus emarginata

black grama Bouteloua eriopoda

black spruce Picea mariana

blackgum Nyssa sylvatica

blueberry Vaccinium spp.

bluebunch wheatgrass Pseudoroegneria spicata

bluegrass Poa spp.

bluestem Andropogon spp.

buckbrush Ceanothus cuneatus

bur oak Quercus macrocarpa

California red fir Abies magnifica

cattail Typha spp.

chamise Adenostoma fasciculatum

Chapman oak Quercus chapmanii

cheatgrass Bromus tectorum

chinkapin oak Quercus muehlenbergii

cholla Opuntia fulgida

deerbrush Ceanothus integerrimus

Douglas-fir Pseudotsuga menziesii

eastern redcedar Juniperus virginiana

eastern white pine Pinus strobus

eastern hemlock Tsuga canadensis

fir Abies spp.

Gambel oak Quercus gambelii

Geyer’s sedge Carex geyeri

giant sequoia Sequoiadendron giganteum

grand fir Abies grandis

gray birch Betula populifolia

greenleaf manzanita Arctostaphylos patula

hickory Carya spp.

huckleberry (western species) Vaccinium spp.

Idaho fescue Festuca idahoensis

incense cedar Calocedrus decurrens

Indian ricegrass Achnatherum hymenoides

Indiangrass Sorghastrum nutans

jack pine Pinus banksiana

Jeffrey pine Pinus jeffreyi

juniper Juniperus spp.

loblolly pine Pinus taeda

lodgepole pine Pinus contorta
80 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

longleaf pine Pinus palustris

manzanita Arctostaphylos spp.

mesquite Prosopis spp.

mullein species Verbascum spp.

myrtle oak Quercus myrtifolia

northern red oak Quercus rubra

oak Quercus spp.

oneseed juniper Juniperus monosperma

paper birch Betula papyrifera

pinegrass Calamagrostis rubescens

pinyon pines Pinus cembroides, P. edulis, P. monophylla

pitch pine Pinus rigida

plains reedgrass Calamagrostis montanensis

pond pine Pinus serotina

ponderosa pine Pinus ponderosa

prairie dropseed Sporobolus heterolepis

pricklypear Opuntia spp.

quaking aspen Populus tremuloides

red alder Alnus rubra

red huckleberry Vaccinium parvifolium

red maple Acer rubrum

red pine Pinus resinosa

red spruce Picea rubens

redwood Sequoia sempervirens

rough dropseed Sporobolus clandestinus

rough fescue Festuca altaica (subspecies F. hallii, F. campestris)

runner oak Quercus margarettiae

sagebrush Artemisia spp.

salal Gaultheria shallon

salmonberry Rubus spectabilis

sand live oak Quercus geminata

sand pine Pinus clausa

saw palmetto Serenoa repens

sawgrass Cladium spp.

Scouler’s willow Salix scouleriana

sedge species Carex spp.

shadbush Amelanchier arborea

shortleaf pine Pinus echinata

silverberry Eleagnus commutata

Sitka spruce Picea sitchensis

slash pine Pinus elliottii

southern magnolia Magnolia grandiflora

spruce species Picea spp.

spurge species Euphorbia spp.

subalpine fir Abies lasiocarpa

sugar maple Acer saccharum

sugar pine Pinus lambertiana

sweetgum Liquidambar styraciflua

thickspike wheatgrass Elymus macrourus

thimbleberry Rubus parviflorus

tobosagrass Pleuraphis mutica

tuliptree Liriodendron tulipifera

turkey oak Quercus laevis

Appendix B

Common name Scientific name
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 81

Appendix B

Common name Scientific name

Utah juniper Juniperus osteosperma

velvet mesquite Prosopis velutina

vine maple Acer circinatum

wedgeleaf ceanothus Ceanothus cuneatus

western hemlock Tsuga heterophylla

western juniper Juniperus occidentalis

western larch Larix occidentalis

western snowberry Symphoricarpos occidentalis

western wheatgrass Pascopyrum smithii

white fir Abies concolor

white spruce Picea glauca

whitebark pine Pinus albicaulis

wild lupine Lupinus perennis

willow species Salix spp.

winterfat Krascheninnikovia lanata

yellow birch Betula alleghaniensis
82 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000

Appendix C: Glossary______________________________________________

The definitions here were derived from the following: fuels and fire behavior from Agee (1993), Brown and others

(1982), Helms (1998), National Park Service and others (1998), Ryan and Noste (1985); fire occurrence from Agee

(1993), Johnson (1992), and Romme (1980); plant reproduction from Allaby (1992), Sutton and Tinus (1983); other

terms from Lincoln and others (1998).

abundance: The total number of individuals of a species in an area or community.

climax: A biotic community that is in equilibrium with existing environmental conditions and represents the

terminal stage of an ecological succession.

cohort: A group of individuals of the same age, recruited into a population at the same time; age class.

connectivity: Accessibility of suitable habitat from population centers. All patches of suitable habitat that can

be reached and occupied are considered connected.

crown fire: Fire that burns in the crowns of trees and shrubs, usually ignited by a surface fire. Crown fires are

common in coniferous forests and chaparral shrublands.

density: The number of individuals within a given area.

dominance (dominant): The extent to which a given species predominates in a community because of its size,

abundance, or coverage.

duff: Partially decomposed organic matter lying beneath the litter layer and above the mineral soil. It includes

the fermentation and humus layers of the forest floor (02 soil horizon).

duration of fire: The length of time that combustion occurs at a given point. Relates closely to downward heating

and fire effects below the fuel surface as well as heating of tree boles above the surface.

fire cycle: Used in this volume as equivalent to fire rotation.

fire exclusion: The policy of suppressing all wildland fires in an area.

fire frequency: A general term referring to the recurrence of fire in a given area over time. Sometimes stated

as number of fires per unit time in a designated area. Also used to refer to the probability of an element

burning per unit time.

fire intensity: Used in this volume as equivalent to fireline intensity.

fire regime: General pattern of fire frequency, season, size, and prominent, immediate effects in a vegetation type

or ecosystem.

fire return interval: Number of years between fires at a given location.

fire rotation: The length of time necessary for an area equal in size to the study area to burn.

fire severity: A qualitative measure of the immediate effects of fire on the ecosystem. Relates to the extent of

mortality and survival of plant and animal life both above and below ground and to loss of organic matter.

fireline intensity: The rate of energy release per unit length of the fire front expressed as BTU per foot of fireline

per second or as kilowatts per meter of fireline. This expression is commonly used to describe the power of

wildland fires.

flame length: The length of flames in the propagating fire front measured along the slant of the flame from the

midpoint of its base to its tip. Mathematically related to fireline intensity and the height of scorch in the tree

crown.

fuel: Living and dead vegetation that can be ignited. For descriptions of kinds of fuels and fuel classification, see

“Effects of Fire on Flora” in the Rainbow Series.

fuel continuity: A qualitative description of the distribution of fuel both horizontally and vertically. Continuous

fuels readily support fire spread. The larger the fuel discontinuity, the greater the fire intensity required for

fire spread.

fuel loading: Weight per unit area of fuel often expressed in tons per acre or tonnes per hectare. Dead woody fuel

loadings are commonly described for small material in diameter classes of 0 to 1/4-, 1/4 to 1-, and 1 to 3-inches

and for large material in one class greater than 3 inches.

ground fire: Fire that burns in the organic material below the litter layer, mostly by smoldering combustion.

Fires in duff, peat, dead moss, lichens, and partly decomposed wood are typically ground fires.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 1. 2000 83

herpetile: Amphibian or reptile.

ladder fuels: Shrubs and young trees that provide continuous fine material from the forest floor into the crowns

of dominant trees.

litter: The top layer of the forest floor (01 soil horizon); includes freshly fallen leaves, needles, fine twigs, bark

flakes, fruits, matted dead grass, and a variety of miscellaneous vegetative parts that are little altered by

decomposition. Litter also accumulates beneath rangeland shrubs. Some surface feather moss and lichens

are considered to be litter because their moisture response is similar to that of dead fine fuel.

mast: Fruits of all flowering plants used by wildlife, including fruits with fleshy exteriors (such as berries) and

fruits with dry or hard exteriors (such as nuts and cones).

mean fire return interval: The arithmetic average of all fire intervals in a given area over a given time period.

mesic: Pertaining to conditions of moderate moisture or water supply.

mixed severity fire regime: Regime in which fires either cause selective mortality in dominant vegetation,

depending on different species’ susceptibility to fire, or vary between understory and stand replacement.

prescribed fire: Any fire ignited by management actions to meet specific objectives. Prior to ignition, a written,

approved prescribed fire plan must exist, and National Environmental Protection Act requirements must be

met.

presettlement fire regime: The time from about 1500 to the mid- to late-1800s, a period when Native American

populations had already been heavily impacted by European presence but before extensive settlement by

European Americans in most parts of North America, before extensive conversion of wildlands for

agricultural and other purposes, and before fires were effectively suppressed in many areas.

rhizome: A creeping stem, not a root, growing beneath the surface; consists of a series of nodes with roots

commonly produced from the nodes and producing buds in the leaf axils.

scatter-hoard: Seed cached in scattered shallow holes, a common behavior for kangaroo rats and pocket mice.

secondary cavity nester: Animal that lives in tree cavities but does not excavate them itself.

sere: A succession of plant communities leading to a particular association.

snag: A standing dead tree from which the leaves and some of the branches have fallen.

stand replacement fire regime: Regime in which fires kill or top-kill aboveground parts of the dominant

vegetation, changing the aboveground structure substantially. Approximately 80 percent or more of the

aboveground dominant vegetation is either consumed or dies as a results of fires. Applies to forests,

shrublands, and grasslands.

succession: The gradual, somewhat predictable process of community change and replacement, leading toward

a climax community; the process of continuous colonization and extinction of populations at a particular site.

surface fire: Fire that burns in litter and other live and dead fuels at or near the surface of the ground, mostly

by flaming combustion.

top-kill: Kills aboveground tissues of plant without killing underground parts from which the plant can produce

new stems and leaves.

total heat release: The heat released by combustion during burnout of all fuels in BTU per square foot or

kilocalories per square meter.

underburn: Understory fire.

understory fire regime: Regime in which fires are generally not lethal to the dominant vegetation and do not

substantially change the structure of the dominant vegetation. Approximately 80 percent or more of the

aboveground dominant vegetation survives fires. Applies to forest and woodland vegetation types.

wildland fire: Any nonstructure fire, other than prescribed fire, that occurs in a wildland.

xeric: Having very little moisture; tolerating or adapted to dry conditions.
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